Grasses of Southern Africa

Grasses of southern Africa

nasionale botaniese

INS TITUUT

Private Baq Xin<

1991 -01- 23

Prfvaatsak X 101 KHuTCfiiA 00

NATIONAL BOTANICAL
INSTITUTE

GRASSES OF SOUTHERN AFRICA

Digitized by the Internet Archive
in 2016 with funding from

South African National Biodiversity Institute Libraries

https://archive.org/details/grassesofsoutherOOgegi

Republic of South Africa

Republiek van Suid-Afrika

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA No. 58
MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA No. 58

GRASSES OF SOUTHERN AFRICA

an identification manual with

keys, descriptions, distributions, classification and
automated identification and information retrieval
from computerized data

G.E. Gibbs Russell, L. Watson, M. Koekemoer, L. Smook,
N.P. Barker, H.M. Anderson, M J. Dallwitz

Illustrations / Illustrasies

W. Roux, C. Bartman, C.E. Smith, B. Loutit and others

Editor / Redakteur: O.A. Leistner

Editorial Committee / Redaksiekomitee

Dr O.A. Leistner
Dr B. de Winter
Dr D.J.B. Killick
Dr M.C. Rutherford
Dr C.J. Scheepers
Mrs B.A. Momberg

ISBN 0 620 14846 2

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j DEPT. VAR IAKC30U (N VVATERVOORSIENING |

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1990 -08- 02

PRETORIA I

I BOTANICAL RESEARCH ».-?TITUT£

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National Botanic Gardens / Botanical Research Institute
Nasionale Botaniese Tuine / Navorsinginstituut vir Plantkunde
South Africa - Suid-Afrika

1990

Dedicated to

LUCY KATHLEEN ARMITAGE CHIPPINDALL CROOK

author of

A Guide to the Identification of Grasses in South Africa
1955

tv

FOREWORD

The aim of the amalgamated National Botanic Gardens and Botanical Research Institute is to improve the quality of
life of the peoples of southern Africa through the use of our plants. It is therefore fitting that this first publication of the
new organization should be on such an economically important plant family as the grasses. This manual for grass
identification is the long-awaited successor to L.K.A. Chippindall’s ‘A guide to the identification of grasses in South
Africa' in Meredith’s Grasses and Pastures of South Africa. Published in 1955, ‘Chipp’ has long been out of print and
the price it commands as Africana puts it beyond the means of those who need to use it. More important, our knowledge
of grasses has increased greatly over the past 35 years, as can be seen from the following: there has been a revolution in
basic classification of the grass family; improvements have been made to the taxonomy of a number of large genera (e.g.
Aristida,Digitaria, Ehrharta , Hyparrhenia , Pentaschistis , Stipagrostis)\ in southern Africa extensive field study of grasses
in recent years has resulted in a more complete list of known species, in better records of species distributions and in a
better understanding of the relationships of many species.

Concurrent with this increase in knowledge of grasses, the past fifteen years have seen the application of
computerization procedures to plant taxonomy. This book is the next stage of development for the List of species of
southern African plants edns ! and 2, produced from the PRECIS computer system at the National Herbarium, Pretoria,
and is a first step in extending PRECIS to include descriptive characteristics. The linking of our species data to world
generic data recorded by L. Watson at the Australian National University, Canberra, to provide a basic reference for
regional floristics foreshadows increasing efforts of taxonomists worldwide to co-ordinate their work through
computerization. This co-operation is especially important in a plant family such as the grasses, which is of high economic
and ecological importance and has many genera that extend beyond national and continental boundaries.

The Grasslands Research Centre (Department of Agricultural Development) provided the funds that made this
publication possible. The book is therefore an example of cooperation not only of taxonomists, ecologists and computer
scientists within and outside South Africa, but also demonstrates cooperation between a State Department, a Statutory
Board (NBG/BRI) and various Universities. Without the inspiring leadership of Beth Gibbs Russell this would not have
been possible, and she and her co-workers richly deserve the recognition that this publication should bring.

J.N. Eloff

Director of Research

National Botanic Gardens / Botanical Research Institute

Pretoria, January 1990

TABLE OF CONTENTS

FOREWORD v

PREFACE ix

INTRODUCTION 1

Importance of the grasses 1

Objectives of this book 1

Grasses included in this book 1

Relationships, evolution and ecology 1

Classification and nomenclature 4

The grass plant 5

METHODS AND FORMAT 13

KEYS TO GENERA 17

SYNOPSIS OF CLASSIFICATION 29

GENERA AND SPECIES 31

SPIKELET PHOTOGRAPHS 357

APPENDIX 1: SUMMARIZED CLASSIFICATION OF SOUTHERN AFRICAN GRASSES 381

Pooideae 381

Bambusoideae 382

Arundinoideae 384

Chloridoideae 385

Panicoideae 386

APPENDIX 2: DELTA 389

World grass genera - character list 389

Southern African grass species - character list 405

Parameters for generic keys 407

LITERATURE REFERENCES 409

GLOSSARY 413

INDEX TO SCIENTIFIC NAMES, SYNONYMS AND COMMON NAMES 421

vii

PREFACE

Responsibilities of each author

Major responsibilities were borne by each of the seven
authors as listed below, but there has been much critical
exchange and sharing of jobs at all stages of the project.

H.M. Anderson: Arundinelleae, various genera in
Paniceae; preparation of bromides for illustrations, page
lay-out.

N.P. Barker: Arundinoideae; computerization and
production of distribution maps from PRECIS.

M.J. Dallwitz: programming of DELTA and its associ-
ated programs, CONFOR, KEY, INTKEY and TYPSET.

L. Smook (Fish): Aristideae, Eragrostis , Panicum ;
generic keys, herbarium and Specimen-PRECIS curation.

G.E. Gibbs Russell: Andropogoneae, Ehrharteae,

various genera in other tribes; overall planning, additional
descriptive data for Watson’s world generic database,
generic keys, generic descriptions, introductory text,
glossary, index, typesetting, page lay-out.

M. Koekemoer: Chloridoideae, Pooideae, various genera
in Paniceae; spikelet photographs, production of distri-
bution maps, applications of DELTA, continuity of work
at PRE, page lay-out, cover photograph.

L. Watson: Computerization of world generic data;
subfamily and tribal classification; generic descriptions.

Several specialists contributed treatments of genera in
which they have particular expertise: H.P. Linder,
University of Cape Town, and R.P. Ellis, Grassland
Research Centre, Pentaschistis: P.D.F. Kok, University of
Pretoria, Digitaria: T.M. Sokutu, University of Transkei,
Aira, Arrhenatherum, Avena, Deschampsia, Holcus ,
Koeleria, Lophochloa ; and E.R. Robinson, University of
the Witwatersrand, Cortaderia.

Acknowledgements

We thank first O.A. Leistner, editor of the Memoirs , for
his unfailing support for the project and his willingness to

discuss any aspect on short notice. Wilma Roux has not
only drawn many of the illustrations, but has also kept track
of all the illustrative material and has cheerfully worked
long hours to set up the pages. Others whose help has been
essential for the project include Adela Romanowski, who
took pains to make excellent prints of the spikelet
photographs; R.P. Ellis, who provided copies of
photomicrographs of leaf blade anatomy for about 40
species to add information to the generic database, and who
was always willing to discuss problems; R.J. Pankhurst,
who advised on key generation; J. Erasmus, of the Soils and
Irrigation Research Institute, who helped us to get an earlier
version of DELTA running on the VAX 1 1/750 computer
at his Institute; Mr Steyn and Mr Venema of Unisys and
Mr Swanepoel of the Department of Agricultural
Development who solved datacommunications problems
between the Burroughs mainframe and the plotter; Mr P.S.J.
Hartzenberg of the Department of Agricultural Devel-
opment who gave facilities to make the bromides of the
illustrations; Emsie du Plessis and Beverley Momberg who
helped with proofreading; Esme Bense, Irma Bense,
Annemarie van Rensburg and Gary van Rensburg of
Remata D.T.P. Bureau and Printers, who printed the text
bromides from DELTA-generated computer files; and B. de
Winter, R.P. Ellis, O.A. Leistner and H.P. Linder, who
suggested improvements to the Introduction.

Permission to use previously published illustrations has
been granted by the following: British Crown Copyright,
reproduced with permission of the Controller, Her
(Britannic) Majesty’s Stationery Office, and the Trustees,
Royal Botanic Gardens, Kew, for Coix lacryma-jobi ,
Elymandra grallata , Entolasia imbricata , Lepturus repens ,
Monocymbium ceresiiforme , Odontelytrum abyssinicum ,
Oxytenanthera abyssinica , Schizachyrium sanguineum,
Schoenefeldia transiens, Sporobolus subtilis, Streblochaete
longiaristata , Thelepogon elegans, and Vossia cuspidata:
the Editorial Board of Flora Zambesiaca , for Sartidia
angolensis ; Penguin Books Ltd., reproduced by permission,
from C.E. Hubbard, 1954, Grasses, illustrations copyright
Joan Sampson, for Agropyron repens (p. 76), Ammophila
arenaria (p. 262), Cynosurus echinatus (p. 192),

Deschampsia caespitosa (p. 226) and Parapholis incurva
(p. 317). We especially thank M.A.N. Mueller, of the
Namibia Herbarium, Windhoek (WIND), who generously
lent us the original plates by Blythe Pascoe (Loutit) used
for his own book. Grasses of South West Africa / Namibia.

\

IX

INTRODUCTION

Importance of the grasses

The grasses are the most important plant family on earth,
in numbers of individuals, biomass, area covered, diversity
of habitats and value to man. Over 30 % of the land area
of the earth is covered in natural grassland and savanna
vegetation dominated by grasses (Walter 1979). Grasses
occur in the tropics, in the arctic and antarctic, in swamps
and deserts and forests, on mountaintops and seashores. The
most widespread flowering plant species, Phragmites aus-
tralis, is a grass (Good 1974). The major part of the land
area devoted to crops is occupied by the great cereals,
maize, wheat, and rice, with smaller tracts in marginal
climates devoted to oats, barley, rye and the millets. The
first steps toward civilization itself began with the
cultivation of grasses whose seeds had previously been
gathered wild. Where conditions are unsuitable for culti-
vation, livestock subsists on natural grazing, while in more
mesic areas planted pastures of cultivars bred for high
forage value allow a carrying capacity higher than that of
the natural rangeland.

There are about 770 genera and 9 700 species of grasses
in the world (Watson & Dallwitz 1989) and the Poaceae is
thus the fifth largest plant family in number of species,
ranking behind Asteraceae, Fabaceae, Orchidaceae and
Rubiaceae. In southern Africa, the grasses include 194
genera and 967 species and infraspecific taxa, of which 1 15
are naturalized and 847 are indigenous, with 329 endemics.
In the southern African flora, grasses rank second (to Aster-
aceae) in the number of genera and seventh (after Aizo-
aceae, Asteraceae, Fabaceae, Liliaceae ( s.I .), Iridaceae and
Ericaceae) in the number of species (Gibbs Russell 1985).

Objectives of this book

This book has two objectives. The primary purpose is
to provide a means for identifying southern African grasses
to genus and species, and to give a core of basic information
about each taxon. Identification aids at the genus level
include keys, descriptions, line drawings and spikelet
photographs; species level identification aids include keys,
distribution maps and brief contrasting descriptions. The
text is supported by a computer treatment that provides
more complete data sets that can be used for interactive
identification and information retrieval. A secondary
objective of the project is to serve as a prototype for the
computerization of descriptive data in the PRECIS com-
puter system and to assess its utility in transferring infor-
mation about plants from taxonomic specialists to practical
users (Gibbs Russell & Arnold 1989). This book should
therefore be regarded as the next logical extension of the
treatment of the Poaceae in the List of species of southern
African plants , edn 2 (Gibbs Russell et al. 1985).

Potential users of an identification manual for grasses
range from school pupils and farmers to specialists in grass
taxonomy. We have attempted to aim the book somewhere
in the middle: to those with a basic background in the
biological sciences. However, so that the book may serve
a wider range of people we have included the many
drawings and spikelet photographs as well as an illustrated
glossary of specialized terms. The Introduction summarizes
important aspects of grass structure and relationships and
is intended for those with little access to the extensive
specialist literature on these subjects.

It must be emphasized very strongly that this book
represents a state-of-the-art account of our grasses in 1989;
a definitive treatment will require much more research into
the basic biological behaviour and taxonomic relationships
of our species. Throughout the book we have pointed out
particular problems where further study is needed, and we
hope that these comments will be a stimulus to more
research on the grasses of southern Africa.

Grasses included in this book

All grasses that are indigenous in southern Africa, plus
those that are naturalized and form self-sustaining popula-
tions under local conditions, are covered in the book. It is
probable that other naturalized species may occur which
have not yet been collected.

Grasses that grow in southern Africa only under culti-
vation are not included. Grasses are cultivated for pastures,
lawns, garden ornamentals, erosion control and especially
for crops. Grain crops brought to southern Africa from other
continents include maize (Zea mays L.) from central
America, wheat ( Triticum aestivum L.) from Europe, the
Mediterranean area and western Asia, barley ( Hordeum
vulgar e L.) from north temperate regions, oats ( Avena
sativa L.) from temperate parts of the Old World, rye
(Secale cereale L.) from northern Eurasia and rice (Oryza
sativa L.) from tropical Asia. Of all our grain crops, only
the millets (Pennisetum glaucum (L.) R. Br. and Sorghum
bicolor (L.) Moench, which continues to hybridize with its
wild relatives) were brought into cultivation in Africa. It is
noteworthy that species widely grown for pastures (e.g.
Digitaria eriantha, Lolium temulentum ), lawns (e.g. Cyno-
don dactylon , Pennisetum clandestinum ), ornamentals (e.g.
Cortaderia selloana, Pennisetum villosum) and erosion
control (e.g. Ammophila arenaria, Ehrharta villosa ) are
either indigenous or have become naturalized. In contrast,
the crop species cannot live successfully out of cultivation
(with the exception of Avena sativa , which belongs to a
genus with many weed species).

Relationships, evolution and ecology

Grasses have the herbaceous stems, sheathing leaf bases
and vestiges of the 3-merous flowers common to most
monocotyledons. Although grasses were previously class-
ified with the sedges and rushes, which are similar vegeta-
tively and also have highly modified ‘chaffy’ inflorescences
(Cronquist 1981 ), it is now thought that these resemblances
are superficial only. More recent classifications indicate
that the nearest relatives of the grasses are probably to be
found in the tropical families Flagellariaceae and Joinville-
aceae and the southern hemisphere family Restionaceae
(Takhtajan 1969, Campbell & Kellogg 1987, Linder 1987).

The available fossil record of grasses provides little
direct information about grass evolution, so our knowledge
must be deduced by comparing living forms and is therefore
inevitably speculative. The earliest fossils are floret frag-
ments from the Oligocene of North America and leaf
cuticles from the Oligocene and Eocene of Germany

2

(Thomasson 1987). Indirect evidence of the rise in domin-
ance of the grasses and the formation of grasslands comes
from the change in dentition of the grazing animals. Herbi-
vore fossils of Eocene age first showed development of
high-crowned teeth capable of chewing grass plants, which
are abrasive because of silica bodies in the epidermis
(Stebbins 1981). There is evidence that by the Oligocene
grassland was an established vegetation type, with at least
one genus, Stipa, that is still in existence (Clayton 1981 ).

In their co-evolution with animals, the herbaceous habit
and intercalary meristems have made it possible for grasses
to thrive while being eaten, and they lack many of the
various secondary chemical compounds repellent to herbi-
vores that are found in many other plant families. In
common with other wind-pollinated groups, grass flower
structure is much reduced, and there is often an increase in
male flowers over female-fertile flowers. There has thus
been none of the co-evolution with animal pollinators that
seems to have been so important in other plant groups,
which has there resulted in dramatic floral modifications
and chemical compounds that attract pollinators by shape,
colour, and scent. The pollen grains themselves are uniform
in surface structure throughout the grasses and it is not
possible even to distinguish the subfamilies (Watson & Bell
1975). This is another contrast to many other plant families,
where pollen surface structures are often diagnostic for
genera and even species. However, recent photographic
studies show that the highly modified bracts (glumes,
lemmas and paleas) making up the grass florets and spike-
lets are precisely oriented to channel the air currents and
facilitate the capture of wind-borne pollen by the stigmas
(Niklas 1985a, 1985b).

The grasses apparently began to diversify before conti-
nents became separated by wide oceans. The subfamilies
and tribes are rather uniformly distributed across the
continents in broad climatic bands, but the genera, which
are of more recent origin, tend to be restricted to a single
continent (Clayton 1983). It is interesting to speculate that
the initial split between Laurasia and Gondwanaland may
be reflected in the grasses by the distinction between the
Pooideae, which have diversified greatly in the northern
hemisphere, and other subfamilies, which have their
greatest diversity in the tropics (Bambusoideae, Panic-
oideae, Chloridoideae) and in the Gondwanaland continents
(many Arundinoideae).

How has a plant family whose members exhibit such a
remarkably uniform appearance been able to adapt with
great success to an extraordinarily wide range of climates
and habitats, and to the changes brought about by man?
Evidently, aspects of their characteristic vegetative struc-
ture confer on the grasses an advantage over other plant
types, yet their apparent uniformity masks significant
physiological and cytogenetic variation. The enormous
success of the grasses may be based on the following
factors:

1. Herbaceous stems and leaves.

Most grasses are herbaceous. They flourish during
favourable periods of the year, completing their annual
growth and reproduction while the weather is warm and
wet, and dying back in cold or dry seasons. Perennials
negotiate unfavourable periods as dormant rootstocks or
rhizomes, and annuals survive as seeds. In southern Africa,
a trend can be seen in many genera where perennial species
occur in mesic areas, and annuals occur in the dry west
( Heteropogon contortus / H. metanocarpus, Sehima
galpinii / S. ischaemoides), or where some species behave
as perennials in more mesic areas and as annuals in arid
areas ( Fingerhuthia africana, Centropodia glauca). Even in
relatively mesic areas it has been found that a grass plant
grows actively only during the most favourable days
immediately following rain (Danckwerts 1988).

2. Growing points at the bases of internodes and leaves.

Intercalary meristems’ are a particularly significant

adaptation to grazing and to fire. In most plants, the
growing points are confined to the tips of stems and

branches. In grasses, besides the usual apical meristem at
the shoot tips, additional growing points are located near
the base of each internode and at the base of each leaf. (Pull
off a grass stem and chew it: the soft, juicy parts are at the
bottom of each internode.) In both culm internodes and
leaves, cell division and elongation and tissue maturation
take place acropetally; that is, the youngest cells are located
towards the base and the older towards the tip. Thus, the
culm and the leaves grow from below rather than at the tips,
so that growth is relatively protected from injury or
defoliation by grazing animals or by fire because the
actively growing part of the plant is usually not removed.
The leaf sheaths are an important adjunct to the intercalary
meristem of the culms. The sheaths protect the immature
culm tissues, help support the weight of the shoot above,
shield the apical meristem that will produce the inflores-
cence, and overlapping layers of sheaths near the base of
the plant act as a ‘splint’ over the weaker mgristematic
portions of each node.

3. Plant architecture .

The appearance of the leaves is fairly uniform
throughout the family, except for the bamboos. The
relatively few species with broad flat horizontal leaf blades
occur in forests where the light intensity is low ( Panicum
laticomum, Oplismenus hirtellus). In contrast, the more
numerous species of the open veld nearly all have long
narrow vertical leaves that are supposedly most efficient in
strong light (Eragrostis curvula, Cymbopogon plurinodis).
This leaf configuration makes the best possible use of
sunlight by allowing light to penetrate deep inside the leaf
canopy of the whole plant. Light therefore reaches a
relatively large total area of leaf surface, and grasses are
thus able to produce a large biomass per volume of space
occupied (Newton & Blackman 1970, Lonsdale &
Watkinson 1983).

4. Leaf structure , photosynthetic pathway and climatic
adaptation.

Although most grass leaves look superficially similar,
their anatomy varies considerably and major suites of
anatomical characters are associated with differences in the
location and biochemistry of photosynthetic processes (Fig.
1). In the most widespread form of photosynthesis in higher
plants, primary assimilation of C02 from the atmosphere as
well as photosynthetic reduction of carbon both occur in all
the chlorophyll-bearing cells of the leaf mesophyll. This is
called the C3 pathway, because the molecules of C02 are
initially fixed as three-carbon chains. It occurs universally
in the grass subfamilies Bambusoideae and Pooideae, in
many Panicoideae and in most genera of Arundinoideae
( Merxmuellera , Pentaschistis).

The other principal form of higher plant photosynthesis
takes place by the C4 pathway, in which the C02 is fixed
initially into four-carbon chains. Here, there is a division
of labour in the leaf tissues, with the mesophyll cells (PCA
tissue) restricted to primary carbon assimilation from
atmospheric C02, and subsequent photosynthetic carbon
reduction confined to specialised cells (PCR tissue) which
usually ensheath the vascular bundles. C4 photosynthesis
occurs sporadically in many plant families, including
Chenopodiaceae, Euphorbiaceae, Asteraceae and Cyper-
aceae ( i.e . in both monocots and dicots), as well as in
Poaceae. Among the grasses, C4 photosynthesis is concen-
trated in the subfamilies Chloridoideae and Panicoideae.
The Chloridoideae are almost exclusively C4, the only
known exception being Eragrostis walterii (Ellis 1984).
Among the Panicoideae, the supertribe Andropogonodae
seems to be exclusively C4, while the Panicodeae include
large suites of genera that are exclusively C3 ( Oplismenus ,
Sacciolepis) or C4 ( Brachiaria , Setaria), a few genera in
which both C3 and C4 species occur ( Panicum , Alloterop-
sis), a few truly indeterminate species, and even species
(Alloteropsis semialata , Panicum ecklonii) which include
both C3 and C4 forms. Most of the genera of Arundinoideae
are C3, but some of the largest are exclusively C4, notably
Aristida and Stipagrostis. (A less common variant of photo-

3

GROUP 1

non-Kranz, PS, no specialized chloroplasts.
C3

GROUP 2

Kranz, PS, centrifugal chloroplasts.
C4, PEP ck

GROUP 4

Kranz, MS, centrifugal chloroplasts.
C4, NADP me

GROUP 3

Kranz, PS, centripetal chloroplasts.
C4, NAD me

Fig. 1. Transverse sections of leaf blades in Panicum, showing the different anatomical types associated with the C3 and
C4 photosynthetic pathways (R.P. Ellis 1988).

synthesis, termed crassulacean acid metabolism (CAM),
occurs in a wide range of succulent plants but has not been
found in grasses.)

C4 photosynthesis is experimentally shown to be more
efficient than the C3 version at higher temperatures and
higher light intensities (Bjorkman 1976). These differences
are broadly reflected in world and local geographical
distributions and ecological ranges of grass subfamilies,
genera and species. Thus, of the two exclusively C3 sub-
families, the Pooideae reach maximum diversity in the tem-
perate zone, especially in the northern hemisphere, with
major representation elsewhere only at high altitude and in
moist habitats, and the Bambusoideae, though mostly trop-
ical, are mainly confined to humid forest shade. The two
major C4 subfamilies, Chloridoideae and Panicoideae, on
the other hand, are concentrated in the tropics and subtrop-
ics - the former in drier or saline habitats (see below),
whereas the latter is extensively mesic, with the C3
representatives often being aquatic or shade plants. The
Arundinoideae, with large C3 and C4 genera, are wide-
spread: they are particularly diversified in the temperate
southern hemisphere, but the C4 genera are concentrated in
warm regions (Hartley 1958a, 1958b, 1973, Hartley &
Slater 1960).

A C4 grass leaf blade when seen in transverse section
will usually exhibit a rather characteristic appearance,
known as 'Kranz anatomy’. The specialized PCR tissue
here occurs around each main vascular bundle as a single,
conspicuous sheath of generally starch-rich cells with
abundant chloroplasts; the intervening PCA mesophyll cells
commonly exhibit a degree of radiateness about the indiv-

idual bundles; and an inner sheath of smaller cells (the
‘mestome sheath’) is only sometimes present. C3 leaves, by
contrast, are non-Kranz: the inner, mestome sheath is
always present, the outer bundle sheath cells lack or are
deficient in chloroplasts and starch, and the mesophyll is
usually not noticeably radiate. The Kranz/non-Kranz dis-
tinction is rather imprecise, however, and some grasses
(including the common genera Aristida and Arundinella)
have leaf blade tissue arrangements where its application
is ambiguous or impossible. The only universally applic-
able, unambiguous and reliable method of anatomical
assignment to C3 or C4 relies upon counting the number of
cells separating chlorenchymatous mesophyll cells from the
nearest PCR cell. In the C3 leaf blade the mesophyll always
has some (often many) chlorenchymatous cells separated
from the nearest PCR sheath cell by two or more (often
many more) comparable cells. In a C4 leaf blade, by
contrast, no chlorenchymatous mesophyll cell is separated
by more than one other similar cell from the nearest PCR
cell (synonymous with the nearest bundle sheath cell in all
African grass genera except some Arundinelleae, which
may exhibit conspicuous PCR strands in isolation from the
vascular bundles).

Grasses exhibit three biochemical variants of the C4
pathway, which are less precisely associated with certain
anatomical and ultrastructural features. Those exhibiting
NADP-ME type C4 photosynthesis, called 'malate formers’,
tend to predominate in all regions where C4 grasses occur,
but they reach their maximum abundance in mesic areas.
This photosynthetic type most often occurs among the C4
Panicoideae, of which the supertribe Andropogonodae

4

Fig. 2. Grass subfamily regions in southern Africa. The
symbol in each whole degree square denotes the subfamily
with the largest number of species recorded in Specimen-
PRECIS: A = Arundinoideae (including Ehrharta ); C =
Chloridoideae; F = Pooideae; P = Panicoideae. Panicoids
are most abundant in summer rainfall areas with more than
500 mm of rainfall per year; chloridoids are most abundant
in summer rainfall with less than 500 mm of rainfall per
year; arundinoids are most abundant in areas with more than
40 % of rainfall occurring in winter; indigenous pooids are
most abundant in the high Drakensberg and introduced
pooids share abundance with arundinoids in winter rainfall
areas (Gibbs Russell 1988).

seems to be exclusively NADP-ME. The ‘aspartate
formers’ (NAD-ME and PCK photosynthetic types), on the
other hand, reach their maximum diversity in relatively arid
regiorts. This photosynthetic type is concentrated in the
Chloridoideae, where the NADP-ME type is unknown, but
is also represented in the Paniceae. The ecological associa-
tion breaks down in the C4 Arundinoideae, however, since
all the representatives whose photosynthetic type is so far
known have proved to be NADP-ME, notwithstanding their
importance in dry regions ( Aristida and the Australian
Eriachne).

The idea that differentiation into C4 types is fundament-
ally an expression of climatic adaptation has now been
further undermined, with the discovery that many species
had been mis-typed as PCK by erroneous predictions of
biochemistry from anatomy. Thus the genus Eragrostis,
which had been thought to include both NAD-ME and PCK
species (with the latter species occupying habitats inter-
mediate between those typical of NAD-ME and NADP-ME
forms), now seems to be exclusively NAD-ME. Extending
biochemical typing ( e.g . to cover Stipagrostis) will clarify
the picture, but it is already clear that taxonomic groupings
and certain features of leaf anatomy loosely associated with
C4 types are better indicators of ecological adaptation than
are the C4 types themselves. The important structural
features include: presence (XyMS+) or absence (XyMs-) of
mestome sheath cells between the large metaxylem
elements and the PCR sheath cells of primary vascular
bundles; even-versus-uneven outlines of PCR sheaths;
presence or absence of a suberised lamella in the PCR cell
walls; and location (centripetal or centrifugal/peripheral) of
PCR cell chloroplasts (for detailed information, see
Hattersley 1987 and Prendergast & Hattersley 1987).

Comparisons between occurrence of subfamilies and
major climatic regions have been made both on a worldwide
scale (Hartley 1958a, 1958b, 1973, Hartley & Slater 1960)
and in southern Africa (Vogel et al. 1978, Ellis et al. 1980,
Gibbs Russell 1988). Fig. 2 shows the subfamily dominant
in each whole-degree latitude / longitude square in southern
Africa. In general, the three largest subfamilies each
dominate in the region where their characteristic photosyn-
thetic pathway is most efficient. This fundamental differ-
ence in the veld was first noted by Acocks (1953). The
panicoid region corresponds roughly to Acocks’ ‘red grass’

area, and the chloridoid and C4 arundinoid region
corresponds to his ‘white grass’ areas of southern Africa.

Habitats are not sharply divided between the subfamily
regions, and there is a mixture of subfamilies and photosyn-
thetic pathways over large areas. Chloridoid grasses can
grow in dry microhabitats in mesic areas (Microchloa
caffra, Trichonerua grandiglumis), and pooid grasses in
wet places in the desert (Polypogon monspeliensis). Some
panicoids are adapted to arid areas and are widespread and
abundant there ( Cenchrus ciliaris). In the Fynbos of the
winter rainfall areas in the Cape, the grasses form a smaller
component of the vegetation than in summer rainfall
regions, their place being generally taken by Restionaceae.
The naturally occurring Fynbos grasses include many
endemic arundinoids ( Pentaschistis , Pentameris , Merx-
muellera, etc.) and bambusoids ( Ehrharta ) that rarely
extend to regions outside the Fynbos. The cool wet winters
suit pooids, and there are many naturalized genera,
especially from the Mediterranean area ( Avena , Hainardia )
which has a climate similar to the southwestern Cape
(Gibbs Russell 1988, Linder 1989).

5. Hybridization, polyploidy and asexual ly produced
seeds.

Natural hybridization is common in grasses, and
variability is much increased in populations where hybrids
occur. This high level of genetic variability probably allows
grasses to take advantage of new habitats as they become
available (Ehrendorfer 1980). Hybrids are often nearly
sterile because of chromosomal incompatibilities and
because chromosomes may be present in multiple sets
(polyploidy) or be ‘unmatched’ (aneuploidy). This near-
sterility is advantageous in a variant well adapted to a stable
habitat because eliminating the sexual process stops gene
exchange and thus preserves favourable characteristics.
However, it is essential that sterility must not eliminate pro-
duction and dispersal of seeds. Sterile hybrids commonly
set seed through the process of apomixis, in which the
ovules develop without fertilization into seeds which carry
the same genes as the parent. In this way, a favourable
variant can be perpetuated for many ‘generations’ and the
adaptations for seed dispersal in the species can continue
to operate. Furthermore, the sterility resulting from hybrid-
ization and polyploidy is not absolute. There is always a
low incidence of sexual reproduction that maintains
variability. If the environment changes or if a new habitat
becomes available it is likely that yet another form will be
well adapted to the new situation.

There is good reason to suppose that this ability to
hybridize and to exploit the advantages of hybrid species
complexes with ranges of chromosome numbers and gen-
omes is ancient in the grasses. The phenomenon occurs in
all subfamilies and is common in many genera - as many
as 80 % of grass species are of polyploid origin (De Wet
1987). In southern Africa, small genera are often
represented by a widespread, extremely variable species
( Themeda triandra , Heteropogon contortus). Some larger
genera have a number of well-demarcated species with dis-
tinct, restricted distributions, which exist alongside a wide-
spread species that hybridizes with some of them and blurs
the species boundaries ( Hyparrhenia hirta,Digitaria erian-
tha, Eragrostis curvula, Ehrharta calycina, Pentaschistis
pallida).

Classification and nomenclature

Biological classifications cannot be static: they must
change as new data and new interpretations result in new
opinions about the relationships of organisms, and in
response to the changing needs of users. The family
Poaceae has undergone several stages of reclassification, as
information from several disciplines has been added to that
from the basic morphology. Grasses have most recently
been classified into five major subfamilies: Arundinoideae,

5

No. genera

No. species

Pooideae

40

133

Bambusoideae

10

48

Arundinoideae

22

210

Chloridoideae

50

232

Panicoideae

72

334

Total

194

957

Table 1. Number of genera and species (plus infraspecific
taxa) per subfamily in southern Africa.

Bambusoideae, Chloridoideae, Panicoideae, Pooideae,
(Watson et al. 1985) with a sixth smaller subfamily, Cento-
thecoideae, sometimes segregated from the Bambusoideae
(Clayton & Renvoize 1986). Table 1 shows the number of
genera and species in each subfamily. A complete classifi-
cation of the southern African genera, with descriptions of
the subfamilies, supertribes and tribes appears on p. 381,
and a synopsis of the classification is given on p. 29.

In the last century, spikelet structure was the main basis
for higher classification of grasses (Bentham 1883, Hackel
1896). A few grass treatments still in use (e.g. Hitchock &
Chase 1950, Chippindall 1955) follow this classification,
which recognizes two main subfamilies: panicoids vs. the
rest (i.e. Panicoiceae and Festucoideae). However, even
before 1900 it was evident that a classification based on
spikelet characters alone contained artificial groups,
because spikelets of similar appearance occur in more than
one lineage as a result of parallel evolution. In the 1930s
leaf anatomy, cytology and physiology (Avdulow 1931)
were correlated with spikelet structure, and since the 1950s
a number of new classification systems have been
published, based on a wider range of characters including
spikelet structure, leaf blade anatomy, starch grain struc-
ture, cytology, embryo structure, and photosynthetic phys-
iology (Prat 1960, Stebbins & Crampton 1961, Jacques-
Felix 1962, Watson et al. 1985, Clayton & Renvoize 1986,
Tzvelev 1987). However, systematic knowledge of the
‘cryptic’ characters is far from complete, and they are
unrecorded for many genera (Watson 1987). This lack of
basic data introduces an element of uncertainty into even
the most recent subfamily classifications.

Furthermore, although reasonable agreement has been
reached on classification at subfamily level, at least for a
core group of genera in each subfamily, satisfactory classi-
fication of species into genera remains the greatest
challenge for grass taxonomists. Limits between closely
related genera are not settled, for example between Era-
grostis and Stiburus , Ehrharta and Microlaena , Cenchrus
and Pennisetum. A number of ‘satellite’ genera have
recently been united with a larger genus, for example
Pseudobromus with Festuca, Poagrostis with Pentaschis-
tis.Beckeropsis with Pennisetum , Cymbosetaria with Setar-
ia, Rhynchelytrum with Melinis, as well asHypogynium and
Diectomis with Andropogon, (thus decreasing the number
of genera counted for southern Africa). There are several
major problems of generic delimitation which reflect the
need for further taxonomic studies on a world scale, for
example among the Paniceae involving Panicum , Brachi-
aria, Pseudobrachiaria, Leucophrys, and Urochloa and
among the Danthonieae involving Merxmuellera , Karro-
ochloa, Poagrostis , Pentaschistis , Rytidosperma and Dan-
thonia. Unfortunately, the classification of species into
genera often remains contentious, not always through lack
of sufficient taxonomic research, but because there is plenty
of room for different generic interpretations even when
there is agreement on the essential facts. It may seem that
satellite genera are regularly described, submerged,
resurrected, etc., according to personal preferences and
current fashions.

The classification system is related to nomenclature at
the level of genus. When a genus is reclassified, the names
of its species may change. For example, Rhynchelytrum and
Melinis have recently been classified in the same genus
(Zizka 1 988). The International Code of Botanical Nomen-
clature prescribes that the older name must be used for the
combined genus, so all species formerly classified in
Rhynchelytrum must be transferred to the older name Meli-
nis. Occasionally a genus name must change because an
older name is found in the literature even though it has not
been used for many years, for example, the older name Tri-
bolium had to replace Lasiochloa. The same principles of
nomenclature apply at species level also, so that species
names too change as a result of reclassification (‘lumping’
previously separate species, or ‘splitting’ a species into
two) according to the rule of priority.

The grass plant

Grasses may be tufted and erect, creeping and rhi-
zomatous or stoloniferous, floating, climbing, scrambling
or even arborescent. However, these different forms are all
constructed in a modular fashion from repeating units. The
basic construction unit of a grass plant is called a phytomer,
and consists of an internode with its associated node, leaf,
bud and (sometimes) an adventitious root (Clark & Fisher
1987). All parts of the grass plant, excluding only the tiny
flowers hidden in the spikelets, may be considered to be
constructed of phytomers. Fig. 3 illustrates a grass plant
with its parts labelled.

Roots

Grass roots are fibrous, with little modification, and
usually penetrate less than I meter into the soil (Troughton
1957). Each plant has two root systems. Seminal roots arise
from the germinating embryo, and are very soon replaced
by the nodal root system that arises from the culms. Indiv-
idual nodal roots may last one to several years (Clark &
Fisher 1987). Some grasses have stout prop roots arising at
lower nodes on erect culms above the soil surface ( Zea
mays , Hyparrhenia tambaf, others have decumbent culms
that root at the lower nodes ( Enneapogon desvauxii , Cen-
chrus ciliaris)-, and still others have roots arising from
nodes of the stolons or rhizomes (Pennisetum clandestinum,
Cynodon dactyl on).

The root hairs of grasses are often long and persistent,
in contrast to most other plants in which they are short-lived
(Metcalfe 1960). Grasses from arid areas (Brachiaria
serrata, Stipagrostis ciliata) often form rhizosheaths, pro-
tective and absorptive casings around the roots composed
of root hairs, root cap mucilage, sand grains and micro-
organisms. Nitrogen-fixing bacteria can occur in associa-
tion with these rhizosheaths (,Wullsteine/tf/. 1957). Mycor-
rhizal associations on grass roots have been found in Po-
oideae, Arundinoideae, Chloridoideae and Panicoideae, but
not in Bambusoideae (Clark & Fisher 1987).

Stems

Grass stems fall into three general categories, aerial
culms, underground rhizomes and stolons that lie at the soil
surface. The culms are the most conspicuous part of the
grass plant, bearing the leaves and the inflorescence, and
their height, branching pattern and posture largely deter-
mine its overall appearance. Except in the woody bamboos
and a few other forms (Arundo donax), the culms are mostly
annual even though the plant itself may be perennial. The
culms die back every year and the flowering culms die after
flowering. However, sometimes the tillers (side-shoots)
may behave as biennials and flower the following year.

The culms are jointed and nearly always cylindrical,
with elongated internodes connected by short, harder, disc-
shaped nodes. The internodes are most commonly hollow,
but are solid in many panicoid and chloridoid grasses, or

6

sometimes become hollow with age (Brown et al. 1959).
In some grasses the intemodes remain short until the inflor-
escence is developed, when they lengthen rapidly ( Harpo -
chloa falx), but in others the intemodes elongate early
C Phragmites australis). A few grasses have alternating long
and short intemodes ( Stenotaphrum secundatum). As each
intemode matures, the tissues toward its upper end mature
first, leaving an area of undeveloped tissue still capable of
cell division at the base, the intercalary meristem. The
nodes are the point of origin of the buds and leaves. They
are always solid and have a complex vascular organization.
Nodes are often quite different in external appearance from
intemodes, often being wider or narrower and sometimes
hairy ( Setaria incrassata, Trachypogon spicatus), or with
a conspicuous ring of hairs ( Sorghum halepense, Stipa-
grostis ciliata), or have a darker colour ( Eragrostis obtusa.
Sorghum versicolor).

The location of the main branching system of a species
determines not only its appearance but its degree of pro-
tection from grazing and fire. Rhizomatous grasses branch
below the surface of the soil and are extremely well-
protected from fire. Stoloniferous (sward-forming) grasses
branch at the soil surface and thrive under grazing. Tufted
or tussock grasses, including many widespread veld
grasses, branch just above the soil surface. Culms, rhizomes
and stolons arise from lateral buds in the leaf axils. Inside
the leaf sheath, the bud is enclosed in the prophyll, a scale-
like modified leaf with two keels.

The aerial culms may be unbranched (simple) or
branched. The culm branches, called tillers, may arise intra-
vaginally (with the new shoot remaining inside the leaf
sheath and emerging from the top) or extravaginally (with
the new shoot rupturing the base of the leaf sheath).
Worldwide, intravaginal branching is more common than
extravaginal branching, and in southern Africa extravaginal
branching is rare. Intravaginal branching may give the
tillers extra protection from the periodic bums usual in our
area. Valuable pasture grasses produce much herbage, and
are usually freely tillering or branching above the base
(Schmidtia pappophoroides, Cenchrus ciliaris).

The posture of the culms is usually typical for a species
(Fig. 4), and varies from erect ( Miscanthus capensis , Cym-
bopogon plurinodis) through geniculate and bent at the
nodes ( Digitaria sanguinalis, Eragrostis lehmanniana), or
decumbent with the lower part of the culm on the ground
and the upper part erect ( Digitaria debilis, Brachiaria mar-
lothii) to procumbent and lying flat on the ground ( Uro -
chloa panicoides) or even scrambling on other plants ( Pros -
phytochloa prehensilis, Olyra latifolia).

Rhizomes are underground stems with scale leaves and
roots at the nodes. True roots are easily distinguished
because they have no nodes or scale-leaves. The buds of rhi-
zomes may develop into erect leafy shoots, into stolons or
into secondary rhizomes, and their proliferation may result
in complicated rhizome systems ( Imperata cylindrica.
Sorghum halepense). Stolons are above-ground, horizontal
stems that produce roots, leaves, and flowering shoots at
their nodes ( Monelytrum luederitzianum, some forms of
Digitaria eriantha). Although they are usually easy to
distinguish, rhizomes and stolons may occasionally inter-
grade ( Cynodon dactylon). Many species of the open veld
have short stout rhizomes and knotty culm bases, these
structures together being loosely designated the ‘rootstock’
(Andropogon ravus, Aristida junciformis) (Gould 1968).

Rhizomes, stolons and culm tillers all are important in
vegetative reproduction. Culms and roots produced from
the nodes of rhizomes and stolons soon become indepen-
dent plants if the parent structures are severed. Anyone who
has planted a lawn from ‘runners’ (rhizomes and stolons)
knows how easily new plants can grow from the old stems.
Even in erect species that lack rhizomes or stolons, the
aerial culm tillers can be important in vegetative repro-
duction. A vigorous plant produces new tillers toward the

Fig. 4. Typical culm positions.

7

outside. As the diameter of the tuft becomes larger the
centre dies out and the plant assumes a ring-like appear-
ance. Continued growth outward can result in fragmentation
into separate individuals. In England, plants derived from
an original clone of Festuca ovina , with an estimated age
of 1 000 years had spread over 200 meters (Harberd 1961,
1962).

At the base of the internode in many non-pooid grasses
is a swollen area, the pulvinus, which by differential growth
can change the orientation of the intemode above. The pul-
vinus is sensitive to gravity, and when a culm is blown
down the cells on the lower side, stimulated by hormonal
changes, elongate and cause the pulvinus to bend upward,
thereby re-orienting the culm to its normal position (Clark
& Fisher 1987). Pulvini also occur at the base of inflores-
cence branches and cause the inflorescence to open out
quickly when the stamens and stigmas are mature. Occur-
rence of internodal pulvini is a feature of potential taxon-
omic interest (Dayanandan etal. 1977) which merits further
study.

Leaves

Grass leaves consist of three parts (Fig. 5), the sheath
which envelops the culm, the blade which extends from it,
and the collar and ligule located at the junction of sheath
and blade. Blade and sheath are the main sites of photosyn-
thesis in a grass plant. Leaves are nearly always initiated
alternately on opposite sides of the apical meristem, and are
therefore initially 2-ranked (in contrast to the vegetatively
similar sedge family, Cyperaceae, which commonly has 3-
ranked leaves). The leaves may be basal or cauline. Some
species have only basal leaves (Microchloa caffra , Corta-
deria selloana), others have only culm leaves {Pseudopen-
tamer is brachyphylla, Trichopteryx dregeana) and
probably most have both basal and culm leaves (Cymbo-
pogon excavatus). The uppermost culm leaf below an in-
florescence is often somewhat different in form and is
commonly called the flag leaf. Leaves, like internodes, also
have intercalary meristems and increase in length from
growing points near the base. This is especially advan-
tageous because growth continues unimpeded by grazing at
the leaf tips.

The sheaths overlap when the culm is young, and form
an integral part of its support structure. Usually the sheath
margins are rolled together around the culm and not joined,
but in a few genera the sheath margins are fused and the
sheaths are therefore tubular for much of their lergth
(. Melica , Bromus). The sheaths at the bottom of the culm
are called basal sheaths, and may be variously modified.
They may be persistent ( Sporobolus nebulosus, Ehrharta
dura), sometimes becoming split into fibres (Styppeiochloa
gynoglossa, Festuca costata ), or forming a thick bulbous
base around the culm ( Alloteropsis semialata subsp. semi-
alata). When the basal sheaths are strongly keeled, the base
of the plant has a flat, fanlike appearance {Eustachys pas-
paloides, Heteropogon contortus). Sometimes basal and/or
upper culm leaves have reduced blades or consist of
bladeless sheaths ( Ehrharta ramosa, Stipagrostis gemini-
folia).

The ligule is located on the inner (adaxial) side of the
leaf at the point where the sheath becomes the blade. This
unique structure is without homology in other plant families
(Philipson 1935). The ligule may be either a membrane
(Bromus catharticus, Hyparrhenia hirta ), a membrane
fringed with hairs (Cynodon dactylon ,Digitaria tricholaen-
oides) or a line of hairs ( Eustachys paspaloides, Finger-
huthia africana), with all gradations between these states.
Occasionally the ligule may be absent or present only on
the lower leaves ( Echinochloa ). Variations in ligule form
are very useful taxonomically, for example, any grass leaf
with a ligule consisting of hairs or hair-fringed is almost
certainly non-pooid. The ligule type is generally constant
in a genus, though in the large genus Panicum it may be
membranous, hairy or missing. Pooid grasses usually have
pale, translucent ligules ( Puccinellia , Helictotrichon),
while the ligule in panicoid grasses is usually firm, papery.

Fig. 5. Parts of a generalized grass leaf.

dry, and/or brownish ( Digitaria , Cymbopogon). The
function of the ligule is not clear, but it presumbaly
obstructs the entry of water, insects and bacteria (Tsvelev
1983).

The ‘collar’ is the area on the outer (abaxial) side of the
leaf opposite the ligule. It is often thickened, and is some-
times darker in colour than the rest of the leaf. Infrequently
there is a line of hairs ( Alloteropsis semialata, Karroochloa
curva ) or Hap of tissue on the collar which is called the
contraligule or abaxial ligule. Also there may be an
abscission zone in the collar, the blade being shed while the
sheath remains on the plant ( Arundo donax, Phragmites
australis, Ehrharta rupestris). Auricles are small append-
ages in the collar area. They may arise from the sheath
mouth ( Hordeum murinum), from the sides of the collar
(Pentameris thuarii), or from the base of the blade (Ehr-
harta microlaena).

Leaf blades are generally long and narrow, and this
shape is significant to the productivity of grass plants, as
mentioned above. Species with short broad blades tend to
be annuals or to occur in habitats such as forests or
watersides. The lamina may be constricted near the blade
base ( Sorghastrum stipoides), or even be absent entirely,
the blade consisting of the midrib only (Miscanthus
junceus). The prominence of the veins is variable. Many
grass leaf blades have uniformly developed nerves, but
broad-leaved non-pooid species usually have a strong
midrib (Zea mays. Sorghum halepense). Forest grasses
often have inconspicuous, short transverse veins connecting
the longitudinal veins (Olyra latifolia,Megastachya mucro-
nata). The base of the blade, where it joins the collar, may
be straight ( Cymbopogon plurinodis), rounded ( Perotis
patens ), auriculate ( Cymbopogon excavatus, Dihetero-
pogon amplectens ), sagittate ( Setaria appendiculata), or
pseudopetiolate (Setaria sagittifolia, Thamnocalamus tes-
sellatus). The margins may be thickened and undulate
(Brachiaria serrata, Ehrharta capensis ), or ciliate with stiff
hairs (Sporobolus nitens). The tips may be rounded (Chi oris
pycnothrix, Paspalidium obtusifolium), hooded (Hetero-
pogon contortus), attenuate (Phragmites australis) or
pungent (Phragmites mauritianus , Cladoraphis spinosa).

Many grass leaf blades inroll or infold in response to
water stress, and such reactions presumably restrict water
loss from the stomata of their upper surfaces. Inrolling can
be involute from both margins (Diplachne fusca) or
convolute from one side, with one margin wrapped round
the other (Leersia hexandra). Simple folding along the
midrib is quite common (Themeda triandra, Heteropogon

contortus), and sometimes the blades may be plicate and
folded accordion-fashion ( Setaria megaphylla). However,
in many species with the ‘underside’ (abaxial epidermis)
permanently exposed, this surface has at least as many
stomata as the protected upper surface. Search of the com-
puterized descriptive data associated with this book yielded
a list of only 31 southern African genera with species
lacking (or with very few) abaxial stomata, and twelve of
these genera apparently consist exclusively of such species
(e.g. Ammophila, Merxmuellera, Odontelytrum, Oxy-
rhachis, Pentameris, Sphenopus, Styppeiochloa).
Experiments could be devised to investigate whether these
species are particularly efficient in controlling water loss
through inrolling or infolding. This is an example of the
wide opportunities the computerized data provides for
further pursuit of this and many other aspects of morpho-
logical and anatomical structure/function relationships, and
for generating testable hypotheses.

White or brown scale leaves (cataphylls) occur on the
rhizomes and stolons and reduced bladeless sheaths may
occur at the plant base or on the culms. However, the most
obvious reduced leaves are the several kinds of bracts that
occur in the inflorescence. Flowers with their subtending
bracts form the florets and spikelets that comprise the basic
units of the grass inflorescence. A pair of bracts (glumes)
lies at the base of each spikelet, and another pair of bracts
subtends each flower. The lower of these is the lemma, and
the upper is the palea, which is thought to be homologous
with a prophyll because of its commonly 2-keeled structure
(Clifford 1987). In addition, genera with much-branched in-
florescences, particularly in the tribe Andropogoneae, often
have reduced leaves called spathes and spatheoles below
the inflorescence branches and raceme clusters (Mono-
cymbium ceresiiforme , Themeda triandra).

Leaf blade anatomy and its importance in classificat-
ion and identification

In preparing the printed keys and descriptions for this
book, we have assumed that most users will lack the
equipment and/or the inclination to become involved in
anatomy. However, unlike flowers and fruits, leaf blades
are available on most plants most of the time. Furthermore,
certain materials requiring identification, e.g. fossils and
digestive tract contents, consist predominantly of leaf
fragments. Clearly, there is every incentive to develop
techniques and expertise to use the wealth of anatomical
information available, and computer-aided identification
makes identification of sterile and fragmentary specimens
increasingly possible. Nobody with a serious interest in
practising grass taxonomy (as opposed to merely using
some of its results) can hope to do so effectively without
anatomical understanding, and without access to a suitable
compound microscope.

The conventional distinction between ‘morphology’ and
‘anatomy’ is quite arbitrary, and by extending their data
gathering activities into anatomy, and thence into
ultrastructure, physiology and biochemistry, taxonomists
greatly improve the standard of their classificatory work,
while extending the possibilities for identification into new
dimensions. This principle is true for all plant groups, but
it has been applied more widely and to greater effect in the
grasses than elsewhere. Organized acquisition of compar-
ative data on grass leaf blade anatomy commenced in
earnest in the first third of this century, and gradually
accelerated as the data came to be viewed alongside infor-
mation from other fields and as the taxonomic implications
were understood. The most spectacular result was a
revolution in grass classification, the need for which was
apparent by the 1950s but which has only recently been
comprehensively implemented. The extent to which
anatomical and related physiological considerations are
now part of grass taxonomy is apparent in the group
descriptions provided with modern classifications, and is
reflected in the summarized descriptions in the section on
classification (p. 381).

The automated database from which the generic descrip-
tions in this book are derived carries comparative infor-
mation on about 80 leaf blade anatomical characters, about
half of them requiring a transverse section and half
observable in the abaxial epidermis. Many of the characters
are of great identificatory reliability at generic or higher
group level. Such characters of the transverse section
include the various features indicative of the C3 and C4
photosynthetic pathways, as well as those more loosely
associated with the various C4 types. Important characters
of the epidermis include presence or absence and forms of
microhairs and papillae, arrangements of short cells and
shapes of silica-bodies.

Illustrations of salient anatomical characters are
available in several sources: Watson & Dallwitz (1988),
which is cross-referenced with the character list
accompanying the automated world generic database,
Metcalfe (1960), and Clifford and Watson (1977). The
continuing series of papers by Ellis (quoted in Ellis 1987)
contain superb leaf anatomical illustrations of southern
African grasses.

Scanning electron microscopy is easy to apply in
studying surface features, and may be the only practicable
approach to dealing with some kinds of fossil material.
However, light microscopy generally yields more infor-
mation about the epidermis than does SEM, and most of the
accumulated data, both in the literature and in the database,
against which comparisons can be made, were obtained
from light microscopy.

Leaf blade epidermal preparations and sections should
be taken from the mid-laminar region, avoiding diseased
material, flag leaves, first seedling leaves and others which
seem likely to be ‘atypical’. Dried material can be boiled
for a few mintues in water with a wetting agent (such as a
detergent). Epidermis should be taken from the underside
(abaxial surface) of the blade. It can be prepared by peeling
or by scraping away the tissues from the other side. Sections
can be cut using a razor blade or a sharp hollow-ground
razor, with the leaf blade supported in carrot, pickled elder
pith or expanded polystyrene. Many of the most useful
anatomical features can be satisfactorily observed in
unstained sections and pieces of epidermis, mounted in
water, at magnifications between x 25 and x 400. Indeed,
such features as chloroplast distribution are reliably
interpreted only in this way, and it is recommended that all
preparations from living material should first be examined
unstained. Phloroglucinal plus concentrated hydrochloric
acid provides a simple and rapid temporary stain. It is
particularly useful for an inexperienced observer because
the lignified cells walls are stained bright red and provide
a valuable guide to tissue identification and section
orientation, and the acid renders thick sections more
transparent. Permanent preparations of both epidermis and
sections are conveniently and very effectively stained in
phenolic Bismarck brown, which is particularly useful in
picking out the detailed shapes of silica bodies. Stain for
10-20 minutes, wash in distilled water, dehydrate in the
usual way through a sequence of alcohols ending in
absolute, clear in xylene (avoid inhaling the vapour!), then
mount in Depex or Canada Balsam. Bismarck brown can
be made from the following recipe: 1 g Bismarck brown;
5 g phenol crystals; 100 ml distilled water. Mix and leave
to stand for 1 hour (keeps indefinitely).

Inflorescences

The inflorescence is the part of the plant that bears
flowers, and like the vegetative component it is conven-
iently seen as constructed of phytomers (an internode plus
its associated node, bud and leaf), although in some inflor-
escence parts the leaves and buds are suppressed. The in-
florescence terminates the culm, and it matures from the
apex down (basipetally); that is, the older spikelets will be
found toward the tip of the inflorescence and the younger
spikelets toward its base.

9

In most plant families the basic unit for classifying in-
florescences is the flower, but in grasses the basic unit is
the spikelet. Grass inflorescences vary greatly in general
form, size and shape, and the terminology applied to them
is difficult because the terms are not precisely defined and
because there may be continuous variation between
designated ‘inflorescence types’. Unfortunately, a satis-
factory terminology has yet to be devised and generally
applied. In this treatment the following groupings are used
(Fig. 6). They are reasonably unambiguous, and most in-
florescences are readily referrable to one or other of them.

Spike : a single unbranched central axis with the spikelets
sessile upon it (Lolium temulentum, Oropetium capense).

Spike-like main branches borne on a central axis : the
branches may be narrow spikes, racemes, or panicles.
The main branches may arise digitately or subdigitately
at the apex of the culm {Cynodon dactylon.Digitaria eri-
antha) or they may be spaced along the main axis ( Uro -
chloa mosambicensis, Bothriochloa bladhii , Brachiaria
serrata). The part of the axis, or branch, from which the
spikelets arise is known as the rachis, and the basal part
of the branch below the spikelets is sometimes called the
peduncle.

False spike : with spikelet clusters borne sessile or
subsessile upon the unbranched central axis ( Setaria
sphacelata , Pennisetum sphacelatum). Each spikelet
cluster is actually borne on a branch system with very
short internodes.

Raceme', a single unbranched central axis bearing pedi-
cellate spikelets (Urelytrum agropyroides, Heteropogon
contort us).

Paniculate: the main axis giving rise to branches which
bear the spikelets ( Eragrostis curvula, Panicum
natalense). Panicles are the most common inflorescence
type in the grasses. In some cases the panicle may be
very narrow, with erect and appressed branches, so that
it appears spike-like ( Imperata cylindrica , Fingerhuthia
africana).

A complex of ‘partial inflorescences' and intervening
foliar organs: in a number of genera in the Andropo-
goneae, the basic inflorescence unit terminating each
culm branch is a raceme or cluster of racemes. However,
because the culms are (profusely) branched above, the
whole upper part of the plant becomes a ‘false panicle’
or compound panicle ( Hyparrhenia , Cymbopogon).

The spikelets usually occur singly, but in Andropo-
goneae they are nearly always paired, with one spikelet of
each pair sessile or short-pedicellate and female-fertile, the

other long-pedicellate and often male or sterile. Spikelet
pairs are also found in some Paniceae (e.g. Digitaria) but
both spikelets of the pair are similar and hermaphrodite.
Triads of spikelets occur in some panicoids ( Tristachya ) as
well as in a few pooids (Hordeum), and triads often
terminate the racemes in Andropogoneae ( Chrysopogon
serrulatus).

Genera in different tribes often have inflorescences that
are quite similar in appearance. For example, Cynodon
(Chlorideae), D/g/tana (Panicea e),Dichanthium (Andropo-
goneae) all have digitately arranged, spike-like main
branches. Inflorescence type is therefore a poor character
tor classification, but it is a good character for identification
because it is such a striking feature of the grass plant. This
is why, despite the drawbacks of imperfect terminology and
lack of classificatory significance, inflorescence type
features prominently in the keys to genera in this book.

Spikelets

The spikelet consists of the rachilla, or central axis,
which bears distichous glumes at the base and florets above.
Each floret consists of a pair of bracts, the lemma and palea,
which conceal a single delicate flower. Although grass
spikelets vary greatly in their outward appearance, there is
a remarkable constancy of structure (Fig. 7). Rachillas,
glumes, lemmas and paleas are usually readily identifiable
in spikelets though they exhibit many (sometimes
spectacular) modifications.

Spikelet differences provided the main characters for
classification of grasses into genera, tribes and subfamilies
until about 60 years ago (Avdulov 1931). Even though there
has been considerable reclassification of grasses at the sub-
family level based on anatomy, physiology and
cytogenetics, spikelet differences are still the most
convenient characters to use in identifying genera and
species. In order to make a positive identification it is
usually necessary to examine spikelets with at least 10 x
magnification. In this treatment, the underlying computer-
ized data affords the possibility of making identifications
to genera using vegetative and anatomical characters, but
observation of spikelet characters is still necessary to use
the printed keys to genera and species.

Four spikelet differences are particularly important for
identification:

1. Plane of flattening. A laterally flattened spikelet lies
on its side when tossed on a flat surface, while a
dorsiventrally flattened spikelet lies on its ‘back’ or ‘front’
(abaxial or adaxial surface). Generally, chloridoid ( Era-
grostis superba ) and pooid ( Bromus catharticus) grasses
have laterally flattened spikelets whereas panicoid grasses
( Hyparrhenia hirta, Panicum maximum) have dorsi-
ventrally flattened spikelets. Some genera (Aristida, Stipa-
grostis) have nearly cylindrical spikelets.

spike spikelike

main branches

false spike

raceme

panicle

complex of

‘partial inflorescences’

Fig. 6. Diagrams of grass inflorescence types.

10

Fig. 7. Comparative diagrams of generalized grass spikelets.

2. Number of florets. Some genera have only a single
floret in each spikelet ( Sporobolus , Agrostis). There are
always two florets per spikelet in the great subfamily Panic-
oideae ( Panicum , Andropogon) and there are a larger
number of florets in other subfamilies, up to about 15 in
Megastachya and over 50 in Eragrostis.

3. Disarticulation. The mature spikelet may separate
from the plant below the glumes ( Heteropogon , Poly-
pogon), above the glumes ( Chloris , Bromus) or between
each of the florets ( Helictotrichon ), and in some species of
Eragrostis the lemma disarticulates separately, leaving the
palea behind on the rachis. In genera where the spikelets
are aggregated, the whole cluster sometimes disarticulates,
sometimes together with the associated bristles ( Anthe -
phora, Pennisetum, Cenchrus). The place where the spike-
let disarticulates at maturity is of great importance in seed
dispersal because this determines which accessory struc-
tures (paleas, lemmas, rachilla segments, glumes, pedicels)
accompany the seed when it is shed from the parent plant.

4. Sexuality of the spikelets and their florets is less easy
to observe than the others. In the grass family there seems
to be a general evolutionary trend towards increase in
number of male spikelets and comparative reduction in
number of the female-fertile florets. Male or sterile spike-
lets are generally smaller in size with reduced bracts, while
the female-fertile spikelets tend to be larger and have more
elaborately specialized bracts. These differences appear to
be biologically sound in a wind-pollinated family: there are
more pollen-producing male flowers and a variety of seed-
dispersal mechanisms associated with the female flowers.

A few grasses are dioecious, with separate male and
female plants having spikelets and florets of a single sex
(Cortaderia jubata, Festuca scabra ) and a few others have
spikelets of a single sex borne in different inflorescences
(in Zea mays the tassel has only male spikelets and the cob
has only female-fertile spikelets). The majority of grasses,
however, have spikelets of different sexes within the same
inflorescence ( Olyra latifolia) or florets of different
sexuality within the same spikelet ( Panicum deustum,
Melica racemosa ). In the tribe Andropogoneae ( Hemarth -
ria, Andropogon, Themeda, Hyparrhenia, Heteropogon,
etc.) both conditions exist together: the pedicellate spikelets
are usually male or sterile and the sessile spikelets have two
florets, the lower male or sterile and the upper female-

fertile. Thus in the sessile-and-pedicellate spikelet pair
which comprises a seed dispersal unit, there is only one
female-fertile floret, this being the upper floret of the sessile
spikelet.

The location of the vestigial, sterile or male-only florets
in a spikelet is important in both classification and identi-
fication. They are always at the base of the spikelet (prox-
imal) in the panicoid grasses (Panicum, Andropogon, etc.)
and are usually at the apex of the spikelet (distal) in the
genera of other subfamilies (Enneapogon, Eustachys, Fes-
tuca), with some exceptions ( Ehrharta , Phalaris). Some
genera may have both proximal and distal sterile or male
florets (Phragmites, Entoplocamia). The reduced florets
may be variously modified for seed dispersal, with awns
(Holcus, Ehrharta) or hairs (Melica).

Spikelets have three kinds of bracts: glumes, lemmas
and paleas. Glumes and lemmas have been likened to
modified leaf sheaths. Both, but more commonly the
lemmas, may bear awns, which can be envisaged as
modified leaf blades. There are usually a pair of glumes at
the base of the spikelet, and they may be distinguished from
lemmas because they are empty, that is they enclose no
palea or flower (Clifford 1987). Sometimes one (Folium
multiflorum, Eriochloa stapfiana) or both (Oryza longi-
staminata) glumes may be missing. The size and thickness
of the glumes relative to the lemmas are important charac-
ters. The glumes may be longer than the rest of the spikelet
(Avena, Hemarthria, Merxmuellera disticha) or much
shorter even than the adjacent lemmas (Eragrostis, Cyno-
don). The glumes may be firmer than the lemmas ( Themeda ,
Heteropogon) or the lemmas may be firmer than the glumes
(Panicum, Digitaria).

Lemmas are always present. They are more diverse
throughout the family than are the glumes, but are generally
very similar within a genus. The shape, texture, and number
of veins of the lemmas, and the presence, type, number and
location of their awns vary greatly and are therefore taxon-
omically important. Very hard lemmas that persistently
clasp the mature fruit are common in the Paniceae. These
hard lemmas often have an area of weakness on the back,
the germination flap, which opens when the root of the
germinating embryo emerges from the enclosed seed
(Brachiaria, Digitaria, Loudetia).

The palea may be envisaged as a modified prophyll and
lies with its back against the rachilla (Clifford 1987). It is
almost always smaller than the lemma and its margins are

1 1

usually hidden inside the lemma except when the floret
opens to expose the mature stamens and stigmas. In contrast
to the glumes and lemmas, which commonly have an odd
number of nerves and are 1 -keeled, the palea is usually 2-
nerved and 2-keeled. The lemma and palea together enclose
the much-reduced flower. In many cases they continue to
enclose the mature fruit, and are often modified to aid its
dispersal. In genera where the glumes are relatively large
and thick ( Rottboellia , Cymbopogon) the lemma may be
much reduced and the palea vestigial or absent. Conversely,
where the glumes are absent the palea is exposed and
thickened ( Leersia hexandra , Oryza longistaminata),
although the interpretation of organs in oryzoid spikelets is
debatable).

Flowers

The grass flower is composed of lodicules, stamens and
a pistil (Fig. 8). The lodicules are small organs that lie
between the lemma and the stamens. They swell when the
flower is mature and force apart the lemma and palea,
allowing the anthers and stigmas to emerge. After anthesis
the lodicules lose turgidity and the lemma and palea close
again around the developing fruit. There are commonly two
lodicules, but some genera may have one (Melica) or three
( Olyra , Thamnocalamus). The derivation of lodicules is
controversial. They have in the past been considered a
modified perianth but the evidence is not conclusive, and
a recent interpretation considers them to be organs peculiar
to the grasses (Clifford 1-987).

Most grasses have three stamens, but there may be one
(sometimes in Imperata cylindrica ), two (Diandrochloa
namaquensis), four (Microlaena stipoides ) or six ( Oryza
longistaminata , Ehrharta erecta). The bamboo Ochlandra
of India, Ceylon and Madagascar may have as many as 120
stamens. Stamen number tends to be reduced in
cleistogamous spikelets ( Bothriochloa insculpta). Anther
length varies from about 0.1 to 14 mm and is often a
convenient character to separate species. Anthers are
usually small in the chloridoid grasses, and anther size
tends to be reduced in cleistogamous florets. Pollen surface
morphology is remarkably constant throughout the family,
and the genera or even the subfamilies cannot be
distinguished on pollen surface characteristics (Watson &
Bell 1975). However, subfamilies and tribes are
distinguishable in terms of pollen antigens and allergens -
an example of how taxonomy can exchange information
with other disciplines to mutual advantage (Watson & Knox
1976).

The pistil terminates the flower, and has a single locule
containing one ovule. The ovary is generally barrel-shaped
or fusiform and is usually glabrous. However, there is
sometimes a tuft of hairs at its apex ( Festuca , Bromus, and
Pentameris). There are usually two styles, which are
normally separate but may be fused at the base ( Elymandra ,
Entoplocamia). In a few cases the stigmas too are fused

(. Zea mays has a single fused stigma up to 75 mm long, the
longest known for flowering plants (Clifford 1987)). The
stigmatic hairs vary in colour from white to purple-black,
and the colour may change as the stigma ages.

Pollination

All grasses are wind pollinated, except for a few forest-
floor genera that do not occur in southern Africa. Grass
pollen tends to retain its viability only over short distances.
However, it can travel enormous distances; as hayfever
sufferers are uncomfortably aware, grass pollen may travel
many kilometers from stands of heavy pollen-producing
species, retaining its allergenic properties (Gregory 1973,
Knox 1979). The daily period of flowering and pollination
is fixed within narrow time limits for each species, which
presumably increases the chances of pollination. Closely
related species may be reproductively isolated by flowering
at different times of the day (Tsvelev 1983). In short-
distance wind pollinated species outbreeding may be main-
tained by complex incompatibility mechanisms in the
stigma and style (Heslop-Harrison & Heslop-Harrison
1987). Few studies of flowering biology, however, have
been carried out for even the most economically important
southern African species.

In cleistogamous spikelets the lemma and palea do not
open and self-fertilization occurs within the closed floret.
Cleistogamy is not often reported for southern African
grasses. Pits on the lower glumes in Bothriochloa may
restrain stamen emergence (Heslop-Harrison 1961), and
Enneapogon desvauxii has cleistogamous spikelets in the
leaf sheaths as well as ordinary (chasmogamous) inflores-
cences. In Pennisetum clandestinum the spikelets are all
hidden in the leaf sheaths, but they are not cleistogamous
because the stamens and stigmas are exposed.

Fruits, seeds and embryos

The grass grain, or caryopsis, consists of one seed
closely surrounded by the pericarp, the thin adherent outer
layer of the fruit. The caryopsis is the characteristic fruit
type of the grasses, and is unique to them. Throughout the
family there are a number of variations in the pericarp,
which in rare cases may be berry-like, achene-like or nut-
like (Sendulsky et al. 1987). These variations may be
readily interpreted as derived from a caryopsis in which an
inner tissue layer collapses at a relatively late stage of
development, resulting in a free or readily removable
pericarp. For example, especially in the tribe Chlorideae,
the pericarp is soft and separable from the seed (e.g.
Eleusine and Sporobolus, in which the pericarp of the wet
utricle splits and extrudes the seed). On the adaxial side of
the fruit is the hilum, a round ( Eragrostis , Panicum ) or
elongated ( Ehrharta , Stipagrostis) scar where the seed is
attached to the pericarp. Inside the seed, food for the
developing embryo is stored in the endosperm and is com-

Fig. 8. Parts of a spikelet that has a single floret, and its flower ( Cynodon dactylon).

12

posed of starch, oils and proteins. The embryo is small
relative to the volume of endosperm, and lies on the
opposite side to the hilum. Differences in embryo size and
structure tend to characterize subfamilies (see pp.
381-388).

Seed dispersal

Plants are generally sessile, rooted organisms, but
relocation occurs twice in their life cycle, at pollination and
at seed dispersal. In contrast to the apparently unspecialized
wind pollination in grasses, transport of the seed to a
favourable place for germination is effected by a remark-
able range of strategies. However, in most grasses the seeds
and fruits themselves are not modified for dispersal;
instead, it is the inflorescence bracts and branches that show
an array of adaptations involving transport by wind and
water, by other organisms, (in fur, skin, feathers, clothing
and digestive tracts), and even by self-propulsion. Parallel
evolution of the many different kinds of dispersal
mechanisms in various subfamilies and tribes has been a
conspicuous feature in the evolution of the large number of
grass genera and species (Davidse 1987). A few genera
show no apparent adaptation for seed dispersal, and the
caryopsis may be dispersed unaccompanied by accessory
structures ( Agrostis , Eragrostis).

The glumes or lemmas occasionally have long hairs
which sail the fruit long distances ( Stipagrostis , Imperata
cylindrica , Phragmites australis). In other adaptations to
wind dispersal, the entire inflorescence breaks off and rolls
about as a tumbleweed ( Trichonerua grandiglumis, Pani-
cum volutans), or the inflorescence falls together with the
flag leaf, which acts as a sail (Urochlaena pusilla).

Several aquatic genera have corky inflorescence axes,
which break up into short sections each bearing a sessile
and a pedicellate spikelet which float to a new location
( Hemarthria altissima , Rottboellia cochinchinensis).

There are many adaptations to dispersal by animals. The
bare seed may be extruded from the viscous pericarp and
positioned at the spikelet tip where it can adhere to a
passing animal ( Sporobolus ). Other mechanisms to disperse
fruits by clinging include hooks on the glumes or lemmas
(' Tragus racemosus, Pseudechinolaena polystachya );
tangles of large scabrous awns from several spikelets
( Heteropogon contortus)\ clusters of spikelets with sca-
brous subtending bristles ( Setaria verticillata, Cenchrus

brownii)', and calluses with clinging retrorse hairs that
penetrate skin with a pungent tip ( Aristida stipitata, Hetero-
pogon contortus). Note that structures collectively termed
calluses represent modifications of different spikelet parts,
depending on where the spikelet disarticulates: the base of
the lemma (Vulpia), the base of the lemma plus part of the
rachilla (Helictotrichon), the base of the spikelet ( Dihetero -
pogon) or the base of the spikelet plus part of the pedicel
( Schismus barbatus , Polypogon).

Callus hairs and bent and twisted awns act together in
self-propulsion. The callus hairs allow the cylindrical floret
or spikelet to move in only one direction, and the twisting
of the awn as a result of hygroscopic changes drills the
floret or spikelet into the soil ( Pentaschistis , Heteropogon).
Fruits dispersed by this mechanism can thus benefit from
both animal transport and self-propulsion.

In many Paniceae, the fruit is closely surrounded by
glumes, a lower lemma and palea and a hard upper lemma
and palea. These protective layers possibly ensure its safe
passage through the digestive tracts of animals. Herbivory
in this case is part of the dispersal mechanism, and many
species with spikelets of this kind have soft, palatable
herbage (e.g. in Panicum, Brachiaria , Paspalum).
Elaiosomes, oil-containing appendages that mature at the
same time as the caryopsis, are another adaptation for
dispersal which attract insects with the offer of food. Ants
carry the seed underground along with the accessory struc-
ture bearing the elaiosome. In Rottboellia cochinchinensis
the elaiosome is on the ‘peg’ at the base intemode which
accompanies the dehiscent spikelet; in Eriochloa meyeriana
it is a beadlike structure at the base of the spikelet formed
from a reduced glume and the adjacent intemode; Ehrharta
calycina has an ear-like appendage at the base of the sterile
lemmas that may be an elaiosome.

Finally, one of the most remarkable factors contributing
to widespread dispersal of grasses is a consequence of the
copious endosperm of the seed. The nutritious grains
attracted the attention of hunter-gatherer people, and led to
the cultivation and improvement of cereal crops, which has
ultimately resulted in today’s mechanized agricultural
industry based on highly selected hybrid cultivars.
Monocultures of cereal crops now extend over vast areas
of the earth’s surface. These few species have reached
continents far from their places of origin, and have replaced
naturally occurring plants to an extent that represents an
alarming loss of genetic and ecological diversity.

13

METHODS AND FORMAT

What is unusual about this book?

This volume is unique in several ways. First, it is at this
time the only identification manual for a major plant family
occurring in a large area to be produced from computerized
data. The data is consistently recorded and fully compara-
tive, and automation has been applied at all levels:
gathering and recording data, preparing keys and
descriptions, plotting distribution maps, and typesetting.
Second, the data backing up the book and the programs for
manipulations are made freely available on MS-DOS
computer diskettes. The generic descriptive data (Watson
& Dallwitz 1988, 1989) includes copious information on
grass anatomy, biochemistry, physiology, ecology,
host/parasite associations, economic aspects, taxonomic
relationships, nomenclature, phytogeography, source
references, etc. Third, because of this wealth of data at
generic level, sterile and fragmentary specimens can often
be identified to genus using the program INTKEY.
Identification of poor material is impossible in conventional
identification manuals which lack this new dimension of
access to the underlying automated data. Fourth, any part
of the treatment can be easily expanded and updated as
more information is recorded and as taxonomic concepts
change. Fifth, subsets can be generated from the computer-
ized data to cover smaller areas or specific groups of taxa.
These could include additional information present in the
generic and species databases but not published in this
volume.

The resources to produce this co-ordinated treatment,
linking a conventional identification manual with support-
ing computer programs and full data sets, were fully
available at the National Herbarium, Pretoria (PRE). There
has been a long history of grass research at the Botanical
Research Institute, which has accumulated over 70 000
grass specimens from southern Africa, and holds important
collections of illustrations and photographs, rare books
essential to nomenclatural study, as well as slides,
photographs and voucher specimens for comprehensive
anatomical and cytogenetic studies of all southern African
grass species. Besides this specialization in grasses, PRE
is the most advanced herbarium in the world for computer-
ized coverage of its specimens and of the flora of a major
subcontinental region. The PRECIS system holds data for
over 650 000 specimens and 24 000 southern African plant
taxa (Gibbs Russell & Arnold 1989). For this project we
were able to link the southern African grass data at species
level in PRECIS with the automated descriptions of grass
genera developed at the Taxonomy Laboratory of the
Research School of Biological Sciences at the Australian
National University (Watson et al. 1988, Watson &
Dallwitz 1989). Thus all descriptive information has been
handled through the DELTA programs produced at the
Division of Entomology, CSIRO, Canberra (Dallwitz &
Paine 1986). Coupling the data for southern African species
with Watson’s worldwide generic data has resulted in a
package comprising not only traditional-style keys,
descriptions and illustrations, but also a flexible system for
interactive identification and information retrieval.

We believe that the day is past when solitary taxonomic
specialists, working in isolation, produce soon-to-be-
outdated monographs and flora contributions. This book
and its underlying computerized data clearly demonstrate
that a group of scientists with different kinds of expertise
working in widespread localities can co-ordinate their
efforts by contributing to a single database. The result is a
comprehensive taxonomic treatment with greater flexibility

and much wider ranges of applications than would have
been possible using traditional methods.

The DELTA computer system

The DELTA computer system that underlies the book is
a generalized system for handling all the different kinds of
descriptive data used by taxonomists, without information
loss, in an easy-to-use format designed to minimize encod-
ing errors (Dallwitz 1980, Dallwitz & Paine 1986, Watson
& Milne 1972). DELTA was adopted as the standard format
for taxonomic descriptions at the 1988 meeting of the
Taxonomic Databases Working Group for Plant Sciences.

An associated program, CONFOR (Dallwitz & Paine
1986), translates the coded descriptions into natural
language; produces summarized data for specified sets of
taxa, giving for multistate characters the numbers of taxa
exhibiting each character state and for numeric characters
giving the means, ranges and names of taxa exhibiting the
extremes of ranges; and carries out various data mainten-
ance operations, for example changing the sequences of
characters and character states while keeping all the files
consistent with one another. CONFOR can also translate
data coded in DELTA format into formats required by
various other taxonomic programs, including KEY
(Dallwitz 1974, Dallwitz & Paine 1986) and GENKEY
(Payne 1975) for making printed keys; DIST (Dallwitz &
Paine, unpublished) for generating distance matrices; and
PAUP (Swofford 1984) for phylogenetic analysis. DELTA
format is used directly by programs in the PANKEY
package for a number of taxonomic applications (Pankhurst
1986, Pankhurst & Aitchison 1975), and also by the
program TYPSET for automated typesetting from DELTA
data (Dallwitz & Zurcher 1988).

The program INTKEY greatly extends the range of
applications of the DELTA format (Watson et al. 1989).
Besides allowing interactive identification, INTKEY also
provides a flexible system for information retrieval. For
example, it can be used for generating group descriptions
for a specified set of taxa; for determining diagnostic
characters for a taxon or group of taxa; and for finding the
similarities or differences between taxa. INTKEY output
can be read to files as well as to the computer screen, and
information can be changed to the format required for other
programs.

Availability of computerized data and programs

The interactive data set, comprising the generic
descriptions and the species descriptions in INTKEY
format together with the program INTKEY, is available free
of charge from the Data Officer, National Botanic Gardens
/ Botanical Research Institute, Private Bag X101, Pretoria
0001, South Africa, or from L. Watson, Research School
of Biological Sciences, Australian National University,
GPO Box 475, Canberra A.C.T, Australia 2600. The set will
be supplied on 360K 5 1/4-inch floppy disks or 740K 3 1/2
inch stiff disks suitable for MS-DOS microcomputers.
INTKEY requires at least 512K of memory (RAM). The
complete world generic data occupies 71 OK of disk space
and the southern African species data occupies 870K, so a
hard disk or large-capacity floppy is required for data
manipulation. Special subsets could be provided to users
with severe space limitations in their microcomputers.

14

Generic keys

The keys to genera were prepared from the database of
world grass genera maintained in the DELTA system
(Watson 1987, Watson et al. 1989), using the key-
generating program KEY (Dallwitz & Paine 1986). They
continue a series of keys produced from the world database
for several countries: Australia (Clifford & Watson 1977,
Watson & Dallwitz 1980, 1985); Canada (Watson, Aiken
et al. 1985); and Greece (Watson. Damanakis & Dallwitz
1988).

About 100 characters for vegetative and spikelet
morphology were used in the key. These were selected for
ease of observation and usefulness in distinguishing
southern African grasses, from the ca. 480 characters
available in the world grass database (p. 389). KEY was run
a number of times, with adjustments to character
weightings, additional preset characters at desired points
and changes in numeric ranges to improve each run over
the previous one. The program parameters set to produce
each part of the generic key appear on p. 407. The final key
was exhaustively checked against southern African
specimens and minor changes were incorporated by hand.

A number of genera appear more than once in the key,
partly as a result of selecting ‘easy’ characters, especially
for the early choices. The key is therefore longer on the
printed page, but is actually shorter to run: the longest track
through the key to 194 genera reaches an identification in
17 steps, and most tracks are considerably shorter.
‘Backtracking’ after an uncertain outcome is possible
because at each couplet the previous couplet number is
given in parentheses.

We strongly recommend computer-aided identification
using the program INTKEY. The advantages of this method
include use of distribution data to limit the number of
genera to be considered, and access to descriptors for any
plant part in any sequence. It is often possible using
INTKEY to identify sterile or fragmentary material to
genus, which cannot be done through the printed key. (Note
that it is perfectly feasible to use the computerized data to
generate printed keys to genera using vegetative and
anatomical characters.)

The generic keys are not strictly dichotomous. The
number of alternative choices in any set reflects the number
of states recorded for each character. (See the generic
character list on p. 389.) In a few cases the generic key runs
directly to a species, where an anomalous species is quite
distinct from the others in its genus (e.g. Pentaschistis
pusilla , Pennisetum unisetum).

Generic descriptions

The genera appear in alphabetical order so that they can
be easily located in the book. A ‘taxonomic’ sequence was
not used because of the uncertain higher classification of
some genera, and the fact that better generic classifications
will certainly be forthcoming: an attempted ‘taxonomic’
sequence for the genera would soon be out of date. The
most up-to-date classification will be available in the most
recent version of the DELTA database. The classification
and supposed relationships of the genera accepted in 1989
are given in the section on classification of southern African
genera (p. 381), and a synopsis of the subfamilies, tribes
and genera with similar genera listed together, appears just
after the generic key (p. 29).

The descriptions were prepared from the database using
the program CONFOR (Dallwitz & Paine 1986). A core of
characters was included for all genera, as well as additional
characters important in each subfamily. The minimum
diagnostic characters necessary to distinguish each genus
from others in its subfamily are printed in italics, and were
determined by an INTKEY search. These short
descriptions, suitable for a manual such as this, present only
a small fraction of the information available in the database
for each genus.

At the beginning of this project, before generic
descriptions and keys were attempted, the database was
scanned to find southern African genera with missing data.
Much additional morphological information from
herbarium specimens was added for about 70 genera, and
leaf blade anatomical data was provided from copies of
photomicrographs supplied by R.P. Ellis for about 40
genera.

The short list of references for each genus is held in
Taxon-PRECIS, and includes recent revisions followed and
the sources of synonymy for the species.

Illustrations

At least one illustration is included for each genus and
more for large genera, except for four naturalized genera
which are not illustrated ( Bambusa , Microlaena,
Paratheria, Periballia). The drawings were prepared by a
number of artists over the years. The majority (about 140)
have been recently prepared under the supervision of grass
specialists, mostly at PRE and WIND, and some (about 70)
were first published in Chippindall (1955). The 32
‘borrowed’ illustrations are acknowledged in the Preface.
For three naturalized genera, photocopies of herbarium
specimens replace a drawing ( Corynephorus , Craspedo-
rhachis, Lamarckia).

Spikelet photographs

There is at least one spikelet photograph for each genus,
and all photographs were prepared especially for this book.
They were taken with a Wild M400 stereomicroscope
connected to a Wild MP545 photo-automat, using the fixed
time exposure mode and Ilford FP4 film. Spikelets from
herbarium specimens or from fresh plants were mounted on
the tip of a beading needle and positioned on a fine-grained
grey sandpaper. This provided an out-of-focus, uniform,
non-reflecting background. All photographs are grouped
together for reasons of economy. It would have been more
desirable to place each spikelet photograph with the rest of
its generic treatment.

Species keys

The species keys are the only descriptive part of the
book not prepared through the DELTA system. They were
written by hand because the species data available at this
early level of approximation of the species database in
Taxon-PRECIS is presently insufficient for key generation
(see below). All the species keys are strictly dichotomous,
and the previous couplet number is given in parentheses at
each couplet to allow ‘backtracking’.

Species descriptions

The species appear in alphabetical order but cross-
references to related or similar species are provided. The
species treatments were produced from computerized data
held in DELTA format, but the species database is
developed separately from Watson’s world generic data, as
part of Taxon-PRECIS. The DELTA treatment began by
taking over the scientific names, synonyms and references
from Taxon-PRECIS Approximation 2 (Gibbs Russell &
Arnold 1989). Continuing the PRECIS format, the number
after each synonym indicates a source for the synonymy in
the references listed for each genus. As mentioned
previously, the grass species database is a prototype for
adding descriptive data to the Taxon component of
PRECIS, to assess the method before extending it to other
plant families. The species character list (p. 405) is based
on the ‘common knowledge’ characters included as ‘type
one data’ in the ILDIS legume database (ILDIS

15

Coordinating Centre 1986), with the addition of diagnostic
characters and voucher specimens as recommended by the
1987 meeting of the Herbarium Curators Working Group.

The species descriptive data was obtained from original
observations of specimens at PRE (and other herbaria as
necessary) and from literature. It represents the minimum
information required to distinguish each species from others
in its genus. (This is the reason species treatments in large
genera are considerably longer than those in genera with
few species). Data for biomes, distribution, and flowering
time was extracted from Specimen-PRECIS (Gibbs Russell
& Arnold 1989) and verified as each genus was studied. In
most genera ‘flowering’ time represents the spread of
months in which spikelets are present on a range of
herbarium specimens, but in Pentaschistis the time of
anthesis was determined by field' study. The specialist
responsible for the species treatment in each genus is named
at the end of the generic description.

Distribution maps

The maps for each species were plotted from specimen
records held in Specimen-PRECIS, using the graphics
package CA-DISSPLA (Version 9.0) on a Burroughs
B7900 mainframe connected to a Hewlett-Packard
HP7550A plotter through a Burroughs B28 local area

network. The provincial boundaries in South Africa were
added programmatically to the basic ‘cylindrically
equidistant’ map projection produced by DISSPLA.

Each dot on the map represents a record in a quarter
degree latitude/longitude grid in Specimen-PRECIS, but the
maps were plotted at a half degree scale so that the dots
were visible on these small maps. In a few cases the
Specimen-PRECIS data was insufficient, and distributions
were obtained from other herbaria and specialists
(especially in Pentaschistis and a number of introduced
genera). Quality control was carried out at two levels:
specimen identifications were checked and corrected in
Specimen-PRECIS as each genus was studied, and when
draft maps were plotted records that appeared unusual were
checked in the herbarium.

A note on typesetting

Generic keys and generic and species descriptions were
printed on a photosetter as camera-ready bromides directly
from the DELTA format data through the program
TYPSET, which produces ASCII files with embedded
instructions for Postscript printers and photosetters. All
other text has been similarly printed using TYPSET on
ASCII files produced by Wordstar Professional 5.0.

17

KEYS TO GENERA

Identifying grasses by printed keys is not easy. This
book covers 194 genera and 957 species and infraspecific
taxa; to distinguish them it is usually necessary to observe
details of spikelet structure, which often require magnifica-
tion. Furthermore, spikelets at a stage suitable for
identification exist on the plant for only a short time during
the year. Equipment needed to make the observations
essential to use these keys includes a lOx lens with a stand
(or, better, a dissecting microscope), tweezers with very
sharp tips, a dissecting needle, and a ruler accurate to 0.1
mm. The index fingernail is useful to hold down small
structures.

The key consists of sets (usually a pair) of one to four
contrasting characters that lead to further sets of contrasts,
and ultimately to a tentative identification. Each set of
contrasts is designated by a number, and the number
following in parentheses indicates the previous contrast that
led to the present set. In each set of contrasts the characters
most easily observed are generally given first, but it is
necessary to attempt to observe all the characters
mentioned in each set. When a tentative identification is
reached it must be verified by comparison to illustrations,
descriptions or other specimens of known identity.
Beginners may find it helpful to take a known plant and run
it through the key backward, starting with the identification
and working back (using the parenthetical numbers) to gain
practice in making the observations and interpreting the
choices.

We strongly recommend computer-aided identification
using the program INTKEY in place of these printed keys.
The advantages of this method include use of distribution
data to limit the number of genera to be considered, and
access to descriptors for any plant part in any sequence. It
is often possible using INTKEY to identify sterile or
fragmentary material to genus, which cannot be done
through the printed key.

Note that the generic keys are not strictly dichotomous,
and in some sets several choices are offered. Up to four
contrasting characters may be included in each comparison.
These are presented in the order that is assumed to be
easiest to see, or to be most important in making the choice.
The keys were computer-generated through the DELTA
computer system and the program KEY. The method is
briefly described on p. 14, and the KEY program
parameters are given on p. 407.

Key to Keys

1(0). Tall plants, to (2—) 3 m or more in height; culms
woody and persistent, always leafy (reeds and

bamboos) Key 1 (p. 17)

Plants seldom reaching 3 m in height; culms
herbaceous or uncommonly woody, leafy or not

leafy 2

2(1). Female-fertile spikelets with proximal incomplete

florets or empty lemmas Key 2 (p. 17)

Female-fertile spikelets without proximal incomplete

florets or empty lemmas ■ 3

3(2). Inflorescence a single spike (or false spike), a single
raceme or composed of spikelike main branches .

Key 3 (p. 22)

Inflorescence paniculate (occasionally a panicle may
be very contracted, with the branches hidden by the
spikelets) Key 4 (p. 25)

Key 1.

1(0). Leaves pseudopetiolate 2

Leaves not pseudopetiolate 4

2(1). Plants scrambling on forest vegetation; culms less

than 10 mm in diameter Olyra

Plants not scrambling; culms more than 10 mm in

diameter 3

3(2). Plants occurring naturally in mountainous areas ot

Lesotho, Natal and eastern Cape

Thamnocalamus

Plants escaped from cultivation, mainly in Natal. . .

Bambusa

Plants cultivated in the Transvaal, near Sibasa ....

Oxytenanthera

4(1). Female-fertile lemmas conspicuously hairy; ligule
hairs to 0.3 mm long, shorter than their subtending

membrane Arundo

Female-fertile lemmas hairless; ligule hairs longer
than 0.5 mm, longer than their subtending
membrane Phragmites

Key 2.

1(0). Plants bisexual, but monoecious, all the fertile

spikelets unisexual 2

Plants bisexual with bisexual spikelets 4

2(1). Inflorescences in hard, globular, 6-12 mm utricles;
female-fertile lemmas with a single median keel on

the back Coix

Inflorescences not as in Coix\ female-fertile lemmas
rounded, flat or with two or more keels on the back

3

3(2). Tall plants to 3 metres or more high; culms woody and
persistent; leaves pseudopetiolate, blades wider
than 15 mm, with transverse veins readily visible

at least abaxially Olyra

Plants never reaching 3 metres in height; culms
herbaceous; leaves not pseudopetiolate, blades
narrower than 15 mm, with transverse veins very

inconspicuous Andropogon festuciformis

4(1). Female-fertile floret 1 per female-fertile spikelet . 5
Female-fertile florets more than 1 per female-fertile

spikelet 113

5(4). Spikelets with bractiform involucres 6

Spikelets without bractiform involucres, not
subtended by scabrid bristles (vestigial branches)

8

Spikelets subtended by several scabrid bristles

(vestigial branches) 108

Spikelets (or at least some of them) subtended by
solitary scabrid bristles (vestigial branches) . 110
6(5). Inflorescence leafy or spatheate, comprising a
complex of ‘partial inflorescences’ and intervening
foliar organs; female-fertile lemmas mucronate or

awned Themeda

Inflorescence not leafy, not spatheate, not comprising
‘partial inflorescences’ and foliar organs; female-
fertile lemmas neither mucronate nor awned ... 7
7(6). Glumes of female-fertile spikelets awned,
approximately equalling or longer than the adjacent
lemmas; proximal incomplete florets epaleate,

sterile Anthephora

Glumes of female-fertile spikelets not awned, all
shorter than the adjacent lemmas in the intact

18

spikelets; proximal incomplete florets paleate (but
the palea may be reduced or vestigial), male ....

Odontelytrum

8(5). Female-fertile spikelets disarticulating above the

glumes 9

Female-fertile spikelets falling with the glumes . 23
Female-fertile spikelets not disarticulating .... 105
9(8). Female-fertile lemmas with a single median keel on

the back 10

Female-fertile lemmas rounded, flat or with two or

more keels on the back 15

10(9). Tall plants to 3 metres or more high; culms
scandent; leaves clinging by retrorsely scabrid
blade margins; grain longitudinally grooved . .

Prosphytochloa

Plants never reaching 3 metres in height; culms not
scandent; leaves not as in Prosphytochloa ; grain

not grooved 11

1 1(10). Incomplete florets proximal to the female-fertile
florets; hilum long-linear; lodicules membranous

12

Incomplete florets both distal and proximal to the
female-fertile florets; hilum short; lodicules

fleshy Ctenium

12(11). Hairy callus present Microlaena

No hairy callus 13

13(12). Palea thinner than the lemma Ehrharta

Palea similar in texture to the lemma 14

14(13). Leaf auricles present; female-fertile palea back 1-
keeled; stamens 5-6 per floret; spikelets very
unconventional and hard to interpret . . . Oryza
Leaf auricles absent; female-fertile palea back
rounded; stamens 3 per floret; spikelets more or
less conventional, with readily identifiable

glumes, lemmas and paleas Phalaris

15(9). Ligule not fringed; embryo less than 1/3 the length

of the grain 16

Ligule fringed; embryo at least 1/3 as long as the

grain 17

16(15). Panicle contracted; female-fertile lemmas
becoming distinctly indurated when mature;

palea entire; fresh shoots aromatic

Anthoxanthum

Panicle open; female-fertile lemmas not becoming
indurated; palea apically notched; fresh shoots

not aromatic Arrhenatherum

17(15). Female-fertile lemmas neither mucronate nor

awned 18

Female-fertile lemmas mucronate or awned .. 19
18(17). Proximal lemmas of the female-fertile spikelets
awned, similar in texture to the female-fertile
lemmas; female-fertile palea apically notched .

Melinis

Proximal lemmas of the female-fertile spikelets not
awned, less firm than the female-fertile lemmas;

female-fertile palea entire Tricholaena

19(17). Mature female-fertile lemmas with a clear
germination flap; stamens 3 per female-fertile

floret 20

No germination flap in the female-fertile lemmas;
stamens 2 per female-fertile floret (or 3 in
Loudetia filifolia, L. flavida, L. pedicellata) . 22
20(19). Callus short; glumes very unequal in the intact
female-fertile spikelet; ovary apex glabrous . 21
Callus long; glumes equal or subequal in length in
the intact female-fertile spikelet; ovary apex

hairy Tristachya

21(20). Female-fertile lemmas conspicuously hairy, deeply
cleft; palea apically notched, the keels winged;

hilum long-linear Danthoniopsis

Female-fertile lemmas not conspicuously hairy, not
deeply cleft; palea entire, the keels wingless;

hilum short Arundinella

22(19). Female-fertile lemmas deeply cleft; panicle

branchlets capillary; stigmas white

Trichopteryx

Female-fertile lemmas not deeply cleft; panicle
branchlets not capillary; stigmas golden-brown

Loudetia

23(8). Female-fertile lemmas less firm than the glumes .

24

Female-fertile lemmas and glumes of similar

texture 70

Female-fertile lemmas decidedly firmer than the

glumes 80

24(23). Female-fertile lemmas neither mucronate nor

awned 25

Female-fertile lemmas mucronate or awned . . 40
25(24). Inflorescence comprising a complex of ‘partial
inflorescences’ and intervening foliar organs .

26

Inflorescence not comprising ‘partial

inflorescences’ and foliar organs 29

26(25). Pedicels of the ‘pedicellate’ spikelets discernable,

but fused with the rachis 27

Pedicels of the ‘pedicellate’ spikelets free of the

rachis 28

27(26). Lower glume of female-fertile spikelet globose,
lacunose; ‘pedicellate’ spikelets reduced,
herbaceous; lemmas of proximal incomplete

florets nerveless Hackelochloa

Lower glume of female-fertile spikelet flattish, not
lacunose; ‘pedicellate’ spikelets similar to the
female-fertile spikelets; lemmas of proximal

incomplete florets 2-nerved Hemarthria

Lower glume of female-fertile spikelet flattish, not
lacunose; ‘pedicellate’ spikelets reduced,
herbaceous; lemmas of proximal incomplete

florets 3-4-nerved Rottboellia

28(26). Hairy callus present; lemmas of proximal
incomplete florets decidedly longer than the
female-fertile lemmas; articulations of the
spikelet-bearing axes distinctly oblique; culms
unbranched vegetatively above; female-fertile

lemmas nerveless Elionurus

No hairy callus; lemmas of proximal incomplete
florets more or less equalling the female-fertile
lemmas; articulations of the spikelet-bearing
axes more or less transverse; culms branching
vegetatively above; female-fertile lemmas 5-

nerved Coelorhachis

29(25). Inflorescence a single spike, with no pedicellate

spikelets Oxyrhachis

Inflorescence of spike-like main branches .... 30

Inflorescence a single raceme 34

Inflorescence paniculate 36

30(29). Ligule not fringed 31

Ligule fringed 32

31(30). Lower glume of the pedicellate spikelet with an
awn 5-10 mm (or longer); articulations of the
spikelet-bearing axes distinctly oblique; callus
long, hairy; fresh shoots aromatic . Urelytrum
Lower glume of the pedicellate spikelet awnless;
articulations of the spikelet-bearing axes more or
less transverse; callus short, not hairy; fresh

shoots not aromatic Phacelurus

32(30). Inflorescence digitate or subdigitate; glumes very
unequal in the intact female-fertile spikelet,
lower glume flattened on the back; proximal
incomplete florets of the female-fertile spikelets
paleate (but the palea may be reduced or

vestigial), male Vossia

Inflorescence neither digitate nor subdigitate;
glumes equal or subequal in length in the intact
female-fertile spikelet, lower glume convex on
the back; proximal incomplete florets of the
female-fertile spikelets epaleate, sterile .... 33
33(32). Panicle open; spikelet-bearing rachises slender;

female-fertile spikelets compressed laterally;
lemmas of proximal incomplete florets 2-nerved;

culm internodes solid Vetiveria

Panicle contracted; spikelet-bearing rachises

19

substantial; female-fertile spikelets compressed
dorsiventrally; lemmas of proximal incomplete
florets nerveless; culm internodes conspicuously

hollow Eriochrysis

34(29). Articulations of the spikelet-bearing axes more or

less transverse; stigmas golden-brown

Rhytachne

Articulations of the spikelet-bearing axes distinctly
oblique; stigmas pink, red, purple or black . 35
35(34). Ligule not fringed; lower glume of the pedicellate
spikelet with an awn 5-10 mm (or longer);
proximal incomplete florets of the female-fertile
spikelets male, the lemmas more or less equalling
the female-fertile lemmas; leaf auricles present

Urelytrum

Ligule fringed; lower glume of the pedicellate
spikelet awnless; proximal incomplete florets of
the female-fertile spikelets sterile, the lemmas
decidedly longer than the female-fertile lemmas;

leaf auricles absent Elionurus

36(29). Spikelets consistently in long-and-short
combinations; glumes equal or subequal in length

in the intact female-fertile spikelet 37

Spikelets not in distinct long-and-short
combinations; glumes very unequal in the intact

female-fertile spikelet 39

37(36). Spikelets in pedicellate/sessile combinations;

glumes of female-fertile spikelets very dissimilar
in form or texture; styles free to their bases;

lodicules present 38

Spikelets all pedicellate; glumes of female-fertile
spikelets more or less similar in form and texture;

styles fused basally; lodicules absent

Imperata

38(37). Female-fertile spikelets compressed dorsiventrally;

the male and female-fertile spikelets overtly
different in form; leaves not distinctly basally

aggregated; lodicules ciliate Sorghum

Female-fertile spikelets compressed laterally; the
male and female-fertile spikelets externally
similar in form; leaves mostly basal; lodicules

glabrous Vetiveria

39(36). Female-fertile lemmas with a single median keel on
the back; spikelets 1-2 mm long; fresh shoots

aromatic

Melinis macrochaetia, minutiflora, tenuissima
Female-fertile lemmas rounded, flat or with two or
more keels on the back; spikelets 2-12 mm long;

fresh shoots not aromatic Melinis

40(24). Spikelets of sexually distinct kinds on the same
plant — e.g. female or hermaphrodite and sterile

or male-only 41

Spikelets alike in sexuality on the same plant . 66
41(40). Female-fertile spikelets compressed laterally . 42
Female-fertile spikelets not noticeably compiessed

47

Female-fertile spikelets compressed dorsiventrally

52

42(41 ). Proximal incomplete florets of the female-fertile
spikelets male, paleate (but the palea may be

reduced or vestigial) Sehima

Proximal incomplete florets of the female-fertile

spikelets sterile, epaleate 43

43(42). Inflorescence digitate or subdigitate . Arthraxon
Inflorescence neither digitate nor subdigitate . 44
44(43). The spikelet-bearing units very much reduced,
bearing one or a few spikelets . . Chrysopogon

The spikelet-bearing units ‘racemes’ 45

45(44). The male and female-fertile spikelets overtly
different in form; inflorescence leafy or
spatheate; grain compressed dorsiventrally . 46
The male and female-fertile spikelets externally
similar in form; inflorescence not leafy, not
spatheate; grain not noticeably compressed . . .
Vetiveria

46(45). Palea lacking within the female-fertile lemma; fresh

shoots aromatic Cymbopogon

Palea present within the female-fertile lemma (but
may be minute); fresh shoots not aromatic . . .

Andropogon

47(41). Lower glume of female-fertile spikelet distinctly 2-

keeled to the middle or below 48

Lower glume of female-fertile spikelet not
distinctly 2-keeled below the upper quarter . 50
48(47). Spikelets all pedicellate; female-fertile lemmas
entire; callus pointed; inflorescence not leafy, not
spatheate; mature spikelet-bearing axes not

disarticulating Trachypogon

Spikelets in pedicellate/sessile combinations;
female-fertile lemmas incised; callus blunt;
inflorescence leafy or spatheate; mature spikelet-

bearing axes disarticulating 49

49(48). Palea lacking within the female-fertile lemma; fresh

shoots aromatic Cymbopogon

Palea present within the female-fertile lemma (but
may be minute); fresh shoots not aromatic . . .

Andropogon

50(47). Ligule not fringed; racemes paired; female-fertile

lemmas incised, 1 -nerved 51

Ligule fringed; racemes solitary; female-fertile

lemmas entire, 3-4-nerved Heteropogon

51(50). The spikelet-bearing rachises slender; glumes of
female-fertile spikelets not awned; lemmas not
deeply cleft; panicle branchlets capillary; styles

free to their bases Hyparrhenia

The spikelet-bearing rachises substantial; glumes of
female-fertile spikelets awned; lemmas deeply
cleft; panicle branchlets not capillary; styles

fused basally Elymandra

52(41 ). Lower glume of female-fertile spikelet distinctly 2-

keeled to the middle or below 53

Lower glume of female-fertile spikelet not
distinctly 2-keeled below the upper quarter . 59

53(52). Female-fertile lemmas entire 54

Female-fertile lemmas incised 56

54(53). Some spikelet pairs or triplets similar in form

(homogamous) Dichanthium

All the spikelet pairs or triplets differing in form

(heterogamous) 55

55(54). Spikelets in pedicellate/sessile combinations; callus
blunt; female-fertile lemmas not conspicuously
hairy; pedicels and internodes of the rachis with
a longitudinal, translucent furrow; lemmas of

proximal incomplete florets nerveless

Bothriochloa

Spikelets all pedicellate; callus pointed; female-
fertile lemmas conspicuously hairy; pedicels and
internodes of the rachis without a longitudinal,
translucent furrow; lemmas of proximal

incomplete florets 2-nerved .... Trachypogon

56(53). Racemes solitary Schizachyrium

Racemes paired 57

Racemes in groups Ischaemum

57(56). Fresh shoots aromatic; palea lacking within the
female-fertile lemma Cymbopogon

Fresh shoots not aromatic; palea present within the
female-fertile lemma (but may be minute) . . 58
58(57). Palea 3/4 or more of female-fertile lemma length

Ischaemum

Palea conspicuous but less than 3/4 of female-

fertile lemma length Diheteropogon

Palea minute Andropogon

59(52). Proximal incomplete florets of the female-fertile
spikelets male, paleate (but the palea may be
reduced or vestigial); leaf blades cordate at the

base Thelepogon

Proximal incomplete florets of the female-fertile
spikelets sterile, epaleate; leaf blades not cordate
at the base 60

20

60(59). Female-fertile lemmas entire Heteropogon

Female-fertile lemmas incised 61

61(60). Inflorescence leafy or spatheate, comprising a
complex of ‘partial inflorescences’ and

intervening foliar organs 62

Inflorescence not leafy, not spatheate, not
comprising ‘partial inflorescences’ and foliar

organs 65

62(61). Racemes paired; some spikelet pairs or triplets

similar in form (homogamous) 63

Racemes solitary; all the spikelet pairs or triplets

differing in form (heterogamous)

Monocymbium

63(62). Lower glume of female-fertile spikelet convex on

the back, upper glume 2-3-nerved 64

Lower glume of female-fertile spikelet sulcate on
the back, upper glume 1 -nerved . . Hyperthelia
64(63). The spikelet-bearing rachises slender; glumes of
female-fertile spikelets not awned; panicle
branchlets capillary; styles free to their bases;

female-fertile lemmas not deeply cleft

Hyparrhenia

The spikelet-bearing rachises substantial; glumes of
female-fertile spikelets awned; panicle
branchlets not capillary; styles fused basally;
female-fertile lemmas deeply cleft . Elymandra
65(61). Spikelets apparently solitary, each accompanied by

a barren pedicel; lodicules glabrous

Sorghastrum

Spikelets paired, all pedicels spikelet-bearing;

lodicules ciliate Sorghum

66(40). Female-fertile lemmas conspicuously hairy ... 67
Female-fertile lemmas not conspicuously hairy . .

68

67(66). Perennial; spikelets consistently paired, in long-
and-short combinations; lower glume of female-
fertile spikelet distinctly 2-keeled to the middle

or below; lemmas entire Miscanthus

Annual; spikelets solitary, not in distinct long-and-
short combinations; lower glume of female-
fertile spikelet not distinctly 2-keeled below the
upper quarter; lemmas incised . . . Cleistachne

68(66). The spikelet-bearing rachises slender 69

The spikelet-bearing rachises substantial

Ischaemum

69(68). Articulations of the spikelet-bearing axes more or

less transverse Microstegium

Articulations of the spikelet-bearing axes distinctly

oblique Eulalia

70(23). Female-fertile lemma margins lying flat and

exposed on the palea (Digitaria-type) 71

Female-fertile lemma margins tucked in onto the

palea (Paspalum-typt) 76

71(70). Ligule not fringed 72

Ligule fringed 73

72(71). Proximal lemmas less firm than the female-fertile
lemmas; mature lemmas with a clear germination
flap; leaves not distinctly basally aggregated;
mature spikelet-bearing axes not disarticulating

Digitaria

Proximal lemmas similar in texture to the female-
fertile lemmas; no germination flap in the
lemmas; leaves mostly basal; mature spikelet-

bearing axes disarticulating Tarigidia

73(71). Inflorescence of spike-like main branches; spikelets
secund, consistently in long-and-short

combinations; leaves mostly basal

Alloteropsis

Inflorescence paniculate; spikelets not secund, not
in distinct long-and-short combinations; leaves

not distinctly basally aggregated 74

74(73). Fresh shoots aromatic; female-fertile lemmas with

a single median keel on the back Melinis

Fresh shoots not aromatic; female-fertile lemmas
rounded, flat or with two or more keels on the
back 75

75(74). Female-fertile spikelets compressed laterally;

lemmas neither mucronate nor awned, 5-nerved,
without a germination flap; the palea apically

notched Melinis

Female-fertile spikelets compressed dorsiventrally;
lemmas mucronate or awned, 6-7-nerved, with a
clear germination flap when mature; the palea

entire Oryzidium

76(70). Hairy callus present Oplismenus

No hairy callus 77

77(76). Upper glume of female-fertile spikelet spiny;

transverse veins readily visible in the leaf blade
at least abaxially; female-fertile spikelets

compressed laterally Pseudechinolaena

Upper glume of female-fertile spikelet not spiny;
transverse veins very inconspicuous in the blade;
female-fertile spikelets compressed

dorsiventrally 78

78(77). Female-fertile lemmas conspicuously hairy

Entolasia

Female-fertile lemmas not conspicuously hairy . .

79

79(78). Inflorescence of spike-like main branches

Paspalum

Inflorescence paniculate Panicum

80(23). Female-fertile lemmas neither mucronate nor

awned 81

Female-fertile lemmas mucronate or awned . 102
81(80). Female-fertile lemma margins lying flat and

exposed on the palea (Digitaria- type) 82

Female-fertile lemma margins tucked in onto the

palea (Paspalum- type) 87

82(81). Proximal lemmas of the female-fertile spikelets

awned; lodicules absent Stereochlaena

Proximal lemmas of the female-fertile spikelets not

awned; lodicules present 83

83(82). Ligule not fringed 84

Ligule fringed 85

84(83). Inflorescence of spike-like main branches; spikelets
secund; female-fertile spikelets compressed
dorsiventrally; glumes not 1 -keeled: rounded, flat
or with more than one keel; lemmas rounded, flat

or with two or more keels on the back

Digitaria

Inflorescence paniculate; spikelets not secund;
female-fertile spikelets compressed laterally;
glumes 1 -keeled to middle or below; lemmas

with a single median keel on the back

Phalaris

85(83). Spikelets consistently in long-and-short
combinations; glumes equal or subequal in length
in the intact female-fertile spikelet; lemmas of
proximal incomplete florets becoming indurated

Megaloprotachne

Spikelets not in distinct long-and-short
combinations; glumes very unequal in the intact
female-fertile spikelet; lemmas of proximal
incomplete florets not becoming indurated . 86
86(85). Female-fertile spikelets not secund, compressed
laterally, spikelets not all embedded in the rachis;
mature spikelet-bearing axes not disarticulating

Tricholaena

Female-fertile spikelets secund, compressed
dorsiventrally, more or less embedded in the
rachis; mature spikelet-bearing axes

disarticulating Stenotaphrum

87(81). Inflorescence of spike-like main branches .... 88

Inflorescence paniculate 96

88(87). Female-fertile spikelets abaxial (with the lower
glume on the side away from the rachis or with

the upper lemma against the rachis) 89

Female-fertile spikelets adaxial (with the lower
glume against the rachis or with the upper lemma

on the side away from the rachis) 92

89(88). Leaf blades cordate at the base; female-fertile
lemmas crested at the tip Acroceras

21

Leaf blades not cordate at the base; female-fertile

lemmas not crested 90

90(89). Hairy callus present Oplismenus

No hairy callus 91

91(90). Glumes and/or sterile lemmas awned or acuminate-

mucronate Echinochloa

Spikelets awnless, muticous Paspalum

Spikelets awnless, the female-fertile lemmas

pointed or apiculate but not mucronate

Paspalidium

92(88). Female-fertile spikelets with one glume; all
spikelets slightly embedded in the rachis; leaf

tips rounded Axonopus

Female-fertile spikelets with two glumes; spikelets
not embedded in the rachis; leaf tips not rounded

93

93(92). Upper glume of female-fertile spikelet spiny;

transverse veins readily visible in the leaf blade

at least abaxially Pseudechinolaena

Upper glume of female-fertile spikelet not spiny;
transverse veins very inconspicuous in the blade

94

94(93). Female-fertile lemmas conspicuously hairy

Entolasia

Female-fertile lemmas not conspicuously hairy . .

95

95(94). Female-fertile lemma smooth Echinochloa

Female-fertile lemma striate or rugose

Brachiaria

96(87). Glumes of female-fertile spikelets with distinct

rows of hairs Leucophrys

Glumes of female-fertile spikelets without

conspicuous tufts or rows of hairs 97

97(96). Upper glume distinctly saccate Sacciolepis

Glumes not saccate 98

98(97). Female-fertile lemmas crested at the tip

Acroceras

Female-fertile lemmas not crested 99

99(98). Female-fertile spikelets compressed laterally 100
Female-fertile spikelets compressed dorsiventrally
101

100(99). Ligule not fringed; glumes equal or subequal in
length in the intact female-fertile spikelet;
female-fertile lemmas with a single median
keel on the back; proximal incomplete florets
of the female-fertile spikelets sterile, the
lemmas shorter than the female-fertile lemmas

Phalaris

Ligule fringed; glumes very unequal in the intact
female-fertile spikelet; female-fertile lemmas
rounded, flat, or with two or more keels on the
back; proximal incomplete florets of the
female-fertile spikelets male, the lemmas
decidedly longer than the female-fertile

lemmas Tricholaena

101(99). Female-fertile lemmas conspicuously hairy;

pedicel apices discoid Entolasia

Female-fertile lemmas not conspicuously hairy;

pedicel apices cupuliform Panicum

102(80). Inflorescence of spike-like main branches;

spikelets secund; lemmas of the proximal
incomplete florets less firm than the female-
fertile lemmas; female-fertile lemma margins
tucked in onto the palea (Paspalum- type) . .

103

Inflorescence paniculate; spikelets not secund;
lemmas of the proximal incomplete florets
similar in texture to the female-fertile lemmas;
female-fertile lemma margins lying flat and

exposed on the palea (Digitaria- type)

Oryzidium

103(102). Female-fertile spikelets abaxial (with the lower
glume on the side away from the rachis or with

the upper lemma against the rachis)

Urochloa

Female-fertile spikelets adaxial (with the lower
glume against the rachis or with the upper
lemma on the side away from the rachis) . . .

104

104(103). Proximal lemmas more or less equalling the
female-fertile lemmas; no beadlike ‘callus’

supporting the spikelets Brachiaria

Proximal lemmas decidedly longer than the
female-fertile lemmas; spikelets supported on

a hardened beadlike ‘callus’ Eriochloa

105(8). Female-fertile spikelets compressed laterally .

106

Female-fertile spikelets compressed

dorsiventrally 107

106(105). Ligule not fringed; female-fertile lemmas with a
single median keel on the back; glumes equal
or subequal in length; proximal incomplete
florets of the female-fertile spikelets sterile,
the lemmas shorter than the female-fertile

lemmas Phalaris

Ligule fringed; female-fertile lemmas rounded,
flat or with two or more keels on the back;
glumes very unequal; proximal incomplete
florets of the female-fertile spikelets male, the
lemmas decidedly longer than the female-

fertile lemmas Tricholaena

107(105). Glumes very unequal in the intact female-fertile
spikelet; spikelets not in distinct long-and-
short combinations; lemmas of the proximal
incomplete florets less firm than the female-
fertile lemmas; spikelets alike in sexuality on
the same plant; female-fertile lemmas entire

Panicum

Glumes equal or subequal in length in the intact
female-fertile spikelet; spikelets consistently
in long-and-short combinations; lemmas of the
proximal incomplete florets similar in texture
to the female-fertile lemmas; spikelets of
sexually distinct kinds on the same plant —
e.g. female or hermaphrodite and sterile or
male-only; female-fertile lemmas incised . .

Sorghum

108(5). The ‘bristles’ spiny, markedly coalescent basally
(but bristles slender, ciliate, coalescent only at

the extreme base in Cenchrus ciliaris)

Cenchrus

The ‘bristles’ relatively slender, not spiny . 109
109(108). The ‘bristles’ persisting on the rachis; mature
spikelet-bearing axes not disarticulating;
female-fertile lemmas becoming distinctly
indurated when mature, the margins tucked in
onto the palea (Paspalum- type); paleas

indurated Setaria

The ‘bristles’ deciduous with the spikelets;
mature spikelet-bearing axes disarticulating;
female-fertile lemmas not becoming indurated,
the margins lying flat and exposed on the palea

(Digitaria- type); paleas not indurated

Pennisetum

1 10(5). Glumes of female-fertile spikelets all minute .

Paratheria

Glumes of female-fertile spikelets large . . Ill
111(110). Lower glume of female-fertile spikelet nerveless

112

Lower glume of female-fertile spikelet 2— 5(— 6)-

nerved Setaria

Lower glume of female-fertile spikelet 7-10-

nerved Sorghastrum

112(111). Proximal incomplete florets of the female-fertile
spikelets male, paleate (but the palea may be
reduced or vestigial), the lemmas less firm than
the female-fertile lemmas; glumes pointed;

palea back rounded Odontelytrum

Proximal incomplete florets of the female-fertile
spikelets sterile, epaleate, the lemmas similar
in texture to the female-fertile lemmas; glumes

22

not pointed; palea back 2-keeled

Pennisetum unisetum

113(4). Plant a reed; inflorescence a plumose panicle .

Phragmites

Plant not a reed; inflorescence of spikelike main

branches; lemmas awnless Tetrachne

Plant not a reed; inflorescence a raceme or much
contracted panicle; lemmas with hard prickly
awns Entoplocamia

Key 3.

1(0). Inflorescence a single spike 2

Inflorescence of spike-like main branches 20

Inflorescence a false spike, with clusters of spikelets

49

Inflorescence a single raceme 55

Inflorescence paniculate 69

2(1). Female-fertile floret 1 per female-fertile spikelet . 3
Female-fertile florets more than 1 per female-fertile

spikelet 11

3(2). Female-fertile lemmas with a single median keel on

the back 4

Female-fertile lemmas rounded, flat or with two or

more keels on the back 6

4(3). Female-fertile spikelets disarticulating above the
glumes; glumes very unequal, upper glume 2-3-
nerved; female-fertile lemmas and glumes of
similar texture; lemmas conspicuously hairy ....

Harpochloa

Female-fertile spikelets falling with the glumes;
glumes equal or subequal in length, upper glume 1-
nerved; female-fertile lemmas less firm than the

glumes; lemmas not conspicuously hairy 5

5(4). Plants caespitose; glumes of female-fertile spikelets
awned, more or less similar in form and texture;
palea conspicuous but less than 3/4 of female-

fertile lemma length, nerveless Perotis

Plants long-rhizomatous or stoloniferous; glumes of
female-fertile spikelets not awned, very dissimilar
in form or texture; palea 3/4 or more of female-
fertile lemma length, with 2 well separated nerves

Mosdenia

6(3). Glumes of the female-fertile spikelets dorsiventral to

the rachis 7

Glumes of the female-fertile spikelets lateral to the

rachis 10

Glumes displaced, lateral to each other on the side

away from the rachis Parapholis

7(6). Female-fertile lemmas mucronate or awned; hairy

callus present 8

Female-fertile lemmas neither mucronate nor awned;

no hairy callus 9

8(7). Spikelets biseriate on one side of the rachis; female-
fertile spikelets compressed dorsiventrally; lower
glume 1 -nerved; lemmas decidedly firmer than the

glumes Enteropogon

Spikelets distichously arranged on opposite sides of
the rachis; female-fertile spikelets compressed
laterally; lower glume nerveless; lemmas less firm

than the glumes Oropetium

9(7). Annual; female-fertile spikelets abaxial (with the
lower glume on the side away from the rachis or
with the upper lemma against the rachis), with
female-fertile florets only; pericarp fused; embryo
less than 1/3 the length of the grain . . Hainardia
Perennial; female-fertile spikelets adaxial (with the
lower glume against the rachis or with the upper
lemma on the side away from the rachis), with
incomplete florets in addition to the female-fertile
florets; pericarp free; embryo at least 1/3 as long
as the grain Lepturus

10(6). Female-fertile lemmas 2-nerved; spikelets with
female-fertile florets only; embryo at least 1/3 as

long as the grain Microchloa

Female-fertile lemmas 3^f-nerved; spikelets with
incomplete florets in addition to the female-
fertile florets; embryo less than 1/3 the length of

the grain Rendlia

11(2). Spikelets biseriate on one side of the rachis . . 12
Spikelets distichously arranged on opposite sides of

the rachis 15

Spikelets not two-ranked (not biseriate, not

distichously inserted) Tribolium

12(1 1). Inflorescence digitate or subdigitate 13

Inflorescence neither digitate nor subdigitate . 14
13(12). Female-fertile spikelets compressed dorsiventrally;

glumes very unequal; lemmas not conspicuously
hairy, back rounded, flat or with two or more

keels; pericarp fused Enteropogon

Female-fertile spikelets compressed laterally;
glumes equal or subequal in length; lemmas
conspicuously hairy, back with a single median

keel; pericarp free Tetrapogon

14(12). Rachilla of the female-fertile spikelets not
disarticulating between the florets; glumes not

pointed Prionanthium

Rachilla of the female-fertile spikelets
disarticulating between the florets; glumes

pointed Tribolium

15(1 1). Glumes of the female-fertile spikelets dorsiventral
to the rachis; all spikelets more or less embedded
in the rachis; ovary apex glabrous; lodicules

glabrous 16

Glumes of the female-fertile spikelets lateral to the
rachis; spikelets not all embedded in the rachis;

ovary apex hairy; lodicules ciliate 18

16(15). Hairy callus present; female-fertile lemmas with a
single median keel on the back; leaves mostly

basal Tripogon

No hairy callus; female-fertile lemmas rounded, flat
or with two or more keels on the back; leaves not

distinctly basally aggregated 17

17(16). Female-fertile spikelets compressed laterally,
disarticulating above the glumes; leaf auricles
present; culm intemodes conspicuously hollow;

stigmas white Lolium

Female-fertile spikelets compressed dorsiventrally,
falling with the glumes; leaf auricles absent;
culm intemodes solid; stigmas pink, red, purple

or black Lepturus

18(15). Annual; plants caespitose; glumes subulate;

lemmas with a single median keel on the back,

the nerves apically non-confluent Secale

Perennial; plants long-rhizomatous or
stoloniferous; glumes not subulate; lemmas
rounded, flat or with two or more keels on the
back, the nerves confluent towards the tip . . 19
19(18). Fresh shoots aromatic; mature spikelet-bearing axes

disarticulating Thinopyrum

Fresh shoots not aromatic; mature spikelet-bearing

axes not disarticulating Elytrigia

20(1). Inflorescence digitate or subdigitate 21

Inflorescence neither digitate nor subdigitate . 34

21(20). Hairy callus present 22

No hairy callus 29

22(21 ). Female-fertile lemmas with a single median keel on

the back 23

Female-fertile lemmas rounded, flat or with two or

more keels on the back 26

23(22). Glumes very unequal in the intact female-fertile

spikelet 24

Glumes equal or subequal in length in the intact

female-fertile spikelet 25

24(23). Spikelets with conventional intemode spacings;

callus blunt; the distal florets reduced in size but

23

not awnlike; leaves not distinctly basally

aggregated; stigmas white Chloris

Spikelets with distinctly elongated rachilla
internodes between the florets; callus pointed;
the distal florets reduced to awns; leaves mostly
basal; stigmas golden-brown .... Lophachme
25(23). Female-fertile floret 1 per female-fertile spikelet;

glumes very dissimilar in form or texture;

pericarp fused Brachyachne

Female-fertile florets more than 1 per female-fertile
spikelet; glumes more or less similar in form and

texture; pericarp free Tetrapogon

26(22). Female-fertile spikelets compressed laterally, the
lemmas conspicuously hairy; pericarp free . 27
Female-fertile spikelets compressed dorsiventrally,
the lemmas not conspicuously hairy; pericarp

fused 28

27(26). Ligule fringed; female-fertile floret 1 per female-
fertile spikelet; glumes approximately equalling
or longer than the adjacent lemmas; grain

compressed laterally Schoenefeldia

Ligule not fringed; female-fertile florets more than
1 per female-fertile spikelet; glumes shorter than
the adjacent lemmas; grain compressed

dorsiventrally Lintonia

28(26). Spikelets solitary; female-fertile spikelets
disarticulating above the glumes; glumes very
unequal, hairless; lemmas decidedly firmer than

the glumes Enteropogon

Spikelets consistently paired; female-fertile
spikelets falling with the glumes; glumes equal
or subequal, hairy; lemmas less firm than the

glumes Eulalia

29(21). Female-fertile floret 1 per female-fertile spikelet;

pericarp fused 30

Female-fertile florets more than 1 per female-fertile

spikelet; pericarp free 32

30(29). Inflorescence axes ending in spikelets; female-
fertile spikelets adaxial (with the lower glume
against the rachis or with the upper lemma on the
side away from the rachis), disarticulating above
the glumes; glumes all shorter than the adjacent

lemmas; styles free to their bases 31

Inflorescence axes not ending in spikelets; female-
fertile spikelets abaxial (with the lower glume on
the side away from the rachis or with the upper
lemma against the rachis), falling with the
glumes; glumes approximately equalling or
longer than the adjacent lemmas; styles fused

basally Spartina

31(30). Plants long-rhizomatous or stoloniferous; glumes
pointed, not awned, more or less similar in form
and texture; culm internodes conspicuously

hollow Cynodon

Plants caespitose; glumes not pointed, awned, very
dissimilar in form or texture; culm internodes

solid; plants caespitose Eustachys

32(29). Inflorescence axes ending in spikelets; styles free

to their bases 33

Inflorescence axes not ending in spikelets; styles

fused basally Dactyloctenium

33(32). Glumes very unequal in the intact female-fertile
spikelet; lemmas entire; grain not grooved . . .

Eleusine

Glumes equal or subequal in length in the intact
female-fertile spikelet; lemmas incised; grain

longitudinally grooved Acrachne

34(20). Female-fertile floret 1 per female-fertile spikelet

35

Female-fertile florets more than 1 per female-fertile

spikelet 39

35(34). Hairy callus present 36

No hairy callus 37

36(35). Female-fertile spikelets compressed laterally,
falling with the glumes; glumes lateral to the
rachis, 1 -keeled to middle or below; lemmas
neither mucronate nor awned Catalepis

Female-fertile spikelets compressed dorsiventrally,
disarticulating above the glumes; glumes
dorsiventral to the rachis, not 1 -keeled: rounded,
flat or with more than one keel; lemmas

mucronate or awned Polevansia

37(35). Glumes very unequal in the intact female-fertile

spikelet Willkommia

Glumes equal or subequal in length in the intact

female-fertile spikelet 38

38(37). Annual; female-fertile spikelets compressed
laterally; glumes awned; spikelets with
incomplete florets in addition to the female-

fertile florets Dinebra

Perennial; female-fertile spikelets compressed
dorsiventrally; glumes not awned; female-fertile

spikelets with female-fertile florets only

Craspedorhachis

39(34). Glumes all shorter than the adjacent lemmas in the

intact female-fertile spikelets 40

Glumes approximately equalling or longer than the
adjacent lemmas in the intact female-fertile

spikelets 46

40(39). Female-fertile lemmas with a single median keel on

the back 41

Female-fertile lemmas rounded, flat or with two or

more keels on the back 45

41(40). Female-fertile spikelets compressed laterally . 42
Female-fertile spikelets not noticeably compressed

Leptochloa

42(41). Glumes of the female-fertile spikelets dorsiventral

to the rachis 43

Glumes of the female-fertile spikelets lateral to the

rachis 44

43(42). Spikelets subsessile; spikelet-bearing rachises
substantial; callus absent; grain longitudinally
grooved; surface of grain sculptured Acrachne
Some spikelets pedicellate; spikelet-bearing
rachises slender; callus short; grain not grooved;

surface of grain smooth Leptochloa

44(42). Spikelets biseriate on one side of the rachis; glumes
very unequal in the intact female-fertile spikelet;
lemmas entire, pointed; the ‘racemes’ without

spikelets towards the base Pogonarthria

Spikelets distichously arranged on opposite sides of
the rachis; glumes equal or subequal in length in
the intact female-fertile spikelet; lemmas incised,
blunt; the ‘racemes’ spikelet-bearing to the base

Brachychloa

45(40). Upper glume of female-fertile spikelet 1 -nerved .

Diplachne

Upper glume of female-fertile spikelet 2-3-nerved
Coelachyrum

46(39). Ligule not fringed 47

Ligule fringed 48

47(46). Some spikelets pedicellate; spikelets not all
embedded in the rachis; rachilla of the female-
fertile spikelets not disarticulating between the
florets; lemmas with a single median keel on the

back; leaves mostly basal Bewsia

Spikelets subsessile; all spikelets more or less
embedded in the rachis; rachilla of the female-
fertile spikelets disarticulating between the
florets; lemmas rounded, flat or with two or more
keels on the back; leaves not distinctly basally

aggregated Trichoneura

48(46). Glumes of female-fertile spikelets about equalling
or longer than the spikelets, equal or subequal in
length, awned; spikelets sessile; no hairy callus

Dinebra

Glumes of female-fertile spikelets markedly shorter
than the spikelets, very unequal, not awned;

spikelets subsessile; hairy callus present

Leptocarydion

49(1). Ligule not fringed 50

Ligule fringed 51

50(49). Spikelets without bractiform involucres; female-
fertile floret 1 per female-fertile spikelet; lemmas

24

rounded, flat or with two or more keels on the
back, 5-nerved; palea 3/4 or more of lemma

length Hordeum 63(62).

Spikelets with bractiform involucres; female-fertile

florets more than 1 per female-fertile spikelet; 64(63).
lemmas with a single median keel on the back,
3-^1-nerved; palea conspicuous but less than 3/4

of lemma length Elytrophorus

51(49). Female-fertile floret 1 per female-fertile spikelet

52

Female-fertile florets more than 1 per female-fertile 65(64).

spikelet 54

52(51). Hairy callus present; glumes of female-fertile
spikelets awned; lemmas mucronate or awned

Monelytrum

No hairy callus; glumes of female-fertile spikelets
not awned; lemmas neither mucronate nor awned

53

53(52). Female-fertile spikelets disarticulating above the
glumes; upper glume not spiny; lower glume 1-
nerved; lemmas not conspicuously hairy; 66(55).

pericarp free Sporobolus

Female-fertile spikelets falling with the glumes;
upper glume spiny; lower glume nerveless;
lemmas conspicuously hairy; pericarp fused . .

Tragus

54(51). Spikelets with bractiform involucres; glumes
approximately equalling or longer than the

adjacent lemmas Elytrophorus

Spikelets without bractiform involucres; glumes all 67(66).
shorter than the adjacent lemmas in the intact

spikelets Eragrostis

55(1). Female-fertile spikelets disarticulating above the

glumes 56

Female-fertile spikelets falling with the glumes .

66

56(55). Ligule not fringed 57 68(67).

Ligule fringed 60

57(56). Sheath margins usually joined to at least 1/4 of their
length; ovary with a conspicuous apical

appendage; grain compressed laterally

Bromus

Sheath margins free; ovary without a conspicuous
apical appendage; grain compressed

dorsiventrally 58

58(57). Glumes of female-fertile spikelets 1 -keeled to 69(1).
middle or below; lemmas neither mucronate nor

awned, blunt; hilum short Catapodium 70(69).

Glumes of female-fertile spikelets not 1 -keeled:
rounded, flat or with more than one keel; lemmas
mucronate or awned, pointed; hilum long-linear

59

59(58). Glumes of female-fertile spikelets very dissimilar
in form or texture; lemmas decidedly firmer than
the glumes; palea apically notched; nodes 71(70).
glabrous; rachises neither flattened nor hollowed,

not winged Vulpia

Glumes of female-fertile spikelets more or less
similar in form and texture; lemmas and glumes
of similar texture; palea entire; nodes

conspicuously hairy; rachises hollowed 72(71).

Brachypodium

60(56). Female-fertile floret 1 per female-fertile spikelet

61

Female-fertile florets more than 1 per female-fertile

spikelet 62

61(60). Inflorescence a solitary raceme, or digitate or

subdigitate; hairy callus present; female-fertile 73(72).
lemmas mucronate or awned; pericarp fused;
grain longitudinally grooved . . . Enteropogon
Inflorescence paniculate; no hairy callus; female-
fertile lemmas neither mucronate nor awned;
pericarp free; grain not grooved . . . Sporobolus
62(60). Rachilla of the female-fertile spikelets not

disarticulating between the florets

Prionanthium

Rachilla of the female-fertile spikelets

disarticulating between the florets 63

Hairy callus present 64

No hairy callus Tribolium

Glumes of female-fertile spikelets 1 -keeled to
middle or below; ovary apex glabrous; pericarp

fused 65

Glumes of female-fertile spikelets not 1 -keeled:
rounded, flat or with more than one keel; ovary

apex hairy; pericarp free Dregeochloa

Inflorescence a solitary raceme or digitate or
subdigitate; female-fertile spikelets compressed
dorsiventrally; glumes very unequal in the intact
female-fertile spikelet; lemmas decidedly firmer
than the glumes, 3-4-nerved . . . Enteropogon
Inflorescence paniculate; female-fertile spikelets
compressed laterally; glumes equal or subequal
in length in the intact female-fertile spikelet;
lemmas and glumes of similar texture, 8-9-

nerved Merxmuellera

Spikelets with the distal incomplete florets and/or
the rachilla apex forming a terminal clavate
appendage; sheath margins joined for at least 1/4
of their length; glumes of female-fertile spikelets
not 1 -keeled: rounded, flat or with more than one

keel Melica

Spikelets without a terminal clavate appendage;
sheath margins free; glumes of female-fertile

spikelets 1 -keeled to middle or below 67

Female-fertile floret 1 per female-fertile spikelet;
lemmas entire, and with a single median keel on

the back 68

Female-fertile florets more than 1 per female-fertile
spikelet; lemmas incised, and rounded, flat or

with two.or more keels on the back

Chaetobromus

Female-fertile lemmas neither mucronate nor
awned; palea conspicuous but less than 3/4 of
female-fertile lemma length, nerveless; leaves
not distinctly basally aggregated; styles fused

basally Perotis

Female-fertile lemmas mucronate or awned; palea
3/4 or more of female-fertile lemma length, with
2 well separated nerves; leaves mostly basal;

styles free to their bases Fingerhuthia

Ligule not fringed 70

Ligule fringed 74

Spikelets with the distal incomplete florets and/or
the rachilla apex forming a terminal clavate
appendage; female-fertile spikelets falling with

the glumes; lodicules fused Melica

Spikelets without a terminal clavate appendage;
female-fertile spikelets disarticulating above the

glumes; lodicules free 71

Sheath margins joined to at least 1/4 of their length;
ovary with a conspicuous apical appendage;

grain compressed laterally Bromus

Sheath margins free; ovary without a conspicuous
apical appendage; grain compressed

dorsiventrally 72

Glumes of female-fertile spikelets 1 -keeled to
middle or below; lemmas neither mucronate nor

awned, blunt; hilum short Catapodium

Glumes of female-fertile spikelets not 1 -keeled:
rounded, flat or with more than one keel; lemmas
mucronate or awned, pointed; hilum long-linear

73

Glumes of female-fertile spikelets very dissimilar
in form or texture; lemmas decidedly firmer than
the glumes; palea apically notched; rachises
neither flattened nor hollowed, not winged;

nodes glabrous Vulpia

Glumes of female-fertile spikelets more or less
similar in form and texture; lemmas and glumes
of similar texture; palea entire; rachises

hollowed; nodes conspicuously hairy

Brachypodium

25

74(69). Female-fertile floret 1 per female-fertile spikelet

75

Female-fertile florets more than 1 per female-fertile

spikelet 77

75(74). Female-fertile lemmas neither mucronate nor

awned; glumes hairless 76

Female-fertile lemmas mucronate or awned; glumes

hairy Fingerhuthia

76(75). Female-fertile spikelets disarticulating above the
glumes; lower glume 1 -nerved; lemmas not
conspicuously hairy; no hairy callus; styles free

to their bases Sporobolus

Female-fertile spikelets falling with the glumes;
lower glume nerveless; lemmas conspicuously
hairy; hairy callus present; styles fused basally
Catalepis

77(74). Hairy callus present 78

No hairy callus , 80

78(77). Female-fertile spikelets disarticulating above the
glumes; culms unbrar.ched vegetatively above

79

Female-fertile spikelets falling with the glumes;

culms branching vegetatively above

Chaetobromus

79(78). Glumes of female-fertile spikelets 1 -keeled to
middle or below; palea similar in texture to the
lemma; leaf blades rolled; ovary apex glabrous;

stigmas white Merxmuellera

Glumes of female-fertile spikelets not 1-keeled:
rounded, flat or with more than one keel; palea
thinner than the lemma; leaf blades folded; ovary
apex hairy; stigmas pink, red, purple or black .

Dregeochloa

80(77). Upper glume of female-fertile spikelet 1-nerved .

Eragrostis

Upper glume of female-fertile spikelet 2-3-nerved

Coelachyrum

Upper glume of female-fertile spikelet (4-) 5 (-6)
nerved Tribolium

Key 4.

1(0). Ligule not fringed 2

Ligule fringed 50

2(1). Female-fertile spikelets disarticulating above the

glumes 3

Female-fertile spikelets falling with the glumes . 47
3(2). Female-fertile floret 1 per female-fertile spikelet . 4
Female-fertile florets more than 1 per female-fertile

spikelet 18

4(3). Female-fertile spikelets with readily identifiable

glumes, lemmas and paleas 5

Female-fertile spikelets without glumes,
unconventional and hard to interpret .... Leersia
5(4). Glumes very unequal in the intact female-fertile

spikelet 6

Glumes equal or subequal in length in the intact

female-fertile spikelet 7

6(5). Leaf blades wider than 15 mm, transverse veins
visible at least abaxially; glumes of female-fertile
spikelets markedly shorter than the spikelets . . .

Festuca africana

Leaf blades narrower than 15 mm, transverse veins
very inconspicuous; glumes about equalling or

longer than the spikelets Arrhenatherum

7(5). Glumes conspicuously ventricose basally

Gastridium

Glumes not ventricose 8

8(7). Spikelets secund, sexually distinct kinds present on
the same plant — e.g. female or hermaphrodite and

sterile or male-only Cynosurus

Spikelets not secund, alike in sexuality on the same
plant 9

9(8). Glumes all shorter than the adjacent lemmas in the

intact female-fertile spikelets 10

Glumes approximately equalling or longer than the
adjacent lemmas in the intact female-fertile
spikelets 11

10(9). Hairy callus present; female-fertile lemmas
mucronate or awned, becoming distinctly
indurated when mature; palea back rounded;

hilum long-linear Stipa

No hairy callus; female-fertile lemmas neither
mucronate nor awned, not becoming indurated;
palea back 2-keeled; hilum short . . Colpodium
1 1(9). Female-fertile lemmas less firm than the glumes .

12

Female-fertile lemmas and glumes of similar

texture 13

Female-fertile lemmas decidedly firmer than the

glumes 15

12(1 1). Callus hairs present, more than 0.5 mm long . . .

Calamagrostis

Callus hairs absent, or if present less than 0.5 mm

long Agrostis

13(1 1). Perennial; glumes of female-fertile spikelets 1-
keeled to middle or below; paleas with 3 or more
well separated nerves; leaves mostly basal . . .

Ammophila

Annual; glumes of female-fertile spikelets not 1-
keeled; rounded, flat or with more than one keel;
paleas with 2 well separated nerves; leaves not

distinctly basally aggregated 14

14(13). Panicle open; callus pointed; glumes of female-
fertile spikelets not awned; spikelets with
incomplete florets in addition to the female-

fertile florets; ovary apex hairy Avena

Panicle contracted; callus blunt; glumes of female-
fertile spikelets awned; female-fertile spikelets
with female-fertile florets only; ovary apex

glabrous Lagurus

15(1 1). Female-fertile florets gibbous, the lemma awn
placed off-centre; embryo at least 1/3 as long as
the grain; embryo waisted in surface view ....

Nassella

Female-fertile florets not asinMm(?//tf;embryo less
than 1/3 the length of the grain; embryo not

waisted 16

16(15). Rachilla prolonged beyond uppermost female-

fertile floret, or florets more than 2 17

Rachilla not prolonged beyond the solitary female-

fertile floret Stipa

17( 16). Panicle open; callus pointed; glumes of female-
fertile spikelets not awned; spikelets with
incomplete florets in addition to the female-

fertile florets; ovary apex hairy Avena

Panicle contracted; callus blunt; glumes of female-
fertile spikelets awned; female-fertile spikelets
with female-fertile florets only; ovary apex

glabrous Lagurus

18(3). Lemmas awned, the awn apically clavate, bearing

a ring of minute hairs at the middle

Corynephorus

Lemmas without the characteristic Corynephorus

awn 19

19(18). Leaf blades cordate at the base, transverse veins
readily visible in the blade at least abaxially;

lodicules absent in female-fertile florets

Megastachya

Leaf blades not cordate at the base; transverse veins
very inconspicuous in the blade; lodicules

present in female-fertile florets 20

20(19). Lemmas as broad as long, gibbous, cordate at the

base Briza

Lemmas not as in Briza 21

21(20). Female-fertile spikelets compressed laterally . 22
Female-fertile spikelets not noticeably compressed

Streblochaete

26

22(21). Glumes of female-fertile spikelets very dissimilar

in form or texture 23

Glumes of female-fertile spikelets more or less

similar in form and texture 24

23(22). Glumes very unequal in the intact female-fertile
spikelet, not 1 -keeled: rounded, flat or with more
than one keel; lemmas decidedly firmer than the
glumes, mucronate or awned, pointed . Vulpia
Glumes equal or subequal in length in the intact
female-fertile spikelet, 1 -keeled to middle or
below; lemmas and glumes of similar texture, the
lemmas neither mucronate nor awned, blunt . .

Catapodium

24(22). Spikelets of sexually distinct kinds on the same
plant — e.g. female or hermaphrodite and sterile

or male-only Cynosurus

Spikelets alike in sexuality on the same plant . 25
25(24). Glumes all shorter than the adjacent lemmas in the

intact female-fertile spikelets 26

Glumes approximately equalling or longer than the
adjacent lemmas in the intact female-fertile

spikelets 42

26(25). Upper glume of female-fertile spikelet 1 -nerved .

27

Upper glume of female-fertile spikelet 2-3-nerved

29

Upper glume of female-fertile spikelet (4— )5(— 6)-

nerved 40

Upper glume of female-fertile spikelet 7-10-nerved

41

27(26). Lower glume of female-fertile spikelet nerveless

Sphenopus

Lower glume of female-fertile spikelet 1 -nerved .

28

28(27). Female-fertile spikelets less than 3 mm long;

stamens 2 per female-fertile floret; lodicules
fleshy; hilum short; embryo at least 1/3 as long

as the grain Diandrochloa

Female-fertile spikelets more than 6 mm long;
stamens 3 per female-fertile floret; lodicules
membranous; hilum long-linear; embryo less

than 1/3 the length of the grain Festuca

29(26). Female-fertile lemmas and glumes of similar

texture 30

Female-fertile lemmas decidedly firmer than the

glumes 39

30(29). Female-fertile lemmas with a single median keel on

the back 31

Female-fertile lemmas rounded, flat or with two or

more keels on the back 35

31(30). Sheath margins joined to at least 1/4 of their length

32

Sheath margins free 33

32(31). Leaves mostly basal; ovary apex glabrous, without
a conspicuous apical appendage; mature grain
less than 4 mm long; caryopsis free from both

lemma and palea Poa

Leaves not distinctly basally aggregated; ovary
apex hairy, with a conspicuous apical appendage;
mature grain 4-10 mm long; caryopsis adhering

to lemma and/or palea Bromus

33(31). Palea thinner than the lemma 34

Palea similar in texture to the lemma Poa

34(33). Annual Lophochloa

Perennial Koeleria

35(30). Glumes very unequal in the intact female-fertile

spikelet 36

Glumes equal or subequal in length in the intact

female-fertile spikelet 38

36(35). Ovary with a conspicuous apical appendage; grain
compressed laterally; lodicules not toothed . . .

Bromus

Ovary without a conspicuous apical appendage;
grain compressed dorsiventrally; lodicules
toothed 37

37(36). Lemmas 3.5-11.0 mm long, tips firm, acute; hilum

long-linear Festuca

Lemmas 1-3 mm long, tips scarious, blunt, often
somewhat ragged; hilum short .... Puccinellia
38(35). Sheath margins joined to at least 1/4 of their length;

ovary apex hairy, with a conspicuous apical
appendage; caryopsis adhering to lemma and/or
palea; mature grain 4-10 mm long . . Bromus
Sheath margins free; ovary apex glabrous, without
a conspicuous apical appendage; caryopsis free
from both lemma and palea; mature grain less

than 4 mm long Catapodium

39(29). Callus pointed, hairy; female-fertile lemmas
incised, back rounded, flat or with two or more

keels; rachilla hairy Helictotrichon

Callus blunt, not hairy; female-fertile lemmas
entire, back with a single median keel; rachilla

not hairy Dactylis

40(26). Ovary with a conspicuous apical appendage; ligule
not truncate; grain compressed laterally;

lodicules not toothed Bromus

Ovary without a conspicuous apical appendage;
ligule truncate; grain compressed dorsiventrally;

lodicules toothed Festuca

41(26). Sheath margins joined to at least 1/4 of their length;

ovary with a conspicuous apical appendage;
rachises neither flattened nor hollowed, not
winged; grain compressed laterally; lodicules

glabrous Bromus

Sheath margins free; ovary without a conspicuous
apical appendage; rachises hollowed; grain
compressed dorsiventrally; lodicules ciliate . .

Brachypodium

42(25). Female-fertile lemmas with a single median keel on

the back; grain not grooved 43

Female-fertile lemmas rounded, flat or with two or
more keels on the back; grain longitudinally

grooved 44

43(42). Annual Lophochloa

Perennial Koeleria

44(42). Callus pointed; ovary apex hairy; mature grain

4-10 mm long; hilum long-linear Avena

Callus blunt; ovary apex glabrous; mature grain less

than 4 mm long; hilum short 45

45(44). Rachilla hairy; caryopsis free from both lemma and

palea Deschampsia

Rachilla not hairy; caryopsis adhering to lemma

and/or palea 46

46(45). Glumes about equalling the spikelets; spikelets with

conventional internode spacing Aira

Glumes shorter than the spikelets; spikelets with an
elongated rachilla internode between the florets

Periballia

47(2). Spikelets with the distal incomplete florets and/or
the rachilla apex forming a terminal clavate
appendage; sheath margins joined to at least 1/4

of their length; lodicules fleshy Melica

Spikelets without a terminal clavate appendage;
sheath margins free; lodicules membranous • 48
48(47). Spikelets secund; spikelets of sexually distinct
kinds on the same plant — e.g. female or
hermaphrodite and sterile or male-only; mature

spikelet-bearing axes disarticulating

Lamarckia

Spikelets not secund; spikelets alike in sexuality on
the same plant; mature spikelet-bearing axes not

disarticulating 49

49(48). Glumes of female-fertile spikelets 1 -keeled to
middle or below; upper glume 2-3-nerved;
lemmas with a single median keel on the back,
decidedly firmer than the glumes; spikelets with
incomplete florets in addition to the female-

fertile florets Holcus

Glumes of female-fertile spikelets not 1 -keeled:
rounded, flat or with more than one keel; upper

27

glume 1 -nerved; lemmas rounded, flat or with
two or more keels on the back, less firm than the
glumes; female-fertile spikelets with female-

fertile florets only Polypogon

50(1). Female-fertile spikelets disarticulating above the

glumes 51

Female-fertile spikelets falling with the glumes .

88

51(50). Female-fertile floret 1 per female-fertile spikelet

52

Female-fertile florets more than 1 per female-fertile

spikelet 63

52(51). Ovary apex hairy Pentameris sp. 2

Ovary apex glabrous 53

53(52). Female-fertile lemmas neither mucronate nor

awned 54

Female-fertile lemmas mucronate or awned . . 55
54(53). Lower glume of female-fertile spikelet 1 -nerved,
upper glume 1 -nerved; lemmas not
conspicuously hairy; embryo at least 1/3 as long

as the grain Sporobolus

Lower glume of female-fertile spikelet 2-3-nerved,
upper glume 2-3-nerved; lemmas conspicuously
hairy; embryo less than 1/3 the length of the grain

Pentaschistis pusilla

55(53). Awns trifid, usually with a basal column (but not
trifid in Aristida parvula or Stipagrostis

anomala) 56

Awns not trifid 58

56(55). Awns plumose (but not plumose in Stipagrostis
anomala)-, paleas of female-fertile florets

indurated Stipagrostis

Awns not plumose; paleas of female-fertile florets

not indurated 57

57(56). Glumes of female-fertile spikelets 1 -keeled to
middle or below; lower glume 1 -nerved; mature
female-fertile lemmas with a clear germination
flap; grain not grooved; embryo at least 1/3 as

long as the grain Aristida

Glumes of female-fertile spikelets not 1 -keeled:
rounded, flat or with more than one keel; lower
glume 2-3-nerved; no germination flap in the
female-fertile lemmas; grain longitudinally
grooved; embryo less than 1/3 the length of the

grain Sartidia

58(55). Hairy callus present 59

No hairy callus 62

59(58). Female-fertile lemmas and paleas becoming
distinctly indurated when mature; palea back
rounded; female-fertile spikelets with female-
fertile florets only; hilum long-linear .... Stipa
Female-fertile lemmas and paleas not becoming
indurated; palea back 2-keeled; spikelets with
incomplete florets in addition to the female-

fertile florets; hilum short 60

60(59). Female-fertile lemmas conspicuously hairy;

incomplete florets distal to the female-fertile

florets 61

Female-fertile lemmas not conspicuously hairy;
incomplete florets proximal to the female-fertile

florets Arundinella

61(60). Rachilla of the female-fertile spikelets not
disarticulating between the florets; lemmas
decidedly firmer than the glumes, incised, 8-9-
nerved; leaves not distinctly basally aggregated

Enneapogon

Rachilla of the female-fertile spikelets
disarticulating between the florets; lemmas and
glumes of similar texture, the lemmas entire,
3^1-nerved; leaves mostly basal .... Stiburus
62(58). Glumes of female-fertile spikelets awned; lemmas
2-lobed, 5-nerved; palea apically notched;
female-fertile spikelets with female-fertile florets

only Lagurus

Glumes of female-fertile spikelets not awned;
lemmas 9-lobed, 8-9-nerved; palea entire;

spikelets with incomplete florets in addition to

the female-fertile florets Enneapogon

63(51). Glumes all shorter than the adjacent lemmas in the

intact female-fertile spikelets 64

Glumes approximately equalling or longer than the
adjacent lemmas in the intact female-fertile

spikelets 70

64(63). Leaves hard, woody, needle-like, plants prickly .

65

Leaves not needle-like, plants not prickly .... 66
65(64). Glumes very unequal in the intact female-fertile
spikelet, lower glume 1 -nerved, upper glume 1
nerved; lemmas incised, mucronate . . Odyssea
Glumes equal or subequal in length in the intact
female-fertile spikelet, lower glume 2-3-nerved,
upper glume 2-3-nerved; lemmas entire, neither

mucronate nor awned Cladoraphis

66(64). Hairy callus present 67

No hairy callus 69

67(66). Female-fertile lemmas entire Stiburus

Female-fertile lemmas incised 68

68(67). Female-fertile lemmas deeply cleft; grain
trigonous; fruit linear; hilum short; embryo at

least 1/3 as long as the grain Triraphis

Female-fertile lemmas not deeply cleft; grain not
noticeably compressed; fruit fusiform; hilum
long-linear; embryo less than 1/3 the length of

the grain Styppeiochloa

69(66). Upper glume of female-fertile spikelet 1 -nerved .

Eragrostis

Upper glume of female-fertile spikelet 2-3-nerved

Coelachyrum

Upper glume of female-fertile spikelet (4— )5(— 6)-

nerved Tribolium

70(63). Rachilla of the female-fertile spikelets not

disarticulating between the florets 71

Rachilla of the female-fertile spikelets

disarticulating between the florets 73

71(70). Callus absent; female-fertile lemmas not deeply
cleft, becoming distinctly indurated when
mature; palea apically notched . . Kaokochloa
Callus short; female-fertile deeply cleft, not

becoming indurated; palea entire 72

72(71). Female-fertile lemmas 9-lobed, with 9 awns (one
terminating each lobe); anthers up to 2.5 mm

long Enneapogon

Female-fertile lemmas 6-lobed, with 5 awns (one
arising between each pair of lobes); anthers more

than 2.5 mm long Schmidtia

73(70). Panicle open 74

Panicle contracted 77

74(73). Callus short, blunt 75

Callus long, pointed Cortaderia

75(74). Leaf blades wider than 15 mm; plants long-
rhizomatous or stoloniferous; young vegetative
shoots bursting through the bases of subtending

sheaths Arundo

Leaf blades narrower than 15 mm; plants usually
caespitose; young vegetative shoots emerging
from between the subtending sheaths and the

stem 76

76(75). Ovary apex hairy; female-fertile lemmas deeply
cleft; fruit with a hard, thick pericarp; hilum

long-linear Pentameris

Ovary apex glabrous; female-fertile lemmas not
deeply cleft; fruit with a membranous pericarp;

hilum short Pentaschistis

77(73). Female-fertile lemmas entire 78

Female-fertile lemmas incised 79

78(77). Hairy callus present; lower glume of female-fertile
spikelet 1 -nerved, upper glume 1 -nerved; grain

not noticeably compressed Stiburus

No hairy callus; lower glume of female-fertile
spikelet (4-)5(-6)-nerved, upper glume
(4-)5(-6)-nerved; grain compressed
dorsiventrally Tribolium

28

79(77). Palea thinner than the lemma 80

Palea similar in texture to the lemma 82

80(79). Spikelets with conventional internode spacings .

81

Spikelets with a distinctly elongated internode

between the glumes Centropodia

81(80). Hairs of the female-fertile lemmas in tufts in
transverse rows; grain longitudinally grooved;
pericarp free; ovary apex hairy . . Dregeochloa
Hairs of the female-fertile lemmas not in tufts, not
in transverse rows; grain not grooved; pericarp

fused; ovary apex glabrous Schismus

82(79). Female-fertile lemmas and glumes of similar

texture 83

Female-fertile lemmas decidedly firmer than the

glumes 87

83(82). Ovary apex glabrous 84

Ovary apex hairy Pentameris

84(83). Female-fertile spikelets with female-fertile florets

only Pentaschistis

Spikelets with incomplete florets in addition to the

female-fertile florets 85

85(84). Female-fertile lemmas with a bent awn, twisted
below (except Merxmuellera macowanii) . . 86
Female-fertile lemmas awnless, mucronate or with

a short straight awn Schismus

86(85). Spikelets 8-25 mm long, inflorescence longer than
60 mm; lodicules membranous Merxmuellera
Spikelets 4-6(-7) mm long, inflorescence 10-60

mm long; lodicules fleshy Karroochloa

87(82). Ovary apex hairy; callus short, blunt; female-fertile
lemma margins lying flat and exposed on the
palea (Digitaria- type); styles free to their bases

Pentameris

Ovary apex glabrous; callus long, pointed; female-
fertile lemma margins tucked in onto the palea

(Paspalum- type); styles fused basally

Pseudopentameris

88(50). Lower glume of female-fertile spikelet nerveless

Catalepis

Lower glume of female-fertile spikelet 1 -nerved .

89

Lower glume of female-fertile spikelet (4— )5(— 6)-

nerved 91

Lower glume of female-fertile spikelet 7-1 0-nerved
93

89(88). Glumes of female-fertile spikelets markedly shorter
than the spikelets, shorter than the adjacent

lemmas; palea thinner than the lemma

Eragrostis

Glumes of female-fertile spikelets about equalling
or longer than the spikelets and the adjacent
lemmas; palea similar in texture to the lemma .

90

90(89). Upper glume of female-fertile spikelet 1 -nerved;

the distal florets seemingly merely
underdeveloped, neither clearly specialised nor
peculiarly modified in form; leaves mostly basal;

lodicules fleshy Fingerhuthia

Upper glume of female-fertile spikelet 2-3-nerved;
the distal florets clearly specialised, peculiarly
modified; leaves not distinctly basally
aggregated; lodicules membranous . . . Holcus
91(88). Spikelets of sexually distinct kinds on the same
plant — e.g. female or hermaphrodite and sterile
or male-only; glumes of female-fertile spikelets
awned; lemmas entire; inflorescence deciduous
in its entirety as a ‘tumbleweed’; pericarp free

Urochlaena

Spikelets alike in sexuality on the same plant;
glumes of female-fertile spikelets not awned;
lemmas incised; inflorescence not becoming

tumbleweed; pericarp fused 92

92(91). Culms branching vegetatively above; female-fertile
lemmas with a bent awn, twisted below; grain
longitudinally grooved, compressed laterally;

hilum long-linear Chaetobromus

Culms unbranched vegetatively above; female-
fertile lemmas awnless, mucronate or with a short
straight awn; grain not grooved, compressed

dorsiventrally; hilum short Schismus

93(88). Culms branching vegetatively above; female-fertile
lemmas with a bent awn, twisted below; grain
longitudinally grooved, compressed laterally;

hilum long-linear Chaetobromus

Culms unbranched vegetatively above; female-
fertile lemmas awnless, mucronate or with a short
straight awn; grain not grooved, compressed
dorsiventrally; hilum short Schismus

29

SYNOPSIS OF CLASSIFICATION

In the main body of this book the genera appear in
alphabetical order. The convenience of this arrangement
seemed to outweigh the advantages to be gained from
imposing an overall taxonomic sequence, especially given
the instability of higher classification in the grasses. The
main disadvantage to an alphabetical arrangment is that
groups of related genera are not placed together in the text.
The following outline of grass classification is therefore
included to serve as a quick reference to indicate supposed
relationships. The sequences of genera were developed by
means of INTKEY operations in conjunction with the
results of numerical analysis of the descriptions in the world
generic database. They are intended to reflect true taxonom-
ic relationships and thus to indicate where problems may
be encountered in identifying closely related genera.

A full formal classification of the southern African
grasses, with descriptions of subfamilies, supertribes, tribes
and subtribes appears on p. 381. The number of genera and
species occurring in each subfamily in southern Africa is
given in Table 1, p. 5.

Family: POACEAE

Subfamily: POOIDEAE

Supertribe: Triticodae

Tribe: Triticeae. Secale, Hordeum, Elytrigia,
Thinopyrum

Tribe: Brachypodieae. Brachypodium
Tribe: Bromeae. Bromus
Supertribe: Poodae

Tribe: Aveneae ( including Agrostideae, Phalarideae).
Avena, Arrhenatherum, Helictotrichon,
Anthoxanthum, Holcus, Phalaris, Koeleria,
Lophochloa, Agrostis, Polypogon, Calamagrostis,
Ammophila, Gastridium, Lagurus, Corynephorus,
Deschampsia, Aira, Periballia

Tribe: Meliceae. Melica , Streblochaete

Tribe: Poeae ( including Hainardieae, Monermeae).
Puccinellia, Colpodium, Festuca, Folium, Vulpia,
Catapodium, Poa, Sphenopus, Briza, Dactylis,
Cynosurus, Lamarckia, Hainardia, Parapholis

Subfamily: BAMBUSOIDEAE
Supertribe: Oryzodae

Tribe: Oryzeae. Leersia, Oryza, Prosphytochloa

Tribe: Olyreae. Olyra

Tribe: Centotheceae. Megastachya

Tribe: Ehrharteae . Ehrharta, Microlaena
Supertribe: Bambusodae

Tribe: Bambuseae. Thamnocalamus , Oxytenanthera,
Bambusa

Subfamily: ARUNDINOIDEAE

Tribe: Stipeae. Stipa, Nassella

Tribe: Arundineae . Phragmites, Arundo

Tribe: Danthonieae . Prionanthium, Tribolium,
Styppeiochloa, Pentaschistis, Schismus,
Karroochloa, Chaetobromus, Dregeochloa,
Centropodia, Pentameris, Merxmuellera,
Pseudopentameris, Elytrophorus, Urochlaena,
Cortaderia

Tribe: Aristideae. Aristida, Sartidia, Stipagrostis

Subfamily: CHLORIDOIDEAE

Tribe: Pappophoreae . Enneapogon, Kaokochloa,
Schmidtia

Tribe: Chlorideae ( including Cynodonteae,

Eragrosteae, Sporoboleae , Leptureae, Trageae,
Spartineae). Tetrachne, Fingerhuthia,
Entoplocamia, Eragrostis, Diandrochloa,
Cladoraphis, Triraphis, Stiburus, Sporobolus,
Eleusine, Pogonarthria, Leptochloa, Diplachne,
Odyssea, Brachychloa, Tetrapogon, Acrachne,
Dactyloctenium, Enteropogon, Schoenefeldia,
Leptocarydion, Lintonia, Chloris, Brachyachne,
Dinebra, Coelachyrum, Harpochloa, Bewsia,
Trichoneura, Lophachme, Craspedorhachis,
Willkommia, Polevansia, Ctenium, Microchloa,
Eustachys, Cynodon, Rendlia, Spartina, Lepturus,
Catalepis, Tragus, Monelytrum, Perotis, Mosdenia,
Tripogon, O rope ti urn

Subfamily: PANICOIDEAE
Supertribe: Panicodae

Tribe: Paniceae . Pseudechinolaena, Tricholaena,
Melinis, Entolasia, Acroceras, Oplismenus,
Sacciolepis, Panicum, Paspalum, Oryzidium,
Leucophrys, Brachiaria, Urochloa, Eriochloa,
Digitaria, Tarigidia, Stereochlaena, Alloteropsis,
Megaloprotachne, Axonopus, Echinochloa,
Paspalidium, Stenotaphrum, Paratheria, Setaria,
Pennisetum, Cenchrus, Anthephora, Odontelytrum

Tribe: Arundinelleae . Arundinella, Danthoniopsis,
Loudetia, Tristachya, Trichopteryx

30

Supertribe: Andropogonodae
Tribe: Andropogoneae

Subtribe: Andropogoninae. Miscanthus, Imperata,
Eulalia, Eriochrysis, Microstegium, Sorghum,
Sorghastrum, Chrysopogon, Vetiveria,
Cleistachne, Trachypogon, Heteropogon,
Arthraxon, Dichanthium, Bothriochloa,
Andropogon, Schizachyrium, Cymhopogon,
Diheteropogon, Monocymhium, Themeda,
Hyperthelia, Elymandra, Hyparrhenia,
lschaemum, Sehima, Thelepogon

Subtribe: Rottboelliinae. Phacelurus, Vossia,

Rhytachne, Urelytrum, Hemarthria, Elionurus,
Rotthoelha, Coelorhachis, Hackelochloa,
Oxyrhachis

Tribe: Maydeae. Coix

31

GENERA AND SPECIES

Acrachne Wright & Arn. ex Chiov.

Arthrochloa Lorch, Camusia Lorch, Normanboria
Butzin.

Annual ; caespitose. Culms 120-800 mm high; herba-
ceous. Leaf blades linear (broadly, tapered to a hairlike tip);
flat. Ligule a fringed membrane to a fringe of hairs.

Inflorescence of spike-like main branches', digitate or
subdigitate (usually with the lower spikes scattered, but
becoming subdigitate above), or non-digitate (A.
racemosa)\ espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; 5.5-13 mm long; com-
pressed laterally; disarticulating above the glumes, or
falling with the glumes, or not disarticulating (the lemmas
falling acropetally from the rachilla, but the spikelet often
falling wholly or in part before all the lemmas have been
shed); not disarticulating between the florets, or disarticu-
lating between the florets (the rachilla tough or breaking
irregularly, the paleas persistent). Glumes two; relatively
large; more or less equal', markedly shorter than the
spikelets; awnless (but subulate via an excurrent mid-
nerve); similar (thinly cartilaginous). Incomplete florets
distal to the female-fertile florets; proximal incomplete
florets absent.

Female-fertile florets 8-20. Lemmas similar in texture
to the glumes to decidedly firmer than the glumes
(cartilaginous); 3 nerved (the laterals closer to the margins
than to the mid-nerve, and excurrent as small teeth);
incised; mucronate (from the midnerve). Palea present
(lanceolate). Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small (0.8-1. 1 mm long); ellipsoid;
hilum short; pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR cell chloroplasts
centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 3 species. Abyssinia, southern
Africa, Indochina, Indomalayan region, Australia. Meso-
phytic; in shade and in open habitats (sandy savanna);
glycophytic. Namibia, Botswana, Transvaal, and
Swaziland. 1 indigenous species.

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Acrachne racemosa (Roem. & Schult.) Ohwi

Fig. 10. PI. 1.

(=A. verticillata (Roxb.)

Chiov.) 2.

Annual; tufted (culms erect or
geniculately ascending); 120-800
mm tall. Leaf blades 120-200
mm long; 8-15 mm wide. Spike-
lets 6-9 mm long. Inflorescence
with 5-10 spike-like racemes in
1-3 whorls; spikelets 10-1 5-flowered; glumes and lemmas
extending into awns 1/3-2/3 their length.

Flowering January to April. Sandy soil in moist and
shady places. Locally common. Biome: Savanna and Nama-
Karoo. Old world tropics. West Indies.

Description: Chippindall & Crook 1976 (41), Chippin-
dall 1955 (132), Clayton et al. 1970-1982 (258).
Illustration: Chippindall 1955 (fig. 105), Clayton et al.
1970-1982 (fig. 71 ). Voucher: Pienaar 253. PRECIS code
9903311-00100.

Fig. 10. Acrachne racemosa

32

Acroceras Stapf

Neohusnotia A. Camus.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or decumbent. Culms 100-1250 mm high; herba-
ceous (often much-branched). Leaf blades linear-lanceolate
to ovate-lanceolate; cordate (somewhat amplexicaul).
Ligule a fringed membrane ( very narrow ), or a fringe of
hairs. Plants with hermaphrodite florets. The spikelets all
alike in sexuality.

Inflorescence of spike-like main branches ( racemes or
panicles), or paniculate', open; espatheate. Spikelet-bearing
axes persistent.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations (in lower parts of panicle), or not in distinct ‘long-
and-short’ combinations; abaxial; compressed laterally to
not noticeably compressed ( terete below)', falling with the
glumes. Glumes two; very unequal; awnless; similar
(membranous). Proximal incomplete florets 1\ paleate,
palea fully developed; male, or sterile.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes; smooth to striate; becoming indurated, or not
becoming indurated; hairless (shiny); having the margins
tucked in onto the palea; with a clear germination flap; 5
nerved; entire; crested at the tip', awnless (the apex blunt,
hard, laterally compressed). Palea present (the tip reflexed);

Fig. 1 1 . Acroceras macrum

relatively long. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Flilum long-linear (half to two thirds the
fruit length).

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. 15
species. Africa, Madagascar, Indomalayan region.
Hydrophytic to mesophytic; in shade and in open habitats
(shallow water, damp places and forests); glycophytic.
Namibia, Botswana, Transvaal, Natal, and Cape Province.
1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

Acroceras macrum Stapf

Fig. 11. PI. 2.

Nile grass.

Perennial; rhizomatous and
tufted; 400-1100 mm tall. Leaf
blades to 120 mm long; 10 mm
wide. Spikelets 4— 5 mm long; 1.5
mm wide. Rhizome creeping
extensively; ligule a very short
rim of hairs; spikelets with con-
spicuous, indurated, rounded appendages at the laterally
compressed apex of the glumes and lemmas; lower glume
2/3 the length of the spikelet; upper glume equalling the
length of the spikelet.

Flowering November to July. Grows in flooded areas
near rivers, swamps or vleis. Infrequent. Biome: Savanna.
Tropical Africa. Cultivated pasture.

Description: Chippindall 1955 (386). Illustration: Chip-
pindall 1955 (fig. 329). Voucher: Smith 413. PRECIS code
9901121-00100.

Agrostis L.

Agraulus P. Beauv., Agrestis Bub., Anomalotis Steud.,
Bromidium Nees, Candollea Steud., Chaetotropis Kunth,
Decandolea Batard, Didymochaeta Steud., Lachnagrostis
Trim, Neoschischkinia Tsvelev, Notonema Raf.,
Pentatherum Nabelek, Podagrostis (Griseb.) Scribn.,
Senisetum Koidz., Trichodium Michaux, Vilfa Adans.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 30-1500 mm
high; herbaceous; unbranched above. Leaf blades linear;
usually flat, or rolled (convolute, or canaliculate). Ligule an
unfringed membrane.

Inflorescence paniculate', open, or contracted (e.g.,
Bromidium)', espatheate. Spikelet-bearing axes persistent.

Spikelets 0.8-4 mm long', compressed laterally; disartic-
ulating above the glumes. Callus hairs absent, or if present
less than 0.5 mm long. Glumes two; more or less equal;
nearly always about equalling the spikelets to much
exceeding the spikelets (very rarely shorter); awnless;
similar (usually narrow, membranous). All florets female-
fertile', proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes (thinly membranous to hyaline ); 3-5 nerved; entire
to incised (usually truncate or emarginate, sometimes
toothed via excurrent veins); awnless, or mucronate, or
awned. Awns when present 1, or 3 ( Bromidium ), or 5
(rarely); median, or median and lateral (by extension of the
lateral veins). The median awn different in form from the
laterals (when laterals present); dorsal; geniculate; much
shorter than the body of the lemma to about as long as the
body of the lemma, or rarely much longer than the body of
the lemma. Palea nearly always present; relatively long, or
conspicuous but relatively short, or very reduced. Lodicules
2; membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; embryo small.

33

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. About 220
species. Temperate. Helophytic, or mesophytic, or xero-
phytic (rarely); in shade and in open habitats (grassland,
light woodland); maritime-arenicolous (rarely), or
glycophytic. Namibia, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. Indigenous
species (9), naturalized species (2-3).

Intergeneric hybrids with Polypogon (X Agropogon P.
Fourn.), Calamagrostis.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA. 3. Smook & Stirton. 1979. Bothalia 12: 637.

Species treatment by G.E. Gibbs Russell.

1(0). Glumes 3.5-5 mm long 2

Glumes to 3.5 mm long 7

2(1). Glumes with middle nerve extending into a short awn

A. polypogonoides

Glumes acute, without an awn 3

3(2). Panicle narrow and spikelike, the spikelets closely
imbricate on appressed branches . . A. continuata
Panicle spreading, spikelets not closely imbricate . 4

4(3). Plant annual, lemmas 4-nerved A. avenacea

Plant perennial, lemmas 5-nerved 5

5(4). Leaf blades 2-6 mm across; lemmas awned from near
the base; rachillas produced as a hairy bristle;
panicle branches flexuous, spreading at maturity

A. barbuligera var. barbuligera

Leaf blades 1-2 mm across; lemmas awned from
about the middle; rachillas not produced; panicle

branches rigid, straight, ascending 6

6(5). Leaf blades folded A. eriantha var. eriantha

Leaf blades flat A. eriantha var. planifolia

7(1). Pedicels more than 5 mm long, spikelets widely

separated; panicles very diffuse 8

Pedicels less than 5 mm long, the spikelets close
together; panicles not diffuse open or compact .

10

8(7). Pedicels 20 mm long or more; paleas absent

A. montevidensis

Pedicels 5-10 mm long; paleas present 9

9(8). Glumes 1.5-2. 5 mm long

A. bergiana var. bergiana

Glumes 2. 5-3.0 mm long

A. bergiana var. laevisulca

10(7). Panicles 150-400 mm long, narrow and sinuous, the

branches held nearly erect 11

Panicles to 200 mm long, open or compact, the

branches ascending to spreading 12

11(10). Lemmas hairy . . A. lachnantha var. lachnantha
Lemmas glabrous ... A. lachnantha var. glabra
12(10). Lemmas glabrous; rachillas not produced .... 13

Lemmas hairy; rachillas produced 14

13(12). Plants 400-750 mm tall, perennial, often
stoloniferous; leaf blades flat, 3-5 mm wide;
glumes greenish or purple-tinged . . A. gigantea
Plants 50-300 mm tall, delicate, annual or weakly
perennial; leaf blades folded, 0. 5-1.0 mm wide;
glumes usually dark purple .... A. subulifolia
14(12). Rachillas 1/2-3/4 length of floret; glumes tinged
with purple; southwestern Cape . A. schlechteri
Rachillas less than 1/2 length of floret; glumes light
green; plants not restricted to southwestern Cape

15

15(14). Plant an annual weed; lemma hairs not spreading,
usually not longer than lemmas; lemmas 4-

nerved A. avenacea

Plant an indigenous perennial; lemma hairs
spreading, longer than lemmas; lemmas 5-nerved
A. barbuligera var. longipilosa

Fig. 12. Agrostis eriantha var. eriantha

34

Agrostis avenacea Gmel.

Bent grass, blown grass.

Annual; 180-600 mm tall.

Leaf blades to 170 mm long;
about 2 mm wide. Spikelets
2. 5^1.0 mm long. Inflorescence
open, 80-190 mm long; rachilla
less than 1/2 the length of the flo-
ret; lemmas hairy, 4-nerved.

Flowering July to March. Disturbed areas or wet places.
Infrequent. Naturalized from Australia. Biome: Fynbos and
Savanna. Weed (of cultivation).

Description: Smook & Stirton 1979 (637). Voucher: Van
der Walt 398. PRECIS code 9902430-00050.

Agrostis barbuligera Stapf var. barbuligera

Perennial; tufted; 200-800
mm tall. Leaf blades to 250 mm
long; 2-6 mm wide (flat). Spike-
lets 4. 0-5. 5 mm long. The basal
sheaths splitting into fibres; pani-
cle branches flexuous, spreading
at maturity; rachilla produced as
a hairy bristle; lemmas awned
from near base.

Flowering November to March. Mountain grassland. In-
frequent, or locally common. Biome: Grassland. Endemic.

Description: Stapf 1898-1900 (548), Chippindall 1955
(99). Voucher: Acocks 21079. PRECIS code

9902430-00100.

Agrostis barbuligera Stapf var. longipilosa Goossens &

Papendorf

Spikelets 3. 0-3. 5 mm long.

Differs from the typical variety in
its smaller spikelets, with more
hairy lemmas.

Biome: Grassland. Endemic.

Voucher: Van der Schijff

4776. PRECIS code 9902430-
00200.

Agrostis bergiana Trin. var. bergiana

Delicate, weak perennial, or
annual; 150-300(-600) mm tall.

Leaf blades to 90 mm long; 1-2
mm wide. Spikelets 1.5-2. 5 mm
long. Panicle very diffuse, the
branches hairlike; pedicels 5-10
mm long; paleas present.

Flowering November to Feb-
ruary. Mountain grassland in
sheltered or wet places. Locally common. Biome: Fynbos
and Grassland. Endemic.

Description: Stapf 1898-1900 (547), Chippindall 1955
(101). Voucher: Huntley 422. PRECIS code

9902430-00300.

Agrostis bergiana Trin. var. laevisulca Stapf

Agrostis continuata Stapf

(=A. natalensis Stapf) 2.

Coarse perennial; tufted; 600-
900 mm tall. Leaf blades to 250
mm long; to 6(— 8) mm wide.

Spikelets about 5 mm long. In-
florescence dense, narrow and
spikelike, the spikelets overlap-
ping.

Flowering December to April. Vlei grassland and wet
places, sometimes at high altitudes. Biome: Savanna and
Grassland. North to Tanzania. The spikelike panicle
resembles Phalaris arundinacea, which has no awns, and
Koeleria capensis, which has 2-4-flowered spikelets.

Description: Stapf 1898-1900 (548), Chippindall 1955
(99), Clayton et al. 1970-1982 (111). Illustration: Chippin-
dall 1955 (fig. 70). Voucher: Pole-Evans 1968. PRECIS
code 9902430-00450.

Agrostis eriantha Hack. var. eriantha

Fig. 12. PI. 3.

Perennial; rhizomatous and
tufted; to 700 mm tall. Leaf
blades to 180 mm long; 1-2 mm
wide (folded). Spikelets 3. 5-5.0
mm long. Panicle branches rigid,
straight, held ascending at maturi-
ty; rachillas not produced; lem-
mas awned from the middle.

Flowering January to April.

Wet places, sometimes in disturbed areas or cultivation. In-
frequent. Biome: Savanna and Grassland. ?Endemic.
Includes var .planifolia, which possibly has flat leaf blades
and slightly longer callus hairs.

Description: Chippindall 1955 (99). Illustration: Chip-
pindall 1955 (fig. 71). Voucher: Potter 1745. PRECIS code
9902430-00500.

Agrostis eriantha Hack. var. planifolia Goossens &

Papendorf

Doubtfully separate from the
typical variety, but with flat leaf
blades and somewhat longer
callus hairs.

Biome: Savanna. ?Endemic.

PRECIS code 9902430-00600.

Agrostis gigantea Roth

Perennial; usually stolonifer-
ous (culms decumbent); 400-750
mm tall. Leaf blades to 90 mm
long; 3-5 mm wide. Spikelets
1.5-2. 5 mm long. Panicle open,

70-200 mm long, the branches
ascending; rachilla not produced;
lemmas glabrous.

Wet disturbed places. Infre-
quent. ? Naturalized from Europe. Biome: Fynbos and Sa-
vanna. This name is applied to our specimens by matching
at Kew; probably related to A. schimperiana of east Africa.

Voucher: Burtt Davy 9233. PRECIS code

9902430-00650.

Spikelets 2. 5-3.0 mm long.

Similar to the typical variety
except for the longer spikelets.

Flowering October to Feb-
ruary. Wet places in mountain
grassland. Rare. Biome: Fynbos,

Grassland and Afromontane.

Endemic.

Description: Stapf 1898-1900
(547), Chippindall 1955 (102). Voucher: De Winter & Codd
209. PRECIS code 9902430-00400.

Agrostis lachnantha Nees var. lachnantha

(=A. huttoniae (Hack.) C.E.
Hubb.) 2; (=A. lachnantha Nees
var. glabra Goossens &
Papendorf).

South African bent grass,
vinkagrostis.

Shortlived perennial, or an-

35

nual (usually robust); loosely tufted; 300-900 mm tall. Leaf
blades 70-200 mm long; 2—4 mm wide. Spikelets
1 .5— 2.5(— 3.0) mm long. Panicle 150—400 mm long, narrow
and sinuous, the branches held nearly erect.

Flowering October to March (occasionally earlier or
later). Riverbanks and wet places. Locally common. Biome:
Fynbos, Savanna, Grassland, Nama-Karoo, and Succulent
Karoo. Northwards through east Africa to Sudan and
Ethiopia. One of the most widespread of all our grass
species. Var. glabra is not considered distinct.

Description: Chippindall & Crook 1976, Stapf

1898-1900 (549), Chippindall 1955 (101), Clayton et al.
1970-1982 (106). Illustration: Chippindall 1955 (fig. 73).
Voucher: Oakes & Scheepers 301. PRECIS code
9902430-00900.

Agrostis montevidensis Spreng. ex Nees
Fog grass.

Annual; 200-600 mm tall.

Leaf blades to 130 mm long; 1-2
mm wide. Spikelets 1.5-2. 5 mm
long. Panicle very diffuse, the
branches hairlike; pedicels 20
mm long or more; paleas absent.

Flowering November to April.

Moist and disturbed places in mountain grassland. Rare.
Naturalized from South America. Biome: Fynbos and
Grassland. Weed.

Description: Smook & Stirton 1979 (637). Voucher:
Story 5438. PRECIS code 9902430-01050.

Agrostis polypogonoides Stapf

Perennial; tufted; to 750 mm
tall. Leaf blades to 1 20 mm long;

3-4 mm wide. Spikelets 4.0— 4.5
mm long. Glumes with an awn
1-3 mm long.

Flowering October to January.

Wet places. Rare. Biome: Fynbos
and Succulent Karoo. Endemic.

This is our only Agrostis species
with awned glumes.

Description: Stapf 1898-1900 (549), Chippindall 1955
(98). Voucher: Acocks 17581. PRECIS code

9902430-01200.

Agrostis schlechteri Rendle

Annual; 30-650 mm tall. Leaf
blades to 100 mm long; 2. 0-2. 5
mm wide. Spikelets 3.0-3. 5 mm
long. Panicle open, 90-150 mm
long, the branches ascending;
rachilla 1/2-3/4 the length of the
floret; lemmas glabrous.

Flowering January to April.

Wet places in mountains. Rare.

Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (762), Chippindall 1955
(101). Voucher: Esterhuysen 27690. PRECIS code
9902430-01300.

Agrostis subulifolia Stapf

Delicate perennial, or annual
(possibly); hydrophyte and tuft-
ed; 50-300 mm tall. Leaf blades
to 70 mm long; 0. 5-1.0 mm wide
(folded). Spikelets 2-3 mm long.

Panicle contracted to open, 15-80
mm long, the branches ascending;
rachilla not produced; lemmas
glabrous.

Flowering January to March. Mountain bogs. Infre-
quent. Biome: Grassland and Afromontane. Endemic.

Description: Chippindall 1955 (102). Voucher: Coetzee
574. PRECIS code 9902430-01400.

Aira L.

Airella (Dumort.) Dumort., Aspris Adans. ,Caryophyllea
Opiz, Fiorinia Park, Fussia Schur, Salmasia Bub.

Annual ; caespitose (small, slender). Culms 20—400 mm
high; herbaceous; unbranched above. Leaf blades linear;
setaceous ; flat, or folded, or rolled. Ligule an unfringed
membrane .

Fig. 13. Aira cupaniana

36

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 1.6-3. 5 mm long; compressed laterally; disar-
ticulating above the glumes. Rachilla hairless. Glumes two;
more or less equal; about equalling the spikelets-, awnless;
similar (membranous, delicate). All florets female-fertile;
proximal incomplete florets absent.

Female-fertile florets 2. Lemmas decidedly firmer than
the glumes (becoming papery)-. 5 nerved; entire, or incised;
awnless, or awned. Awns when present 1; dorsal; genicu-
late; much shorter than the body of the lemma, or about as
long as the body of the lemma, or much longer than the
body of the lemma. Palea present; relatively long. Lodicules
2; membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; fusiform; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 8 species.
North and South temperate. Mesophytic to xerophytic; in
open habitats (sandy soils). Orange Free State, Natal,
Lesotho, and Cape Province. 1 naturalized species.

References. 1 . Clayton. 1972. FWTA. 2. Tutin. 1980. FI.
Europ.

Species treatment by T.M. Sokutu.

Aira cupaniana Guss.

Fig. 13. PI. 4.

Annual; tufted; 3CMI00 mm
tall. Leaf blades 10-95 mm long;
to 3 mm wide. Spikelets 2-3 mm
long. Inflorescense open, spread-
ing; spikelets 2-flowered; glumes
pear-shaped; lemmas awned from
the lower third, lower lemma
sometimes awnless.

Flowering September to Jan-
uary. Shallow soils in damp to wet areas. Common. Natural-
ized from Europe. Biome: Fynbos and Grassland. Europe.
Weed. A variable species. See Clayton (1970) for a
comment on the intermediate state of the African material.
In the past our specimens were assigned to two species, A.
cupaniana and A. caryophyllea L. They are treated here
under A. cupaniana pending further research and
confirmation of their identities. The species can be
confused with Periballia minuta , last collected at
Simonstown in 1943, but this genus has an elongated
internode between the florets.

Description: Tutin 1980 (5: 227), Stapf 1898-1900
(463), Chippindall 1955 (86), Clayton et al. 1970-1982
(84). Illustration: Chippindall 1955 (fig. 57). Voucher:
Davidse 33862. PRECIS code 9901850-00100.

Alloteropsis Presl

Bluffia Nees, Coridochloa Nees, Holosetum Steud..
Mezochloa Butzin, Pterochlaena Chiov.

Annual (rarely), or perennial; caespitose, or decumbent.
Culms 200-1500 mm high; herbaceous; unbranched above.
Leaf blades linear to lanceolate. Ligule a fringed membrane
to a fringe of hairs.

Inflorescence of spike-like main branches', digitate or
subdigitate, or non-digitate (in whorls on a short central
axis); espatheate. Spikelet-bearing axes persistent.

Spikelets in triplets, or in pairs; consistently in 7 ong-
and-short' combinations . Spikelets 2.5-7 mm long \abaxial,
compressed dorsiventrally; falling with the glumes. Glumes
two; very unequal; awned, or awnless; very dissimilar (G1
smaller, thinner, often mucronulate; G2 densely ciliate).
Lower glume 3—5 nerved. Proximal incomplete floiets 1,
paleate, palea reduced (deeply bifid, 1 -nerved); male.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes ; smooth to striate; not becoming indurated; hairless;
having the margins lying flat and exposed on the palea; with
a clear germination flap; 5 nerved (1-3 in ~Mezochloa')\
entire; mucronate to awned. Awns when present 1; apical;
non-geniculate; much shorter than the body of the lemma
to about as long as the body of the lemma. Palea present
(auriculate at base); relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit small; hilum

37

Photosynthetic pathway. C4 (in all the material
examined except A. semialata subspecies eckloniana ,
including ‘ Coridochloa ’), or C3 (A. semialata subspecies
eckloniana). The anatomical organization when C4 uncon-
ventional. Organization of PCR tissue when C4 Alloteropsis
type (with an inner PCR sheath, and an outer sparsely
chlorenchymatous sheath of unknown function).
Biochemical type PCK (in Australian C4 A. semialata), or
NADP-ME (in southern African C4 A. semialata : evidently
more biochemical typing needed, in relation to the
intergrading C4 anatomical forms and the problematical
taxonomy); XyMS+ (usually), or XyMS- (in C4 forms of
A. semialata). PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x - 9. Panicoideae; Panicodae; Paniceae. 5-8
species (with complexes around A. semialata and A.
paniculata reflecting specific and generic synomyms).
Tropical Africa, Asia & Australia. Helophytic, or meso-
phytic, or xerophytic; in open habitats (marshy and weedy
places); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, and Cape Province.
3 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA. 2.
Gibbs Russell. 1983. Bothalia 14: 205.

Species treatment by G.E. Gibbs Russell.

1(0). Spikelets 5-8 mm long; palea of upper floret glabrous
or with sparse hairs; leaf blades linear to lanceolate,
bases tapering, margins not ciliate; culms thickly

clad by old leaf sheaths at base 2

Spikelets 3. 0^1.5 mm long; palea of upper floret
papillose; leaf blades lanceolate, bases cordate to
auriculate, margins ciliate; culm bases not thickly

clad by old leaf sheaths 3

2(1). Leaf blades rolled or curved upwards, tapering
gradually to tip, sparsely hairy, linear, hard-
textured; old leaf sheaths at base of culms with
veins forming ribs 0. 5-1.1 mm wide; racemes often
(but not always) longer than 80 mm, usually with

light-coloured spikelets loosely arranged

A. semialata subsp. semialata

Leaf blades held flat, tapering abruptly to tip, densely
hairy, lanceolate, rather soft-textured; old leaf
sheaths at base of culms with veins to 0.3 mm wide;
racemes often (but not always) shorter than 80
mm, usually with dark-coloured spikelets tightly
packed together . A. semialata subsp. eckloniana
3(1). Plant annual; culms often geniculate at base; northern

Namibia and Botswana A. cimicina

Plant perennial; culms erect from a knotted base;
eastern Transvaal and Natal A. papillosa

Alloteropsis cimicina (L.) Stapf

Annual; tufted (culms erect or
ascending); 300-1000 mm tall.

Leaf blades 30-150 mm long
(bases rounded, margins ciliate);

5-10 mm wide. Spikelets 3. 5-5. 5
mm long. Palea with globular
hairs.

Flowering December to May.

Moist open places on sandy clay
soil. Rare (in southern Africa). Biome: Savanna. Old world
tropics.

Description: Clayton et al. 1970-1982 (615).

Illustration: Clayton et al. 1970-1982 (fig. 144). Voucher:
Killick & Leistner 3027. PRECIS code 9900940-00100.

Alloteropsis papillosa Clayton

Perennial; tufted; 400-700
mm tall. Leaf blades 40-250 mm
long (bases rounded to auriculate,
margins ciliate); 2-8 mm wide.

Spikelets 3. 0-4. 5 mm long. Culm
bases knotted, with a few hairy
old leaf sheaths; palea with
globular hairs.

Flowering November to May.

Sandy soil in open or shaded habitats. Biome: Savanna. To
tropical east Africa. Combines the perennial habit and
longer leaves of A. semialata with the ciliate blade margins
and papillate palea of A. cimicina. Possibly of hybrid origin.

Description: Clayton et al. 1970-1982 (615). Voucher:
Ward 4140. PRECIS code 9900940-00150.

Alloteropsis semialata (R. Br.) Hitchc. subsp.

eckloniana (Nees) Gibbs Russell

Fig. 14. PI. 5. PI. 6.

( =A . semialata auctt., non
Gibbs Russell) 2; ( -A . semialata
(R. Br.) Hitchc. var. ecklonii
(Stapf) Stapf) 2.

Perennial; short-rhizomatous
and tufted; 250-1100 mm tall.

Leaf blades 3-12 mm wide (flat,
base tapering, tip tapering
abruptly, velvety to sparsely hairy, margins not ciliate).
Spikelets 5-8 mm long. Basal parts V-shaped in silhouette,
not bulbous; basal sheaths with veins to 0.3 mm wide;
racemes often shorter than 80 mm, with dark-coloured
spikelets tightly packed; palea glabrous or with sparse hairs.

Flowering September to March (sometimes in other
months). Grassland, rocky places and forest margins;
usually at higher altitudes and more acid soil than subsp.
semialata. Common. Biome: Savanna and Grassland. North
to Tanzania at high altitudes. Natural pasture. This
subspecies is unusual in the Paniceae because it has C3
photosynthesis.

Description: Gibbs Russell 1983 (205), Chippindall
1955 (423). Illustration: Chippindall 1955 (fig. 352).
Voucher: Smook 2586. PRECIS code 9900940-00200.

Alloteropsis semialata (R. Br.) Hitchc. subsp. semialata

( =A . semialata auctt., non
Gibbs Russell) 2.

Perennial; short-rhizomatous
and tufted; 300-1 300(— 1 500) mm
tall. Leaf blades 3— 5(— 6) mm
wide (usually curved inward or
loosely rolled, sparsely hairy,
base tapering, tip gradually
tapering, margins not ciliate). Basal parts bulbous, rounded
in silhouette; basal sheaths with ribs 0. 5-1.1 mm wide;
racemes often longer than 80 mm, with light-coloured
spikelets loosely arranged; palea glabrous or with sparse
hairs.

Flowering September to March. Grasslands and
bushveld. Common. Biome: Savanna and Grassland. Old
world tropics. Natural pasture. A. semialata is most unusual
in having two photosynthetic types in one species. This is
the subspecies with C4 photosynthesis.

Description: Chippindall 1955 (423), Clayton et al.
1970-1982 (616). Voucher: Van der Schijff 2035. PRECIS
code 9900940-00250.

38

Ammophila Host

Psamma P. Beauv.

Perennial; long-rhizomatous. Culms 200-1300 mm
high; herbaceous; unbranched above. Leaf blades linear;
rolled (convolute). Ligule an unfringed membrane.

Inflorescence paniculate', contracted', espatheate.
Spikelet-bearing axes persistent.

Spikelets 9-15 mm long', compressed laterally; disarticu-
lating above the glumes. Hairy callus present. Glumes two;
more or less equal; long relative to the adjacent lemmas
(longer); awnless; similar. All florets female-fertile',
proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes; 5 nerved; entire, or incised; mucronate ( with a
subterminal, vestigial awn). Palea present; relatively long.
Lodicules 2; membranous; ciliate, or glabrous. Stamens 3.
Ovary glabrous. Fruit medium sized; hilum long-linear (two
thirds of fruit length); embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 2 species.
North temperate. Commonly adventive. Xerophytic; in
open habitats; maritime-arenicolous. Cape Province. 1
naturalized species.

A. arenaria hybridizes with Calamagrostis epigejos ( X
Ammocalamagrostis P. Fourn., a useful sand stabilizer).

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Fig. 15. Ammophila arenaria

Ammophila arenaria (L.) Link

Fig. 15. PI. 7.

Marram grass.

Robustperennial; rhizomatous
and tufted (the culms creeping
through blown sand); 600-1300
mm tall. Leaf blades rolled and
appearing setaceous, to 750 mm
long. Panicle very narrow and
spikelike.

Flowering October to December (old inflorescences
persistent until autumn). Seaside dunes. Naturalized from
Europe. Widely naturalized. Erosion control (in seaside
dunes).

Description: Chippindall 1955 (93). Illustration: Chip-
pindall 1955 (fig. 65). Voucher: Liebenberg 4024. PRECIS
code 9902560-00100.

Andropogon L.

Anatherum P. Beauv., Arthrostachys Desv.,
Arthrolophis (Trin.) Chiov., Diectomis Kunth,
Dimeiostemon Raf., Eriopodium Hochst., Heterochloa
Desv., Homoeatherum Nees , Leptopogon Roberty.

Annual, or perennial; long-rhizomatous, or caespitose,
or decumbent. Culms 80-2500(-4300) mm high; herba-
ceous; branched above, or unbranched above. The shoots
not aromatic. Leaf blades linear. Ligule an unfringed
membrane to a fringed membrane . Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant', overtly heteromorphic.

Inflorescence of spike-like main branches, or paniculate
(usually with spikelets in paired or digitate ‘racemes' ,
which are often spatheate and aggregated into false
panicles); spatheate (usually); a complex of ‘partial inflo-
rescences’ and intervening foliar organs (often), or not so.
Spikelet-bearing axes ‘racemes’; paired (rarely one or
several, not deflexed); with very slender rachides, or with
substantial rachides: disarticulating at the joints. ‘Articles’
without a basal callus-knob.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, or barren, usually awnless,
occasionally suppressed (sometimes reduced to their
pedicels). Female-fertile spikelets compressed laterally, or
not noticeably compressed, or compressed dorsiventrally;
falling with the glumes. Callus blunt. Glumes two; more or
less equal; awned, or awnless (upper sometimes aristate);
very dissimilar (subcoriaceous to membranous, the lower
flat, concave or canaliculate on the back, the margins folded
and 2-keeled; the upper naviculate, carinate above). Lower
glume two-keeled. Proximal incomplete florets I ; epaleate;
sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline to firm, sometimes substipitate beneath the
awn); incised (usually bifid); awned. Awns 1 ; median; from
the sinus; geniculate; about as long as the body of the
lemma to much longer than the body of the lemma. Palea
present; very reduced (hyaline). Lodicules 2; fleshy; ciliate,
or glabrous. Stamens 1-3. Ovary glabrous. Fruit small;
hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, jc = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. About 100 species.
Tropical. Mesophytic, or xerophytic: mostly savanna, some
in tropical highlands. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 15 indigenous species.

References. 1. Anderson 1960. Bothalia 7: 417. 2.
Clayton. 1964. Kew Bull. 17: 470. 3. Clayton & Renvoize.
1982. FTEA.

Species treatment by G.E. Gibbs Russell.

39

1(0). Plants annual, reddish, with numerous flowering
branches; racemes 1 per spathe; lower glume of
pedicellate spikelets large, flat, papery, reddish,

with an awn 5-7 mm long A. fastigiatus

Plants perennial, not reddish; flowering branches 1-8
per culm; racemes more than 1 per spathe; lower
glume of pedicellate spikelets not as above .... 2

2(1). Glumes with irregular pits between the veins

A. lacunosus

Glumes lacking pits between the veins 3

3(2). Inflorescence white silky plumose; hairs at least as
long as sessile spikelets; pedicellate spikelets

usually conspicuously reduced or lacking 4

Inflorescence not white silky plumose; hairs if present
not longer than sessile spikelets; pedicellate

spikelets not conspicously reduced 6

4(3). Sessile spikelets 2-3 mm long; pedicellate spikelets
lacking; inflorescence hairs at least twice as long

as spikelets A. eucomus

Sessile spikelets 4. 5-6.0 mm long; pedicellate

spikelets reduced; inflorescence hairs as long as

sessile spikelets 5

5(4). Racemes 4-10 per spathe; pedicellate spikelets well-
developed or variously reduced; plants 800-1800

mm tall A. huillensis

Racemes 2-3 per spathe; pedicellate spikelets reduced

or suppressed; plants 300-900 mm tall

A. laxatus

6(3). Lower glume of sessile spikelets with hairs longer

than 1 mm on the back A. amethystinus

Lower glume of sessile spikelets glabrous or
sometimes hispid on the back, rarely with a few

hairs to 1 mm long 7

7(6). Basal sheaths compressed and keeled, plant bases
often flattened; inflorescence rachises and pedicels
linear and rounded, not or only slightly broadened
at the upper end (or narrowly clavate in A.

appendiculatus) 8

Basal sheaths not keeled and plant bases not flattened;
inflorescence rachises often flattened, broadening
upwards so that widest part is just below the

spikelet 11

8(7). Sessile spikelets 0.5-0. 8 mm wide; inflorescences
hairless or nearly so, sometimes with a few hairs

at upper ends of pedicels 9

Sessile spikelets (0.7-)0. 9-1.1 mm wide;
inflorescences hairy on callus, rachis and pedicels

10

9(8). Racemes 1-2 per spathe, short and straight, awnless;
spikelets 4. 0^1. 5 mm long, glumes glabrous; Natal

A. festuciformis

Racemes 4-15 per spathe, long and flexuous, short -
awned; spikelets 5. 5-7.0 mm long, lower glume of
sessile spikelets with appressed stiff hairs on keels;
Namibia and Botswana A. brazzae

10(8). Racemes 2-3 per spathe, 25-60 mm long; callus of
sessile spikelets inserted in a tuft of hairs;
Transvaal mountain grasslands .... A. mannii
Racemes 2-20 per spathe, ( 30— )60— 1 50 mm long;
callus of sessile spikelets (sometimes hairy)
inserted in a membranous-edged socket;

widespread distribution, various habitats

A. appendiculatus

1 1(7). Lower glume of sessile spikelets flat or shallowly
furrowed, keels lateral, backs sometimes with

short hairs or glabrous 12

Lower glume of sessile spikelets deeply grooved,
keels dorsal and sometimes nearly meeting,

backs glabrous 13

1 2( 1 1 ). Lower glume of sessile spikelets wingless, back flat
with a narrow median furrow, tip not toothed, not
awned; bushveld . A. gayanus var. polycladus
Lower glume of sessile spikelets winged at edge in
upper end, back flat, tip 2-toothed, often awned

between teeth; mountain grassland

A. distachyos

13(1 1 ). Upper glume of sessile spikelets with awn 7-10 mm
long, and both glumes of pedicellate spikelets

with awns 4-7 mm long A. chinensis

All glumes awnless or lower glume of pedicellate
spikelets with a minute awn to 2 mm long . . 14
14(13). Inflorescence of 2-20 racemes, dark purple-tinged;

lemma awn 10-15 mm long; damp or shady

habitats A. appendiculatus

Inflorescence of 2(— 5) racemes, glaucous green or
reddish; lemma awn 15-30 mm long; open

habitats on hills and mountains 15

15(14). Sessile spikelets 7-9 mm long, keels of lower
glume held separated, callus broadly obtuse;
basal parts and rhizomes knotted; inflorescences
glaucous grey or green; mountain sourveld . . .

A. ravus

Sessile spikelets 5-7 mm long, keels of lower
glume nearly meeting, callus obtuse to subacute;
basal parts and rhizomes straight; inflorescences

usually reddish-tinged; various habitats

A. schirensis

Fig. 16. Andropogon chinensis

40

Andropogon amethystinus Steud.

( =A . abyssinicus sensu
Chippind., non Fresen.) 3; (=A.
pilosellus Stapf) 3.

Perennial; rhizomatous and
tufted; 80-700 mm tall. Leaf
blades 10-150 mm long; 1-4 mm
wide. Spikelets (sessile) 5. 0-8. 5
mm long (pedicellate slightly
shorter). Racemes 2 per spathe, pedicels linear or slightly
clavate; lower glume of sessile spikelets flat on the back,
with hairs longer than 1 mm, but inflorescence not plumose.

Flowering February to June. Mountain grassland.
Locally common. Biome: Afromontane. Tropical Africa
and India. A. abyssinicus Fresen., a closely-related annual,
does not occur in southern Africa. Our specimens
previously included in this species are perennial and thus
belong in A. amethystinus.

Description: Chippindall 1955 (496), Clayton et al.
1970-1982 (772). Voucher: Edwards 2819. PRECIS code
9900710-00150.

Andropogon appendiculatus Nees

Blougras.

Perennial; densely tufted;

300-1300 mm tall. Leaf blades
1 50-500 mm long; to 6 mm wide.

Spikelets (sessile and pedicellate)

5-7 mm long; (0.7-)0.9-l .2 mm
wide. Basal sheaths keeled,
flattened, yellow, becoming
brown, shining; inflorescence of 1-2 flowering branches
per culm, racemes 4—20 per spathe, (30 — )60 — 1 50 mm long,
dark purple, with short hairs, pedicels rounded, slightly
clavate; callus of sessile spikelets inserted in a
membranous-edged socket, lower glume deeply but broadly
grooved; lemma awn 10—15 mm long.

Flowering October to April. Wet or shady places. Com-
mon. Biome: Fynbos, Savanna, and Grassland. Southern
Africa. A widespread variable species best distinguished by
the flattened basal parts and deeply but widely grooved
lower glumes of the sessile spikelets. Individuals with
particularly hairy racemes may be mistaken for A.
huillensis, which has 5-7 flowering branches per culm.

Description: Chippindall 1955 (500). Illustration: Chip-
pindall 1955 (fig. 403). Voucher: Huntley 456. PRECIS
code 9900710-00200.

Andropogon brazzae Franch.

Perennial; rhizomatous; to
2000 mm tall. Leaf blades to 600
mm long; to 5 mm wide. Spike-
lets (sessile and pedicellate)

5. 5-7.0 mm long; 0.6-0. 8 mm
wide. Basal sheaths keeled,
flattened; inflorescence nearly
glabrous, racemes 4—15 per
spathe, long, slender and flex-
uous, pedicels linear, rounded or slightly clavate; lower
glume of sessile spikelets shallowly concave.

Flowering February to May. Beside permanent rivers.
Rare and conservation status not known. Biome: Savanna.
To Angola and Zaire.

Voucher: Smith 2685. PRECIS code 9900710-00300.

Andropogon chinensis (Nees) Merr.

(=A. schinzii Hack.) 3.

Hairy bluegrass, tweevinger-
gras.

Fig. 16. PI. 8.

Plant glaucous grey.

Perennial; densely tufted;

600-1200 mm tall. Leaf blades
100^100 mm long; to 8 mm wide.

Spikelets (sessile) 5-7 mm long
(pedicellate somewhat shorter),
reddish tinged; base slightly bulbous; culms branched;
racemes 2-3 per spatheole, pedicels cuneate; lower glume
of sessile spikelets deeply and narrowly grooved, upper
glume awned, both glumes of pedicellate spikelets with an
awn 4-7 mm long.

Flowering December to June. Rocky hillsides and often
in poor sandy soil. Common. Biome: Savanna, Grassland,
and Nama-Karoo. Throughout tropical Africa and Asia to
China.

Description: Chippindall 1955 (499), Clayton et al.
1970-1982 (779). Illustration: Clayton et al. 1970-1982
(fig. 180,3). Voucher: De Winter & Leistner 5487. PRECIS
code 9900710-00350.

Andropogon distachyos L.

Mountain andropogon, twee-
vingergras.

Perennial; rhizomatous and
tufted; 300-1000 mm tall. Leaf
blades 70-200 mm long; 2-4 mm
wide. Spikelets (sessile) 9-1 1
mm long (pedicellate shorter).

Plant base with silky hairs;
racemes 2 per spathe, pedicels stout, slightly clavate; lower
glume of sessile spikelets flat on back, broadly winged on
upper 1/3, tip bidentate, often awned.

Flowering January to June. Mountain grassland. Infre-
quent. Biome: Afromontane. Throughout tropical Africa to
Asia.

Description: Chippindall 1955 (496), Clayton et al.
1970-1982 (770). Illustration: Clayton et al. 1970-1982
(fig. 180,1). Voucher: Edwards 2006. PRECIS code
9900710-00400.

Andropogon eucomus Nees

Snowflake grass, kapokgras,
old man’s beard, silver thread
grass.

Perennial; densely tufted;

200-900 mm tall. Leaf blades
40-200 mm long; to 4 mm wide.

Spikelets 2-3 mm long (all ses-
sile, accompanied by a hairy
empty pedicel). Inflorescence plumose, of 2-6 flowering
branches; racemes 2-5 per spathe, with white silky hairs
twice as long as the spikelets; lower glume of sessile
spikelets deeply and narrowly grooved.

Flowering November to May. Vleis and wet places.
Common. Biome: Fynbos, Savanna, and Grassland.
Tropical Africa and Madagascar. Closely related to A.
huillensis and A. laxatus , which both have larger spikelets.

Description: Chippindall 1955 (502), Clayton et al.
1970-1982 (775). Illustration: Chippindall 1955 (fig. 404).
Voucher: Louw 870. PRECIS code 9900710-00500.

41

Andropogon fastigiatus Swartz

(=Diectomis fasti giata
(Swartz) Kunth) 3.

Annual; tufted; 300-500 mm
tall. Leaf blades 50-300 mm
long; 1-4 mm wide. Spikelets
(sessile) 4-5 mm long (pedicel-
late longer and wider). Plants
reddish brown; inflorescences
with numerous flowering branches per culm; racemes
solitary in spathes; lower glume of sessile spikelet very
deeply and narrowly grooved, lower glume of pedicellate
spikelet large, flat, papery, reddish, with an awn 5-7 mm
long.

Flowering April to May. Dry sandy soil. Rare and con-
servation status not known. Locally common. Biome: Sa-
vanna. Throughout tropics. Monocymbium ceresiiforme
also has solitary spatheate racemes, but that is a perennial
species with wider leaf blades and shorter spikelets.

Description: Chippindall 1955 (504), Clayton et al.
1970-1982 (777). Illustration: Chippindall 1955 (fig. 405).
Voucher: Codd 4029. PRECIS code 9900710-00600.

Andropogon festuciformis Rendle

( =Hypogynium schlechteri
(Hack.) Pilg.) 2.

Perennial; densely tufted;

160-1000 mm tall. Leaf blades
50-300 mm long; 1. 5-3.0 mm
wide. Spikelets (sessile) 4. 0^1. 5
mm long (pedicellate larger);

0.5-0. 8 mm wide. Basal sheaths
keeled, flattened; inflorescence glabrous, racemes 1-2 per
spathe, awnless, short, pedicels linear, rounded; lower
glume of sessile spikelets flattish.

Flowering July to January. Moist places. Infrequent.
Southern tropical Africa.

Description: Chippindall 1955 (516). Illustration: Chip-
pindall 1955 (fig. 412). Voucher: Wood 8543. PRECIS
code 9900710-00700.

Andropogon gayanus Kunth var. polycladus (Hack.)

Clayton

(=A. gayanus Kunth var.
squamulatus (Hochst.) Stapf) 3.

Rhodesian bluegrass, Rhode-
sieseandropogonL

Robust perennial; tufted;
culms branched, 1200-3600
mm tall. Leaf blades to 600 mm long; 5-20 mm wide (in
the middle, base narrow). Spikelets (sessile) 6. 0-7. 5 mm
long (pedicellate a little shorter). Plant glaucous; racemes
2 per spathe, pedicels cuneate; lower glume of sessile
spikelet broad, flattish but with a narrow central furrow;
lower or both glumes of pedicellate spikelet with awns to
10 mm.

Flowering December to June. Bushveld. Common.
Biome: Savanna. Tropical Africa.

Description: Chippindall 1955 (499), Clayton et al.
1970—1982 (777). Illustration: Chippindall 1955 (fig. 402),
Clayton et al. 1970-1982 (fig. 180,4). Voucher: De Winter
& Marais 4827. PRECIS code 9900710-00820.

Andropogon huillensis Rendle

Grootwitbaardandropogon,
rietgras, large silver andropogon.

Perennial; tufted; 900-1800
mm tall. Leaf blades 80-400 mm
long; 2-4 mm wide. Spikelets
(sessile) 4—5 mm long (pedicel-
late sometimes slightly longer,
but usually reduced). Inflores-
cence plumose, with 5-7
flowering branches per culm; racemes 4- 1 0 per spathe, with
white silky hairs as long as the sessile spikelet; lower glume
of sessile spikelets deeply and broadly grooved.

Flowering September to June (but usually in autumn).
Wet places, usually on sand. Common. Biome: Savanna and
Grassland. Southern tropical Africa. Closely related to A.
eucomus and A. laxatus, which are smaller and have
suppressed pedicellate spikelets.

Description: Chippindall 1955 (500). Voucher: Repton
4058. PRECIS code 9900710-00900.

Andropogon lacunosus J.G. Anders.

Perennial; straggling; 300-
600 mm tall. Leaf blades 80-150
mm long; 2-5 mm wide. Spike-
lets (sessile) 5-7 mm long (pedi-
cellate longer). Racemes 2-3,
pedicels linear; lower glume of
sessile spikelets broadly rounded
on either side of a deep central
furrow, pitted between veins.

Flowering November to April. Swampy places at high
altitudes. Infrequent. Scattered in tropical Africa. Related
to A. distachyos, which lacks the glume pits.

Description: Clayton et al. 1970-1982 (770). Voucher:
Codd 6441. PRECIS code 9900710-01000.

Andropogon laxatus Stapf

Perennial; tufted; 300-900
mm tall. Leaf blades 20-200 mm
long; 2-3 mm wide. Spikelets
(sessile) 4-6 mm long (pedicel-
late reduced or suppressed). In-
florescence plumose; racemes
2-3 per spathe, with white silky
hairs as long as the sessile
spikelets; lower glume of ses-
sile spikelets deeply and broadly grooved.

Flowering October to March. Wet places. Rare and con-
servation status not known. Tropical Africa. Closely related
to A. eucomus, which has smaller spikelets, and A.
huillensis, which is a larger plant with more racemes.

Description: Chippindall 1955 (501), Clayton et al.
1970-1982 (775). Illustration: Clayton et al. 1970-1982
(fig. 180,2). Voucher: De Winter & Codd 218. PRECIS
code 9900710-01100.

Andropogon mannii Hook, f .

{-A. platybasis J.G.

Anders.) 3.

Perennial; densely tufted;

100-600 mm tall. Leaf blades
20-250 mm long; 2-8 mm wide.

Spikelets (sessile) 4. 5-8.0 mm
long (pedicellate equalling it or
slightly longer); 0. 9-1.1 mm
wide. Basal sheaths keeled, flattened; inflorescence with

42

short hairs; racemes 2-3 per spathe, 25-60 mm long,
pedicels linear, rounded; lower glume of sessile spikelets
shallowly concave below.

Flowering October to December. Mountain grassland in
moist places. Rare and conservation status not known.
Biome: Afromontane. Tropical Africa. The species is
variable over its range, with the forms on the disjunct
highland areas all differing slightly.

Description: Clayton et al. 1970-1982 (774). Voucher:
De Winter & Codd 199. PRECIS code 9900710-01250.

Andropogon ravus J.G. Anders.

Perennial; rhizomatous (rhi-
zomes branched, knotted); 1 50 —

900 mm tall. Leaf blades to 300
mm long; 2-7 mm wide. Spike-
lets (sessile) 7-9 mm long (pedi-
cellate considerably longer).

Plant glaucous grey; racemes
2(-3), pedicels cuneate-clavate;
lower glume of sessile spikelets
deeply grooved, glumes unawned; lemma awn 15-20 mm
long.

Flowering January to March. Mountain sourveld.
Locally common. Biome: Afromontane. Southern Africa.
Only doubtfully distinct from A. schirensis, with which it
intergrades.

Voucher: Killick 1261. PRECIS code 9900710-01400.

Andropogon schirensis A. Rich.

(=A. schirensis A. Rich. var.
angustifolius Stapf) 3.

Gesteektegras.

Perennial; densely tufted;

600-1200 mm tall. Leaf blades
90-600 mm long; 3-14 mm wide.

Spikelets (sessile) 5-7 mm long
(pedicellate slightly longer). Plant reddish; racemes 2(-5),
pedicels clavate; lower glume of sessile spikelets deeply
and very narrowly grooved, glumes unawned; lemma awn
25-30 mm long.

Flowering December to April. Open veld and rocky
hillsides. Common. Biome: Savanna and Grassland.
Tropical Africa. Closely related to Diheteropogon
amplectens , which has very similar sessile spikelets, but is
distinguished by the rounded leaf blade bases.

Description: Chippindall 1955 (497), Clayton et al.
1970-1982 (779). Voucher: Feely, Tinley & Ward 3.
PRECIS code 9900710-01600.

Anthephora Schreber

Hypudaerus A. Br.

Annual, or perennial; long-rhizomatous, or caespitose to
decumbent. Culms 150-1500 mm high: herbaceous;
branched above, or unbranched above. Leaf blades linear
to lanceolate; flat, or rolled. Ligule an unfringed membrane,
or a fringed membrane. Plants bisexual , with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant (the glomerules comprising 1-3 central perfect
spikelets with two or more outer, male, modified involucral
spikelets).

Inflorescence a false spike, with clusters of spikelets on
reduced axes (3-11 spikelets per glomerule)', espatheate.
Spikelet-bearing axes disarticulating (the glomerules being
reduced branches); falling entire (i.e. each glomerule falling
from the persistent main axis).

Spikelets associated with bractiform involucres (these
consisting of the leathery, expanded lower glumes of the
outer, involucral spikelets). The outer, involucral spikelets
of each glomerule are male-only, with a broad leathery
2-15 nerved G, and a setaceous G2- Female-fertile spikelets
compressed dorsiventrally; falling with the glumes. Glumes
two; very unequal, or more or less equal; owned ; very dis-
similar. Proximal incomplete florets I ; epaleate; sterile.

Female-fertile florets 1. Lemmas not becoming indu-
rated (membranous); hairless; having the margins lying flat
and exposed on the palea; with a clear germination flap, or
without a germination flap; 3-5 nerved; entire; awnless.
Palea present; relatively long. Stamens 3. Ovary glabrous.
Hilum short.

Photosynthetic pathway. C4. The anatomical
organization usually conventional, or unconventional
(rarely, doubtfully). Organization of PCR tissue when
unconventional, supposedly Arundinella type (see W.V.
Brown 1977, quoting Johnson 1965). XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, ,v = 9. Panicoideae; Panicodae; Paniceae. 12
species. Tropical and southern Africa, Arabia, tropical

Fig. 17. Anthephora pubescens

43

America. Mesophytic to xerophytic; in open habitats (in
dry. sandy savanna); glycophytic. Namibia, Botswana,
Transvaal, Orange Free State, and Cape Province. 4 indige-
nous species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by H.M. Anderson.

1(0). Plants annual; lower glume 7-13-nerved (rarely 5-
nerved), with a distinct convex lower half,

constricted and curving outwards above

A. schinzii

Plants perennial; lower glume 2-5-nerved, without a
distinct convex lower half, not constricted and

curving outwards above 2

2(1). Culms profusely branched; spikelets sparsely hairy

A. ramosa

Culms unbranched or only sparsely branched;

spikelets sparsely to densely hairy 3

3(2). Leaves blue-green, rigid, often folded, margins
smooth, tips shortly tapering to a stiff point;
spikelets 3-6 mm wide, lower glume acute or rarely

acuminate A. argentea

Leaves green, flat, margins crinkled, tips tapering to
a soft point; spikelets 5-10 mm wide, lower glume
acuminate or shortly awned A. pubescens

Anthephora argentea Goossens

(=A. angustifolia Goossens).

Perennial; tufted; to 1000 mm
tall. Leaf blades 100-150 mm
long; 1-3 mm wide. Spikelets
about 6 mm long; 2 mm wide.

Culms slender and wiry; leaf
blades blue-green, rigid, often
folded, tip shortly tapered to stiff
point; ligule up to 8 mm long and often split; inflorescence
3-6 mm wide; spikelets covered with hairs; lower glumes
acute, rarely acuminate.

Flowering November to April. Sandy soil often on
dunes, confined to Kalahari Thornveld. Infrequent. Biome;
Savanna and Nama-Karoo. Endemic. Natural pasture (high
nutritive value). Often mistaken for Elionurus muticus,
which has paired spikelets and the rachis of the false spike
curls and breaks up at maturity. A. angustifolia is here
synonymised. Chippindall 1955 (438) already remarked
that it was not distinct and differed only from A. argentea
in having the culms scantily branched.

Description: Muller 1984 (54), Chippindall 1955 (436).
Illustration: Muller 1984 (fig. 24), Chippindall 1955 (fig.
363). Voucher: Hansen 3333. PRECIS code 9901380-
00200.

Anthephora pubescens Nees

Wool grass; borseltjiegras.

Perennial; tufted; 300-1500
mm tall. Leaf blades 100-150
mm long; 3-5 mm wide. Spike-
lets about 8 mm long; 3 mm wide.

Culms not branched; leaves long-
tapering to soft point, often curl-
ing; inflorescence straw-coloured
or dull purple, 5-10 mm wide; spikelets densely covered
with hairs; lower glumes acuminate or shortly awned.

Flowering December to April. Shallow acid sandy soils,
often on hillsides. Common. Biome: Savanna, Grassland,
and Nama-Karoo. Southern and east Africa to Sudan and

Fig. 17.

Iran. Pasture (may be very palatable, has potential for
cultivation). Close to Tarigidia aequiglumis, which has a
paniculate inflorescence and glumes approximately equal.

Description: Muller 1984 (56), Chippindall 1955 (436).
Illustration: Muller 1984 (fig. 25), Chippindall 1955 (fig.
362). Voucher: Smook 4441. PRECIS code 9901380-
00300.

Anthephora ramosa Goossens

PI. 9.

Vertakte borseltjiegras.

Perennial; tufted; to 1200 mm
tall. Leaf blades 100-200 mm
long; 2-6 mm wide. Spikelets 6-7
mm long; 2-3 mm wide. Plants
forming lax tufts up to one meter
wide; culms branching profusely
from base; inflorescence 10 mm
wide; spikelets sparsely hairy; lower glume acute, rarely
acuminate.

Flowering February to May. Among rocks on hillsides
and ravines. Common. Biome: Nama-Karoo. Endemic.
Natural pasture. The characteristic branched habit
distinquishes A. ramosa from other Anthephora species.

Description: Muller 1984 (58), Chippindall 1955 (438).
Illustration: Muller 1984 (fig. 26), Chippindall 1955 (fig.
364). Voucher: De Winter 3308. PRECIS code
9901380-00400.

Anthephora schinzii Hack.

Annual wool grass, eenjarige
borseltjiegras.

Annual; tufted; 120-350 mm
tall. Leaf blades 60-100 mm
long; 4-6 mm wide. Spikelets
usually about 10 mm long; 3 mm
wide. Inflorescence 10 mm wide;
spikelets in groups of five; lower
glumes are convex on the lower half, have a central
constricted area and then curve outwards, with the tip being
acute or awned, and length variable up to 15 mm long.

Flowering December to April. Pioneer grass on sandy
soils. Locally common. Biome: Savanna and Nama-Karoo.
Angola. Natural pasture.

Description: Muller 1984 (60), Chippindall 1955 (438).
Illustration: Muller 1984 (fig. 27). Voucher: Maguire 2164.
PRECIS code 9901380-00500.

PI. 10.

Anthoxanthum L.

Flavia Fabric., Foenodorum Kraus e,Xanthonanthus St-
Lager.

Annual, or perennial; caespitose to decumbent. Culms
50-900 mm high; herbaceous; unbranched above. The
shoots aromatic (coumarin-scented). Leaf blades linear to
lanceolate; Hat. Ligule an unfringed membrane . Plants
bisexual, with bisexual spikelets.

Inflorescence a single raceme (rarely), or paniculate ;
contracted; espatheate. Spikelet-bearing axes persistent.

Spikelets 5-10 mm long; compressed laterally; disartic-
ulating above the glumes. Glumes two; very unequal; long
relative to the adjacent lemmas (i.e., the longer glumes);
awriless; similar (membranous). Proximal incomplete
florets 2 .

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes; 1-7 nerved; entire, or incised; awnless, or
awned. Awns when present 1 ; geniculate; much shorter than
the body of the lemma, to much longer than the body of the
lemma. Palea present; relatively long. Stamens 2, or 3
(rarely). Ovary glabrous. Fruit small; hilum short; embryo
small.

44

Cytology, classification, distribution. Chromosome base
number, x — 5. Pooideae; Poodae; Aveneae. 20 species.
North temperate & mountains of tropical Africa & Asia.
Mesophytic; in shade and in open habitats (meadows,
grasslands and in light shade). Transvaal, Orange Free
State, Natal, Lesotho, and Cape Province. Indigenous
species (4), naturalized species (1).

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

1(0). Lower glume 1 -nerved 2

Lower glume 3-5-nerved 3

2(1). Lower lemma about 3 mm long, dark brown and

densely hairy A. odoratum

Lower lemma about 5 mm long, pale to light brown

and usually sparsely hairy A. ecklonii

3(1). Panicle small, oblong, contracted or reduced to a
scanty raceme; plants fine and weak, leaves soft .

A. tongo

Panicle spike-like, fairly dense, occasionally
interrupted near the base; plants usually distinctly

tufted and erect, leaves rigid and pungent

A. dregeanum

Anthoxanthum brevifolium Stapf

Perennial; rhizomatous and
tufted; 150-220 mm tall.

Flowering March. Biome; Sa-
vanna. This species is known only
from the type collection (Galpin
6884). Except for the very short
and broad leaf blades this
specimen cannot be distinguished
from A. ecklonii and therefore it
is not regarded as a distinct taxon. The genus as a whole is
in great need of revision.

Description; Chippindall 1955 (92). PRECIS code
9901640-00100.

Anthoxanthum dregeanum (Nees) Stapf

Perennial; rhizomatous and
tufted; 200-600 mm tall. Leaf
blades to 250 mm long; to 9 mm
wide. Spikelets 6-7 mm long.

Leaf blades rigid, often folded;
panicle spike-like, occasionally
interrupted near the base; lower
glume 3-nerved.

Flowering October to J anuary .

On moist mountainslopes. Infrequent to locally common.
Biome: Fynbos. Endemic. Sometimes not clearly

distinguished from A. tongo, which normally is a very fine
plant with a scanty panicle.

Description: Stapf 1898—1900 (466), Chippindall 1955
(92). Illustration: Chippindall 1955 (fig. 63). Voucher:
Esterhuysen 26575. PRECIS code 9901640—00200.

Anthoxanthum ecklonii (Nees ex Trin.) Stapf

Fig. 18. PI. 11.

Perennial; loosely or densely
tufted and rhizomatous; 350-800
mm tall. Leaf blades 70-250 mm
long; 4-9 mm wide. Spikelets 6—8
mm long. Bases of culms usually
bulbous; panicle spike-like,

40-130 mm long; lower glume 1 -
nerved; lower lemma about 5 mm
long, pale to light brown, usually
sparsely hairy.

Fig. 18. Anthoxanthum ecklonii

Flowering December to April. Usually in moist places
such as streamsides and on grassy mountain slopes,
extending to forest margins. Infrequent (but fairly
widespread). Biome: Fynbos, Savanna, and Grassland.
Possibly Malawi. Resembles A. odoratum, which has
lemmas shorter, darker and densely hairy.

Description: Stapf 1898—1900 (466), Chippindall 1955
(92). Illustration: Chippindall 1955 (fig. 64). Voucher:
Killick 1296. PRECIS code 9901640-00300.

45

Anthoxanthum odoratum L.

Sweet vernal grass.

Perennial; loosely or densely
tufted; 300-600(-1000) mm tall.

Leaf blades 150-300 mm long;

2-8 mm wide. Spikelets 7-10 mm
long. Panicle spike-like, 10-90
mm long; lower glume 1 -nerved;
lower lemma about 3 mm long,
dark brown, densely hairy.

Flowering October to February. Humic soils in moist,
swampy areas. Rare. Locally common. Naturalized from
Europe. Biome: Savanna. Eurasia. Resembles A. ecklonii,
which has the lemma about 5 mm long, lighter coloured and
sparsely hairy.

Description: Hitchcock & Chase 1950 (528), Chippin-
dall 1955 (93). Illustration: Hitchcock & Chase 1950 (fig.
1114). Voucher; Acocks 22118. PRECIS code 9901640-
00400.

Anthoxanthum tongo (Trin.) Stapf

Perennial; culms very fine,
straggling or loosely tufted
(occasionally mat-forming); 100-
400 mm tall. Leaf blades
20— 1 00(— 170) mm long; filiform,
to 2 mm wide. Spikelets 5-7 mm
long. Panicle small, oblong,
contracted or reduced to a scanty
raceme, with very few spikelets;
lower glume 3-5-nerved.

Flowering September to February. In moist shady places
in the shelter of rocks and in shallow crevices. Locally com-
mon. Biome: Fynbos. Endemic. Many specimens deposited
under A. tongo and A. dregeanum at PRE seem to be
misplaced. This problem cannot be solved within the
current classification and stresses the need for a revision of
this genus which is very poorly studied in the FSA region.

Description: Stapf 1898-1900 (467), Chippindall 1955
(92). Illustration: Chippindall 1955 (fig. 62). Voucher:
Esterhuysen 33603. PRECIS code 9901640-00500.

Aristida L.

Aristopsis Catasus , Arthratherum P. Beauv. , Chaetaria
P. Beauv., Curtopogon P. Beauv., Kielboul Adans.,
Moulinsia Raf., Streptachne R. Br., Trixostis Raf.

Annual, or perennial; caespitose. Culms
100— 1000(— 1 800) mm high; herbaceous; branched above,
or unbranched above. Leaf blades linear, or linear-
lanceolate; flat, or rolled. Ligule a fringed membrane to a
fringe of hairs.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 4-30 mm long ; compressed laterally to not
noticeably compressed; disarticulating above the glumes.
Rachilla terminated by a female-fertile floret. Glumes two
(membranous to papery); relatively large; very unequal, or
more or less equal; or at least the G2 about equalling the
spikelets (or longer); awned, or awnless; very dissimilar, or
similar. Lower glume 1 nerved. All florets female-fertile;
proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (narrow, cylindrical); hairy (rarely), or hairless;
with a clear germination flap; 1-3 nerved; entire; awned.
Awns usually triple or trifid, commonly with a basal
column, or not of the triple/trifid, basal column type (the
column sometimes absent, the lateral branches sometimes
reduced or absent); 1 , or 3. Awns apical; non-geniculate (at
least, not geniculate in the normal sense)', hairless (usually

glabrous); much shorter than the body of the lemma, to
much longer than the body of the lemma. Palea present;
conspicuous but relatively short, or very reduced; 1-nerved,
or 2-nerved, or nerveless. Lodicules when present 2;
membranous; glabrous. Stamens 1-3. Ovary glabrous. Fruit
small to large (3-1 1 mm); fusiform; hilum short, or long-
linear; pericarp fused; embryo large.

Fig. 19. Aristida congesta subsp. congesta

46

Photosynthetic pathway. C4. The anatomical
organization unconventional. Organization of PCR tissue
Aristida type. Biochemical type NADP-ME (3 species);
XyMS- (with double PCR sheaths).

Cytology, classification, distribution. Chromosome base
number, x = 11 and 12. Arundinoideae; Aristideae. 290
species. Temperate and subtropical. Xerophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 27 indigenous species.

References. 1. De Winter. 1965. Bothalia 8: 199. 2.
Melderis. 1971. FZ. 3. Giess. 1971. Bothalia 10: 365.

Species treatment by L. Smook.

1(0). Lemma awn solitary or the two lateral awns poorly
developed, less than 1/4 the length of the central

awn 2

Lemma awns three, lateral awns well developed,
longer than 1/4 the length of the central awn . . 3
2(1). Plants annual; lemma articulation between the apex of

the lemma and the base of the column

A. parvula

Plants perennial; lemma without articulation

A. transvaalensis

3(1). Lower glume without an awn or mucro 4

Lower glume with an awn or mucro (which can be

minute) 24

4(3). Lower glume longer than the upper glume (Note: in
some species the long, delicate apex breaks off

early) 5

Lower glume shorter than to equalling the upper

glume 6

5(4). Plants annual; spikelets 25-30 mm long (including
awns); lower glume narrowing abruptly to an acute
apex; lemma not narrowed into a beak or twisted

column A. effusa

Plants perennial; spikelets to 20 mm long (including
awns); lower glume long, tapering to an acuminate
apex; lemma narrowed into a beak or short twisted

column A. monticola

6(4). Lower glume 2/3 as long as to longer than the upper

glume 7

Lower glume 1/2-2/3 the length of the upper glume

10

7(6). Plants perennial 8

Plants annual 9

8(7). Lemma without a column; callus subobtuse to
rounded; lower glume firm throughout, but the

extreme tip membranous

A. canescens subsp. canescens

Lemma with a column; callus emarginate to distinctly
bifid; lower glume firm below, upper 2/3

membranous and often torn A. spectabilis

9(7). Inflorescence narrowly oblong to lanceolate; lemma
usually scabrid only on the keel . A. adscensionis
Inflorescence ovate; lemma usually scabrid all over
except on the lower 1/4 A. effusa

10(6). Plants annual 11

Plants perennial 12

11(10). Lower glume 3. 5-4.0 mm long, broadly oblong,

apex obtuse to truncate or slightly emarginate,
fimbriate; spikelets bright yellow with glume tips

dark A. dewinteri

Lower glume 6-9 mm long, lanceolate, apex acute,
not fimbriate; spikelets dull yellow to purple but
glume tips not with a dark patch . A. stipoides
12(10). Lemma articulation absent or inconspicuous . . 13
Lemma articulation present between the apex of the

lemma and the base of the column 16

13(12). Plants robust; culm diameter 5-6 mm . A. sciurus

Plants slender; culm diameter 1. 5-3.0 mm ... 14
14(13). Culms much branched from the upper nodes; leaves
folded, straight, rigid, erect ... A. dasydesmis
Culms unbranched or sparsely branched from the

upper nodes; leaves flat or rolled, curved or

slightly curled, not straight and erect 15

15( 14). Spikelets to 22 mm long (including awns),
congested on the inflorescence branches; callus

tip naked, rounded and swollen

A. canescens subsp. canescens

Spikelets 25-40 mm long (including awns), distinct
from one another on the inflorescence branches;

callus tip bifid A. diffusa subsp. burkei

16(12). Lower intemodes of culms pubescent to woolly-
hairy, upper intemodes pubescent or glabrous .

17

Lower and upper culm internodes glabrous ... 18
17(16). Lemma column 8-20 mm long; inflorescence
oblong to broadly oblong, symmetrical, to 800
mm long and over 200 mm wide, much branched

A. meridionalis

Lemma column 5-7 mm long; inflorescence
narrowly oblong to narrowly elliptic, usually
asymmetric, to 200 mm long and 120 mm wide

A. vestita

18(16). Callus tip truncate, obliquely truncate or slightly

emarginate; leaves erect and rigid

A. dasydesmis

Callus tip bifid; leaves curved or bent, flexible, not

rigid 19

19(18). Lower glume with upper 1/3 membranous, often

tom and broken A. spectabilis

Lower glume apex firm or only the very tip

membranous 20

20(19). Some leaf auricles with long woolly hairs

A. meridionalis

All leaf auricles glabrous or pubescent but not with

long woolly hairs 21

21(20). Culms much branched

A. engleri var. ramosissima

Culms unbranched or sparsely branched 22

22(21). Lower glume apex acute, finely fimbriate

A. engleri var. engleri

Lower glume apex obtuse, usually entire, only

occasionally coarsely fimbriate 23

23(22). Upper glume 12-18 mm long

A. diffusa subsp. diffusa

Upper glume to 12 mm long

A. diffusa subsp. burkei

24(3). Lemma articulation present (sometimes shown only
by a swollen line or colour differentiation,
usually directly below the branching point of the

awns) 25

Lemma articulation absent 34

25(24). Lemma articulation between the apex of the lemma
and the base of the long column; callus 1. 5-3.0

mm long, tip acuminate, pungent 26

Lemma articulation not as above; callus 0. 5-1.5
mm long, tip narrowly or broadly rounded to

truncate 31

26(25). Lower intemodes of culms woolly to densely

tomentose 27

Lower intemodes of culms glabrous or pubescent

but not with woolly hairs 28

27(26). Inflorescence contracted, spikelike, very dense . .

A. mollissima subsp. mollissima

Inflorescence narrow, lax, more or less divarcately
branched .... A. mollissima subsp. argentea
28(26). Inflorescence spikelike, sometimes interrupted
towards the base, branches closely appressed to

the main axis 29

Inflorescence not spikelike, much interrupted,

branches suberect or spreading 30

29(28). Plants robust, to 1500 mm tall, sparsely branched
at upper nodes; inflorescence usually 150-300

mm long A. stipitata subsp. stipitata

Plants slender, to 600 mm tall, much branched at
the upper nodes; inflorescence usually to 1 50 mm

long A. stipitata subsp. spicata

30(28). Plants robust; culm diameter 2. 5-4.0 mm;

47

inflorescence usually 200-350 mm long

A. stipitata subsp. robusta

Plants slender; culm diameter 1.0-2. 5 mm;

inflorescence usually 100-200 mm long

A. stipitata subsp. graciliflora

31(25). Culm internodes pubescent; lemma slightly
narrowed at apex but column absent, articulation
between the apex of the lemma and the branching

point of the awns A. hordeacea

Culm internodes glabrous; lemma narrowed into a
distinct column, articulation between the apex of
the column and the branching point of the awns

32

32(31). Spikelet clusters linear to oblanceolate (including
awns), longest pedicel to 5.5 mm long; plants

robust and coarse A. pilgeri

Spikelet clusters narrowly obovate (including
awns), longest pedicel to 1.5 mm long; plants

usually slender 33

33(32). Inflorescence very dense, branches closely
appressed and covering main axis except
occasionally at the base where 1-2 subspicate

branches spread from the main axis

A. congesta subsp. congesta

Inflorescence variable, with many side branches
spreading from and exposing the main axis, these
either with a few spikelets laxly clustered or
many spikelets densely clustered at the ends of
the long, naked side branches, or with spikelets

appressed all along the side branches

A. congesta subsp. barbicollis

34(24). Plants annual 35

Plants perennial to subperennial 40

35(34). Lemma narrowly elliptic, dorsally compressed . .

A. hubbardiana

Lemma linear, laterally compressed 36

36(35). Lower glume with a distinct, robust awn 0.8-3. 5

mm long 37

Lower glume mucronate or with a short awn to 0.8

mm long 39

37(36). Spikelets coarse, 35-50 mm long (including awns)

A. rhiniochloa

Spikelets slender and fine, 10-30 mm long

(including awns) 38

38(37). Inflorescence delicate, branchlets and pedicels
spreading, with spikelets distant from one

another at the end of the branches

A. scabrivalvis subsp. scabrivalvis

Inflorescence robust, branchlets and pedicels
appressed, with spikelets densely congested at

the ends of branches

A. scabrivalvis subsp. contracta

39(36). Inflorescence oblong to ovate, 80 mm or wider, side
branches spreading from the main axis with lax
clusters of 2-3 spikelets distant from each other
at the ends; lower glume 3/4 as long to equaling,

sometimes longer than the upper glume

A. effusa

Inflorescence linear to lanceolate, usually 10-50
mm wide, side branches appressed to main axis,
spikelike but interrupted towards the base, or
open and spreading, spikelets densely clustered;
lower glume usually 2/3 or 3/4 the length of the

upper glume A. adscensionis

40(34). Inflorescence open, branches rigid, spreading at 90
degrees from the main axis; lower glume equal
to or longer than the upper glume . A. bipartita
Inflorescence contracted, usually dense, branches
flexible, erect or spreading not more than 45
degrees from the main axis; glumes variable, but
lower glume never longer than the upper glume

41

41(40). Spikelets to 14 mm long (including awns)

A. recta

Spikelets 15^10 mm long (including awns) ... 42

42(41 ). Lemma oblong, almost the same width throughout,
sometimes with a minute constriction at the
branching point of the awns; column absent . .

A. canescens subsp. canescens

Lemma narrowly lanceolate, distinctly tapering
towards the branching point of the awns; column

present or absent 43

43(42). Leaves mainly basal, forming a dense basal tuft in
which the culms are hidden; culms mainly

unbranched in the upper nodes 44

Leaves mainly cauline, or not basally dense and the
culms obvious for most of their length; culms
branched or unbranched in the upper nodes . 45
44(43). Rhizomes long, oblique, thin and creeping;

spikelets 15-30 mm long (including awns);
glumes usually very unequal; high eastern
mountains . . . A. junciformis subsp. galpinii
Rhizomes short and stout; spikelets 20-35 mm long
(including awns); glumes equal to subequal;
bushveld and the highveld ... A. aequiglumis
45(43). Culms much branched, usually at every node;

lateral awns usually rudimentary, very much

shorter and thinner than the central awn

A. transvaalensis

Culms branched, but not at every node; lateral awns
well developed, shorter or subequal to the central

awn, never rudimentary

A. junciformis subsp. junciformis

Aristida adscensionis L.

( =A . curvata (Nees) Dur. &
Schinz ) 1; ( =A . submucronata
Schumach.) 1; (=A. adscensionis
L. subsp. guineensis (Trin. &
Rupr.) Hem'.) 2.

Annual bristle grass, steek-
gras.

PI. 12.

Annual; tufted (erect, often branched); to 1000 mm tall.
Leaf blades to 300 mm long; to 3 mm wide. Spikelets 1 0-40
mm long (including awns). Inflorescence narrowly oblong
to lanceolate, 10-50 mm wide, usually spikelike, interrupted
at base, with the side branches appressed to main axis,
sometimes branches open and spreading, with the spikelets
densely clustered on the branches; lower glume 2/3— 3/4 the
length of the upper glume, sometimes with a mucro or short
awn to 0.8 mm long; lemma laterally compressed, articula-
tion and column absent; awns three, laterals well developed,
shorter than the central awn; callus with rounded, naked tip.

Flowering December to September. Stony, sandy loam,
clayey, calcareous, shallow soils on stony hills, moist areas
along pans and rivers, along roads and other disturbed
ground. Common. Biome: Savanna, Grassland, and Nama-
Karoo. Throughout the tropics. Pasture (only grazed when
very young), or indicator (of advanced retrogression of veld
and disturbed ground), or weed (troublesome in wool, also
causes sores by piercing the sheep's skin). This species has
a very wide geographical distribution and exhibits a
considerable variation in its external morphology. In the
past it has been divided into species, subspecies and
varieties. The treatment of Melderis (1972) is followed
pending a detailed study. Some forms resemble A.
hubbardiana , which has the lemma dorsally compressed,
and forms of A. congesta, which have a lemma articulation
between the apex of the column and the branching point of
the awns.

Description: De Winter 1965, Melderis 1971 (110).
Illustration: Muller 1984 (fig. 28). Voucher: Smook 2781,
Van Jaarsveld 179, Schmitz 1523. PRECIS code
9902620-00050.

48

Aristida aequiglumis Hack.

Curly-leaved three-awned
grass.

Stout and shortly rhizomatous
and tufted (densely); to 800 mm
tall. Leaf blades to 150 mm long;
to 1 mm wide. Spikelets 20-35
mm long (including awns). Culms
usually unbranched; leaves main-
ly basal, forming a dense basal tuft, enclosing the culms for
most of its length; inflorescence contracted, dense, much
branched, branches not spreading more than 45 degrees
from the main axis; glumes equal to subequal, often
pubescent, lower glume mucronate; lemma narrowly lan-
ceolate, tapering upwards, articulation absent; column long
and twisted; awns three, laterals shorter but well developed;
callus tip naked, truncate to slightly emarginate.

Flowering December to May. Sandy, shallow soils on
rocky hillslopes or in seasonally flooded areas. Common.
Biome: Savanna and Grassland (occasionally). Zimbabwe
to Zambia. Indicator (of eroded soils). Although this species
occurs on the highveld, it is more common in bushveld
areas. It resembles A. junciformis subsp. galpinii, which has
a long, thin, oblique rhizome, generally unequal glumes and
is found on the high mountain sourveld.

Description: De Winter 1965 (262), Melderis 1971
(107). Voucher: Smook 2709, Smook 1435. PRECIS code
9902620-00200.

Aristida bipartita (Nees) Trin. & Rupr.

Three-awned rolling grass.

Perennial (to subperennial);
tufted (erect or geniculate); to
650 mm tall. Leaf blades to 200
mm long; to 2 mm wide. Spike-
lets 18-20 mm long (including
awns). Inflorescence open, bran-
ches rigid, naked for most of their
length, spreading at 90 degrees from the main axis, spike-
lets borne at the tips of the long, naked branches; lower
glume equalling or longer than the upper glume, awned;
lemma articulation and column absent; awns three, sub-
equal; callus tip thickened, naked, rounded to obtuse.

Flowering October to May. Sandy, stony, loamy, clayey
and black turf soils in moist areas around vleis and dongas,
and in overgrazed and other disturbed ground. Common.
Biome: Savanna and Grassland. Mozambique. The whole
inflorescence breaks off at maturity and is rolled about as
a tumbleweed by the wind. Resembles A. effusa, which is
a definite annual and has the lower glume with or without
a mucro, and resembles forms of A. scabrivalvis, which is
an annual with the lower glume less than 2/3 the length of
the upper glume (excluding awns).

Description: De Winter 1965 (256), Melderis 1971
(114). Voucher: Scheepers 1574. PRECIS code 9902620-
00400.

Aristida canescens Henr. subsp. canescens

Vaalsteekgras.

Slender perennial; tufted (e-
rect); to 1500 mm tall. Leaf blad-
es to 300 mm long; to 2 mm wide.

Spikelets to 22 mm long (includ-
ing awns). Culms 1. 5-3.0 mm in
diameter, unbranched to sparsely
branched from the upper nodes;
leaves flat or rolled, curved to curled; spikelets congested
on the inflorescence branches; lower glume firm except for
the very tip, which is membranous, 1/2 as long as to nearly
equalling the upper glume, without mucro or awn; lemma

articulation and column absent; awns three; callus tip nak-
ed, subobtuse to rounded.

Flowering December to May. Shallow, sandy, stony
soils on rocky ridges, eroded and disturbed ground. Locally
common. Biome: Savanna and Grassland. Zimbabwe to
Zambia. Differs from subsp. ramosa , which has the culms
branched from the upper nodes and the lower glume awned.
Resembles A. junciformis, which has the lower glume awn-
ed or mucronate, A. pilgeri, which has the lemma articula-
tion present, and A. sciurus , which has a culm diameter of
5-6 mm.

Description: De Winter 1965 (260). Voucher: Smook
2063, De Winter 7561. PRECIS code 9902620-00500.

Aristida canescens Henr. subsp. ramosa De Winter

Perennial; tufted (culms some-
times geniculate); to 600 mm tall.

Leaf blades to 150 mm long; to
1.5 mm wide. Spikelets 15-18
mm long (including awns). Culms
branched from the upper nodes;
inflorescence contracted, branch-
es erect or spreading not more
than 45 degrees from the main ax-
is; lower glume more than 2/3 the length of the upper, awn-
ed; lemma oblong, almost the same width throughout,
sometimes with a minute constriction at the branching point
of the awns, articulation and column absent; awns three,
subequal; callus tip naked, swollen, rounded.

Flowering July, December, February, and March.
Dolerite slopes. Infrequent. Biome: Nama-Karoo. Endemic.
Differs from subsp. canescens , which has the culm mainly
unbranched and the lower glume unawned.

Description: De Winter 1965 (262). Voucher:

Donaldson 318. PRECIS code 9902620-00550.

Aristida congesta Roem. & Schult. subsp. barbicollis

(Trin. & Rupr.) De Winter

(=A. barbicollis Trin. &

Rupr.) 1.

Spreading prickle grass,
witsteekgras.

Perennial, or annual (slender);
tufted; to 750 mm tall. Leaf blad-
es to 200 mm long; to 3 mm wide. Spikelets 20-30(-50)
mm long (including awns). Inflorescence variable, with
many side branches spreading from and exposing the main
axis, either with a few spikelets laxly clustered or with
many spikelets densely clustered at the ends of long, naked
side branches or spreading to the base of the side branches;
spikelet clusters narrow, obovate to ovate (including awns),
longest pedicel to 1 .5 mm long; lower glume awned; lemma
articulation between the apex of the column and the branch-
ing point of the awns, articulation sometimes represented
only by a swollen line or colour differentiation; column
present; awns three, laterals well developed; callus 0.5-1. 5
mm long, tip naked, narrow or broadly rounded to truncate.

Flowering October to May. Deep, sandy clayey soils on
rocky hillsides, old lands and disturbed ground. Common.
Biome: Savanna and Grassland. Northwards to East Africa.
Indicator (poor veld management or other disturbances), or
weed (the floret callus penetrates sheep skins causing
sores). The spikelets of the two subspecies cannot be
distinguished from each other, the inflorescence shape be-
ing the main character used but this tends to intergrade.
Thus there are some plants that cannot be referred to with
certainty to either subspecies. In this treatment, pending
further study, specimens with inflorescences open and lax
or those with most of the side branches spreading from and
exposing the main axis are included in subsp. barbicollis.

Description: De Winter 1965 (296), Melderis 1971
(129). Illustration: Melderis 1971 (tab. 33). Voucher:

(

49

Smook 2787, 5703, Codd 4865, Herbst 50. PRECIS code
9902620-00800.

Aristida congesta Roem. & Schult. subsp. congesta

Fig. 19.

(=A. alopecuroides Hack.) 1;

(-A. longicauda Hack. &

Henriques) 1.

Katstertsteekgras.

Slender perennial, or annual
(occasionally); densely tufted; to
900 mm tall. Leaf blades to 300
mm long; to 5 mm wide. Spikelets 25-30 mm long (includ-
ing awns). Inflorescence very dense, branches closely ap-
pressed, enclosing main axis except occasionally at the base
where 1-2 subspicate side branches spread away from the
main axis; spikelet clusters narrowly obovate to obovate;
lower glume awned; lemma articulation between the apex
of the column and the branching point of the awns, articula-
tion sometimes represented only by a swollen line or a
colour differentiation; column present; awns three, laterals
well developed; callus tip naked, narrowly to broadly
rounded to truncate.

Flowering December to May. Hard or stony loam, sandy
basalt, black clayey soils, Kalahari sands on stony slopes,
open eroded places, old lands, road verges and other
disturbed ground. Common. Biome: Savanna and Grass-
land. Northwards to northeast and east Africa and the
Mediterranean. Pasture (for small stock only), or indicator
(of retrogression of veld), or weed (tangles in wool, and the
floret callus pierces the skin and causes sores). The spike-
lets of the two subspecies cannot be distinguished from each
other, inflorescence shape being the main character used
although it tends to intergrade. Thus there are some plants
that cannot be referred to with certainty to either
subspecies. In this treatment, pending further studies,
specimens with very dense inflorescences, with branches
closely appressed and covering the main axis, occasionally
interrupted at the base by 1-2 spreading branches are in-
cluded in this subspecies. Resembles A. hordeacea , which
has culm intemodes pubescent and column absent, and A.
hubbardiana, which has the lemma articulation absent.

Description: De Winter 1965 (296), Melderis 1971
(127). Illustration: Muller 1984 (fig. 29). Voucher: Theron
1264, Giess, Volk & Bleissner 7027, De Winter & Marais
4128. PRECIS code 9902620-00850.

Aristida dasydesmis (Pilg.) Mez

Slender perennial; densely
tufted; to 800 mm tall. Leaf blad-
es to 300 mm long; about 1 mm
wide. Spikelets 25-30 mm long
(including awns). Culms 1. 5-3.0
mm in diameter, much branched
from the upper nodes; leaves
mainly cauline, folded, straight,
erect and rigid; lower glume just
over 1/2 the length of the upper glume, mucro or awn ab-
sent; lemma articulation between the apex of the lemma and
the base of the column, sometimes inconspicuous or absent;
column to 6 mm long; awns three, laterals well developed;
callus tip naked, truncate, obliquely truncate or slightly
emarginate.

Flowering August to September. Granite slopes in arid
areas. Locally common. Biome: Succulent Karoo. Endemic.
Resembles A. junciformis subsp. junciformis and A.
transvaalensis, which both have the lower glumes awned
or mucronate. Similar to A. vestita, which has the lower
internodes pubescent to woolly-hairy, and A. diffusa , which
has flexible leaves.

Description: De Winter 1965 (275). Voucher: Acocks
19518. PRECIS code 9902620-01000.

Aristida dewinteri Giess

Annual; tufted; to 1000 mm
tall. Leaf blades to 300 mm long;
to 3 mm wide. Spikelets 40-50
mm long (including awns). Spike-
lets bright yellow, with a dark
patch at the apex of the glumes;
lower glume 3. 5^1.0 mm long,
broadly oblong, apex obtuse to
truncate or slightly emarginate,
fimbriate, 1/2 the length of the upper glume, without a mu-
cro or awn; lemma articulation between the apex of the lem-
ma and the base of the column; column 1.3-1. 4 mm long;
awns three, laterals well developed but shorter than the cen-
tral awn; callus tip naked, distinctly bifid.

Flowering April. Rare. Locally common (where found).
Biome: Nama-Karoo. Endemic, or possibly also in Angola.
Only the holotype was available for this treatment.

Description: Giess 1971 (365). Voucher: Giess 9345
(holotype PRE). PRECIS code 9902620-01 100.

Aristida diffusa Trin. subsp. burkei (Stapf) Meld.

(-A. diffusa Trin. var. burkei
(Stapf) Schweick.) 2.

Koperdraadgras, ystergras.

Slender perennial; densely
tufted; to 1000 mm tall. Leaf
blades to 300 mm long; to 2 mm
wide. Spikelets 25-40 mm long
(including awns). Culms 1. 5-3.0 mm in diameter, unbran-
ched or sparsely branched; leaf auricles glabrous or short-
hairy, not woolly; inflorescence open, with spikelets distant
from one another; lower glume obtuse, occasionally coarse-
ly fimbriate at the apex, 1/2-2/3 the length of the upper
glume, mucro or awn absent; upper glume to 12 mm long;
lemma articulation present between the apex of the lemma
and the base of the column, or absent; column present; cal-
lus tip naked, deeply bifid.

Flowering November to April. Dry, sandy, gravelly
loam soils on hilly slopes. Common. Biome: Savanna,
Grassland, and Nama-Karoo. Zimbabwe. Indicator (of
overgrazing). Barely distinguished from subsp. diffusa ,
from which it is separated by the longer upper glume and
the distribution. Further study is needed.

Description: De Winter 1965 (275), Melderis 1971
(118). Voucher: Smook 6404, 3442, Smook & Gibbs
Russell 2275. PRECIS code 9902620-01200.

Aristida diffusa Trin. subsp. diffusa

(=A. diffusa Trin. var.
genuina Henr.) 1; ( -A . diffusa
Trin. var. pseudo-hystrix (Trin.

& Rupr.) Henr.) 1 .

Slender perennial; densely
tufted (erect); to 750 mm tall.

Leaf blades to 300 mm long; to 2
mm wide. Spikelets 25 — 45 mm
long (including awns). Culms unbranched or sparsely bran-
ched; leaves flexible, leaf auricles glabrous or short-hairy,
not woolly; lower glume 1/2-2/3 the length of the upper
glume, apex obtuse, occasionally coarsely fimbriate; upper
glume 12-18 mm long; lemma articulation present between
the apex of the lemma and the base of the column, or absent;
column present; awns three, laterals well developed; callus
tip naked, deeply bifid.

Flowering October. Sandy soils, between rocks and in
disturbed places. Infrequent. Biome: Fynbos. Endemic.
Barely distinguishable from subsp. burkei, from which it is
mainly separated by the shorter upper glume. Further study
is needed.

50

Description: De Winter 1965 275. Voucher: Liebenberg
4240. PRECIS code 9902620-01300.

Aristida effusa Henr.

Spreading steekgras, pluim-
steekgras.

Annual; tufted (branched, e-
rect); to 900 mm tall. Leaf blades
to 300 mm long; to 3 mm wide.

Spikelets 25-32 mm long (includ-
ing awns). Inflorescence open,
ovate, spikelets clustered at the
end of lax, flexible branches; lower glume 2/3 as long to
longer than the upper glume, narrowed abruptly into a short,
acute apex, mucro or awn absent; lemma usually scabrid
except for lower 1/4, articulation absent; column or beak
absent, awns three, laterals shorter; callus tip naked,
swollen, rounded.

Flowering February to May. Calcareous, sandy loam,
stony soils along roadsides. Locally common. Biome: Sa-
vanna. Endemic. Indicator (retrogression of veld).
Specimens from Botswana previously referred to as A.
wildii Meld, have been placed here pending a more detailed
study. Resembles some forms of A. scabrivalvis, which has
the lower glume strongly awned, and A. bipartita , which is
perennial with the lower glume awned.

Description: De Winter 1965 (251). Illustration: Muller
1984 (fig. 30). Voucher: De Winter & Leistner 5174, Field
3051. PRECIS code 9902620-01400.

Aristida engleri Mez var. engleri

Engler’s bristle grass, bristle
three-awn.

Densely tufted (erect to genic-
ulate); to 700 mm tall. Leaf blad-
es to 150 mm long; to 2.5 mm
wide. Spikelets 25-35 mm long
(including awns). Culms unbran-
ched or sparsely branched; leaves
flexible; leaf auricles glabrous to short-hairy, not woolly;
lower glume 1/2 the length of the upper glume, apex acute,
finely fimbriate; lemma articulation between the apex of the
lemma and the base of the column; column present; awns
three, laterals well developed; callus tip naked, deeply bifid.

Flowering February to August. Rocky outcrops. Infre-
quent. Biome: Savanna and Nama-Karoo. Endemic. This
variety is not always distinguishable from var. ramosissima,
which has the culms much branched. Resembles A. diffusa ,
which has the lower glume with an obtuse apex which is
only occasionally coarsely fimbriate, and A. vestita, which
has lower internodes pubescent to woolly-hairy.

Description: De Winter 1965 (281). Illustration: Muller
1984 (fig. 31). Voucher: Giess & Muller 11954, Theron
1972. PRECIS code 9902620-01500.

Aristida engleri Mez var. ramosissima De Winter

Perennial; tufted (to sprawl-
ing); to 900 mm tall. Leaf blades
to 150 mm long; to 2 mm wide.

Spikelets 20-25 mm long (includ-
ing awns). Culms much branched;
leaves flexible, leaf auricles
glabrous or shortly pubescent;
lower glume to 1/2 the length of
the upper glume; lemma artic-
ulation between the apex of the lemma and the base of the
column; column present; awns three; callus tip naked, deep-
ly bifid.

Flowering January to June. Red sandy soils between
rocks on hillsides. Locally common. Biome: Savanna and
Nama-Karoo. Endemic. This variety is not always

distinguishable from var. engleri, which has culms unbran-
ched or sparsely branched.

Description: De Winter 1965 (281). Illustration: Muller
1984 (fig. 32). Voucher: Liebenberg 5228. PRECIS code
9902620-01600.

Aristida hordeacea Kunth

Jakkalsstert, garssteekgras.

Annual; tufted (erect to genic-
ulate); to 900 mm tall. Leaf blad-
es to 300 mm long; to 10 mm
wide. Spikelets 15-45 mm long
(including awns). Culm inter-
nodes pubescent; leaves and
sheaths usually scabrid; inflores-
cence very dense, spikelike, sometimes interrupted at the
base; lower glume to 2/3 the length of the upper glume (ex-
cluding awns), long awned; lemma articulation between the
apex of the lemma and the branching point of the awns;
column absent but lemma narrowed below the branching
point of the awns; awns three, subequal; callus 0. 5-1.5 mm
long, tip naked, narrowly to broadly rounded.

Flowering January to May. Moist heavy soils in shallow
depressions, on edges of pans and vleis, in old farmlands.
Locally common. Biome: Savanna. Throughout tropical
Africa. Indicator (of retrogression of veld). Resembles A.
hubbardiana, which has no articulation on the lemma, and
forms of A. congesta subsp. congesta, which have a distinct
column.

Description: De Winter 1965 (245), Melderis 1971
(116). Illustration: Muller 1984 (fig. 33), Melderis 1971
(tab. 3 1 ). Voucher: De Winter 2729, Giess & Muller 1 1 804.
PRECIS code 9902620-01700.

Aristida hubbardiana Schweick.

Annual: densely tufted (bran-
ched, erect to geniculate); to 500
mm tall. Leaf blades to 100 mm
long; to 2 mm wide. Spikelets
10-30 mm long (including awns).

Culm internodes glabrous; lower
leaf surface and sheaths smooth;
inflorescence dense and spike-
like; lower glume 2/3 as long
as to nearly equaling the upper glume, short-awned; lemma
narrowly elliptic, dorsally compressed, usually very scabrid
in the upper 2/3 with large prickles in rows, articulation ab-
sent; column absent but lemma narrowed into a short beak;
awns three, laterals well developed; callus tip naked,
swollen and rounded.

Flowering March to April. Damp calcareous, clayey
soils around vleis and seasonally flood depressions. Locally
common (but with a limited distribution). Biome: Savanna.
Endemic, possibly found in Angola. Resembles some forms
of A. adscensionis, which has the lemma dorsally compress-
ed, and A. hordeacea , which has a lemma articulation.

Description: De Winter 1965 (246). Voucher: Giess,
Volk & Bleissner 6405, Smith 3641. PRECIS code
9902620-01800.

Aristida junciformis Trin. & Rupr. subsp. galpinii

(Stapf) De Winter

(=A. galpinii Stapf) 1.

Perennial; long, oblique, thin-
ly rhizomatous and tufted (dense-
ly, erect); to 500 mm tall. Leaf
blades to 250 mm long; to 1 mm
wide. Spikelets 15-30 mm long
(including awns). Culms mainly
unbranched at the upper nodes; leaves mainly basal, form-
ing a dense basal tuft in which the culms remain hidden;

51

inflorescence contracted, branches erect or spreading but
not more than 45 degrees from the main axis; glumes usual-
ly very unequal, the lower 1/2-2/3 the length of the upper,
mucronate to shortly awned; lemma narrowly lanceolate,
articulation absent, lemma distinctly tapering towards a
short beak or very short column; awns three, laterals well
developed; callus tip naked, truncate or broadly rounded.

Flowering November to April. Shallow soils and
overgrazed areas on basalt or sandstone, rocky slopes of
very high mountains. Locally common. Biome: Grassland.
Endemic. Indicator (overgrazed and disturbed areas). This
subsp. occurs in very high mountainous sourveld. This
separates it from A. aequiglumis, which occurs mainly in
the bushveld, has a short, stout rhizome and glumes that are
usually subequal to equal.

Description: De Winter 1965 (266). Illustration: Chip-
pindall 1955 (fig. 278). Voucher: Killick 4471, Acocks
21992. PRECIS code 9902620-01900.

Aristida junciformis Trin. & Rupr. subsp. junciformis

Gongoni-steekgras, wire
grass'.

Perennial; stoutly rhizomatous
and tufted (densely, erect); to 900
mm tall. Leaf blades to 300 mm
long; to 3 mm wide. Spikelets
20-30 mm long (including awns).

Culms unbranched to branched at
some nodes; leaves mainly cauline or at least not densely
basal, culms always clearly visible for most of their lengths;
inflorescence contracted, dense to lax, branches erect to
spreading up to 45 degrees from the main axis; lower glume
up to 2/3 the length of the upper, awned; lemma narrowly
lanceolate, distinctly tapering into a beak or column, articu-
lation absent; awns three, laterals well developed; callus tip
naked, swollen, rounded to truncate.

Flowering November to May. Sandy, clayey, stony soils
or shallow soils on stony hillsides, in depressions where
water collects and in other damp places, along roadsides
and other disturbed ground. Common to locally dominant
(widely distributed). Biome: Fynbos, Savanna, and Grass-
land. The species in the broad sense occurs northwards to
East Africa. Domestic use (used for brooms), or indicator
(of mismanagement of veld), or weed (extremely tough
problem, pioneer grass). In this treatment De Winter’s
broad concept of the subspecies has been followed and
specimens that may be referrable to subsp. welwitschii may
be included here. Resembles A. transvaalensis , which is
branched at most nodes and with lateral awns either absent
or shorter and thinner than the central awn. Resembles A.
aequiglumis, which has a dense tuft of basal leaves, and A.
dasydesmis, which has the lower glume without an awn or
mucro.

Description: De Winter 1965 (266). Illustration: Chip-
pindall 1955 (fig. 274). Voucher: Strey & Schlieben 8524,
Smook 4651, De Winter 735. PRECIS code 9902620-
02000.

Aristida meridionalis Henr.

Langbeensteekgras.

Perennial; densely tufted; to
2000 mm tall. Leaf blades to 650
mm long; to 5 mm wide. Spike-
lets 35-50 mm long (including
awns). Lower culm internodes
glabrous or pubescent to woolly-
hairy; leaves flexible, some leaf
auricles with long woolly hairs; inflorescence oblong to
broadly oblong, symmetrical, large, to 800 mm long and ov-
er 200 mm wide, much branched; lower glume less than 2/3
the length of the upper glume; lemma articulation between
the apex of the lemma and the base of the column; column

Fig. 20.

8-20 mm long; awns three, subequal; callus tip naked,
deeply bifid.

Flowering November to May. Deep sandy to stony soils
in open areas, along roadsides and in moist areas around
vleis and damp depressions. Locally common to common.
Biome: Savanna. Angola, Zimbabwe, Mozambique,

northwards to Tanzania. Domestic use (occasionally used
for thatching), or pasture (only grazed when very young).
Similar to A. stipoides, which is annual. Resembles A.
spectabilis, which has the lower glume membranous for the
upper 1/3-2/3, and A. vestita, which has the lemma column
5-7 mm long.

Description: De Winter 1965 (284), Clayton et al.
1970-1982 (153). Voucher: Smook 4344. PRECIS code
9902620-02100.

Fig. 20. Aristida meridionalis

Aristida mollissima Pilg. subsp. argentea (Schweick.)

Meld.

( -A . argentea Schweick.) 1.

Perennial; densely tufted (e-
rect to geniculate); to 1000 mm
tall. Leaf blades to 400 mm long
to 4 mm wide. Spikelets 50-90
mm long (including awns). Lower
internodes of culms woolly to

52

densely tomentose; inflorescence narrow, lax, more or less
divarcately branched; lower glume to 2/3 the length of the
upper glume, awned; lemma articulation between the apex
of the lemma and the base of the column; column 13-30
mm long, twisted; awns three, slightly unequal to subequal;
callus 1. 5-3.0 mm long, tip naked, acuminate, pungent.

Flowering December to March. Light, sandy to stony
sandy soils in open areas, on mountain slopes or disturbed
areas. Locally common. Biome: Savanna. Zimbabwe,
Mozambique. Resembles A. stipitata subsp. graciliflora ,
which has the lower culm internodes glabrous or pubescent
but not woolly-hairy.

Description: De Winter 1965 (289), Melderis 1971.
Voucher: Ellis 3211. PRECIS code 9902620-02150.

Aristida mollissima Pilg. subsp. mollissima

Perennial; densely tufted (e-
rect); to 850 mm tall. Leaf blades
to 300 mm long; to 4 mm wide.

Spikelets 50-95 mm long (includ-
ing awns). Lower internodes of
culms woolly to densely tomen-
tose; inflorescence contracted,
spikelike, very dense; lower
glume to 2/3 the length of the
upper glume, awned; lemma articulation between the apex
of the lemma and the base of the column; column 16-30
mm long; awns three, subequal; callus 1 .5-3.0 mm long, tip
naked, acuminate, pungent.

Flowering December to May. Deep sandy soils especial-
ly red Kalahari sands. Locally common. Biome: Savanna.
Zimbabwe, Kenya. Resembles A. stipitata subsp. stipitata ,
which has the lower culm nodes glabrous or hairy but not
woolly.

Description: De Winter 1965 (287), Melderis 1971.
Voucher: Davidse 6071. PRECIS code 9902620-02200.

Aristida monticola Henr.

Perennial; long rhizomatous
and tufted (erect to geniculate); to
900 mm tall. Leaf blades to 120
mm long; to 2 mm wide. Spike-
lets 18-25 mm long (including
awns). Culms much branched;
lower glume longer than the up-
per glume, long tapering to an
acuminate apex (this often breaks
very early, giving a false idea of the length), no awn or mu-
cro; lemma without articulation, but a beak or short, twisted
column is present; awns three, two laterals shorter than cen-
tral; callus tip naked, rounded to truncate.

Flowering January to May. Moist and shady situations
such as stream banks and seepage areas on mountain slopes.
Locally common (in the Drakensberg). Biome: Grassland.
Endemic. Resembles A. transvaalensis and some forms of
A. junciformis. These two species have the lower glume
awned or mucronate.

Description: De Winter 1965 (265). Voucher: Killick
1382. PRECIS code 9902620-02300.

Aristida parvula (Nees) De Winter

(=Stipa parvula Nees) 1.

Annual; tufted (erect to
geniculate to semi-prostrate); to
400(-800) mm tall. Leaf blades to
120 mm long; to 2 mm wide.
Spikelets 1 8-22 mm long (includ-
ing awns). Glumes unequal, low-
er to 2/3 the length of the upper

glume (excluding awns), lower glume awned; lemma later-
ally compressed, articulation between the apex of the lem-
ma and the base of the column; column to 5 mm long, twist-
ed; awn solitary, usually bent; callus tip naked, broad,
truncate to rounded, with long hairs at the junction between
the lemma and the callus, which are 1/2 the length of the
lemma body.

Flowering August to October and January to May (in
Namibia). Sandy, stony or gravel soils on gravel plains,
along water courses, disturbed areas and rocky hillsides.
Locally common. Biome: Nama-Karoo and Desert.
Endemic. In the past this species has been included in Stipa
because of the single awn, but Stipa has a membranous
ligule. Detailed studies have shown it to be an Aristida with
the lateral awns missing.

Description: De Winter 1965 (242), Chippindall 1955
(290). Voucher: Oliver, Muller & Steenkamp 6620.
PRECIS code 9902620-02400.

Aristida pilgeri Henr.

Robust and coarse perennial;
densely tufted; to 1500 mm tall.

Leaf blades to 500 mm long;

0.9^4. 5 mm wide. Spikelets
10-26 mm long (including awns).

Lower internodes glabrous; in-
florescence with spikelet clusters
linear to obianceolate (including
awns), longest pedicel to 8.5 mm
long; lower glume 1/2-2/3 the length of the upper glume,
long-awned; lemma articulation between the apex of the
lemma and the branching point of the awns, sometimes ar-
ticulation represented only by a swollen line or colour
differentiation; column distinct, to 2.5 mm long; awns
three, slightly unequal; callus 0.5-1. 5 mm long, tip naked,
broadly rounded.

Flowering February to July. Calcareous or sandy, stony
soils in moist depressions, along edges of floodplains, on
river banks. Locally common. Biome: Savanna. Zimbabwe,
Zambia. Indicator (retrogression of veld). Resembles A.
sciurus, forms of A. junciformis subsp. junciformis and A.
canescens subsp. canescens, but all these taxa have no lem-
ma articulation.

Description: De Winter 1965 (293). Illustration: Muller
1984 (fig. 35). Voucher: Smith 3834. PRECIS code
9902620-02500.

Aristida recta Franch.

Perennial; occasionally short-
ly rhizomatous and tufted (erect);
to 500 mm tall (usually shorter).

Leaf blades to 200 mm long; to 1
mm wide. Spikelets 10-12 mm
long (including awns). Basal leaf
sheaths persistent, breaking up
into fibres and forming a dense
tuft at the base of the culms; in-
florescence contracted, branches erect or spreading less
than 45 degrees from the main axis; spikelets brownish-
purple to deep purple; lower glume to 2/3 the length of the
upper glume, awned; lemma articulation absent; column ab-
sent, a short beak may be present; awns three, subequal; cal-
lus very short, tip naked, large, swollen, rounded.

Flowering September to November. Damp ground
around vleis, and seepage areas, usually on slopes of sour
mountain grassland. Infrequent to locally common. Biome:
Grassland. Tropical Africa.

Description: De Winter 1965 (270), Clayton et al.
1970-1982 (145). Illustration: Chippindall 1955 (fig. 277).
Voucher: Kluge 1982. PRECIS code 9902620-02600.

Aristida rhiniochloa Hochst.

(=A. andoniensis Henr.) 1.

Skurwe steekgras.

Annual; tufted (erect); to 900
mm tall. Leaf blades to 200 mm
long; to 4 mm wide. Spikelets
35-50 mm long (including awns).

Plants usually very scabrid, oc-
casionally smooth; spikelets coarse; lower glume usually
slightly longer than the upper glume, awn 0.8-3. 5 mm long;
lemma linear, laterally compressed, articulation absent;
column absent; awns three, laterals well developed; callus
tip naked, broad, swollen, rounded.

Flowering January to May. Sand to sandy loam, stony
to heavier soils, sometimes over calcareous outcrops on dry
ground along rocky slopes, gravel plains and eroded areas.
Locally common. Biome: Savanna. Northwards to tropical
east and west Africa. Indicator (overgrazing, drought and
other disturbances).

Description: De Winter 1965 (250), Clayton et al.
1970-1982 (147). Illustration: Muller 1984 (fig. 36).
Voucher: Smook 4221, Volk 1251. PRECIS code
9902620-02700.

Aristida scabrivalvis Hack, subsp. contracta (De Winter)

Meld.

Annual; tufted (erect to genic-
ulate); to 850 mm tall. Leaf blad-
es to 300 mm long; to 3.2 mm
wide. Spikelets 10-25 mm long
(including awns). Inflorescence
robust, branchlets and pedicels
appressed, with spikelets densely
congested at the ends of the bran-
ches; spikelets slender and fine;
lower glume less than 3/4 the length of the upper glume (ex-
cluding awns), awn 0.8-3. 5 mm long; lemma linear, lateral-
ly compressed, articulation absent; column absent; awns
three, laterals well developed; callus tip naked, swollen,
rounded.

Flowering March to May. Shale, heavy basalt or sandy
soils in open and disturbed places on roadsides and
hillslopes. infrequent. Biome: Savanna and Grassland. The
species occurs northwards to east Africa. Distinguished
from subsp. scabrivalvis, which has a more delicate inflor-
escence with the branchlets spreading and spikelets distant
from one another at the ends of the branches. There appears
to be a complete gradation between specimens conforming
to the ‘typical’ subsp. contracta and specimens that have
been referred to as subsp. borumensis. Pending a more
detailed study, all the specimens are included in subsp. con-
tracta. Resembles A. bipartita , which is a perennial, and A.
effusa , which may have a lower glume awn to 0.8 mm.

Description: De Winter 1965 (255). Voucher: Smith
2361, Ellis 524, Smook 4236, 3114. PRECIS code
9902620-02800 .

Aristida scabrivalvis Hack, subsp. scabrivalvis

Pers steekgras.

Annual; tufted (erect to
geniculate); to 850 mm tall. Leaf
blades to 200 mm long; to 3.5 mm
wide. Spikelets 18-24 mm long
(including awns). Inflorescence
delicate, branchlets and pedicels
spreading, with a few spikelets
distant from one another at the ends of the branches; lower
glume less than 3/4 the length of the upper glume (exclud-
ing awns), awn 0.8-3. 5 mm long; lemma linear, laterally
compressed, articulation absent; column absent; awns three,
laterals well developed; callus tip naked, swollen, rounded.

NATIONAL BOTANICAL INSTITUTE
PRIVATE BAG X 101
PRETORIA 0001

REPUBLIC OF SOUTH AFRICA

53

Flowering January to May. Sandy, sandy loam, clays,
often over limestone, usually in disturbed places such as
roadsides and old lands. Locally common. Biome: Savanna.
Tropical Africa. Distinguished from subsp. contracta,
which has a more robust inflorescence with branchlets
appressed and spikelets clustered at the ends of the branch-
es. Resembles A. effusa, which may have a mucro or short
awn to 0.8 mm long, and A. bipartita, which is perennial.

Description: De Winter 1965 (255), Clayton et al.
1970-1982 (147). Voucher: Giess, Volk & Bleissner

6508. PRECIS code 9902620-02900.

Aristida sciurus Stapf

Tall three-awned grass.

Robust perennial; short rhizo-
matous and tufted (erect); to 1400
mm tall. Leaf blades to 800 mm
long; 2— 3(— 6) mm wide. Spikelets
25-30 mm long (including awns).

Culms 5-6 mm in diameter, un-
branched, lower internodes
woolly-hairy or glabrous; lower glume 1/2 the length of the
upper glume, mucro or awn absent; lemma articulation ab-
sent; lemma narrowed into a beak or very short twisted
column; awns three, subequal; callus tip small, naked,
swollen and rounded.

Flowering January to May. Moist sandy soils mainly in
mountain sourveld. Locally common. Biome: Grassland.
Endemic. Resembles A. pilgeri, which has a lemma articu-
lation, A. spectabilis, which has the lower glume membran-
ous for the upper 1/2-2/3, and A. canescens subsp.
canescens, which is a more slender plant with a culm
diameter of 1 .5-3.0 mm.

Description: De Winter 1965 (273). Voucher: Turner
133, Compton 30594. PRECIS code 9902620-03000.

Aristida spectabilis Hack.

Bergsteekgras.

Perennial; densely tufted; to
1750 mm tall. Leaf blades to 600
mm long; 4-5 mm wide. Spike-
lets 35-40 mm long (including
awns). Lower glume 1/2 the
length to nearly as long as the
upper glume, firm below, upper
1/3-2/3 membranous and often torn, mucro or awn absent;
lemma articulation present between the apex of the lemma
and the base of the column; column 4-6 mm long, twisted;
awns three, equal to subequal; callus tip naked, emarginate
to distinctly bifid.

Flowering February to April. Shallow and sandy soils
mainly derived from quartzite on stony, rocky mountain
slopes. Infrequent to locally common. Biome; Savanna.
Endemic. Resembles A. meridionalis, which has the lower
glume with only the extreme tip membranous, A. pilgeri,
which has the articulation between the apex of the column
and the branching point of the awns, and A. sciurus, which
has no articulation.

Description: De Winter 1965 (283). Voucher: Smook
4755. PRECIS code 9902620-03100.

Aristida stipitata Hack, subsp. graciliflora (Pilg.) Meld.

(=A. graciliflora Pilg.) 1;

( -A . stipitata Hack. var.
graciliflora (Pilg.) De Winter) 2.

Langnaaldsteekgras.

Slender perennial; loosely
tufted (erect); to 900 mm tall.

Leaf blades to 200 mm long; 2-3
mm wide. Spikelets 60-80 mm long (including awns).

54

Culms 1. 0-2.5 mm in diameter, lower intemodes glabrous
or pubescent, not woolly-hairy; inflorescence narrow, not
spikelike, sparse, branches lax, not closely appressed to the
main axis; lower glume to 2/3 the length of the upper glume,
awned; lemma articulation between the apex of the lemma
and base of the column; column present; awns three, laterals
well developed; callus 1. 5-3.0 mm long, tip naked,
acuminate, pungent.

Flowering November to June. Sandy or loamy soils in
rocky situations, seepage zones and disturbed ground. Com-
mon (widespread). Biome: Savanna. Zimbabwe, Zambia,
Mozambique. A polymorphic species which is so variable
that it is often difficult to distinguish between the subsp.
although the extreme variants are distinct. Resembles A.
mollissima subsp. argentea , which has the lower culm inter-
nodes woolly to densely tomentose.

Description: De Winter 1965 (290). Illustration: Muller
1984 (fig. 38). Voucher: Smook 4166, Tinley 609. PRECIS
code 9902620-03300.

Aristida stipitata Hack, subsp. robusta (Stent & Rattray)

Meld.

Robust perennial; tufted; to
1 500 mm tall. Leaf blades to 300
mm long; to 4 mm wide. Spike-
lets 70-90 mm long (including
awns). Plants robust, culms

2. 5- 4.0 mm in diameter; lower
intemodes glabrous or pubescent,
not woolly-hairy; inflorescence
narrow, not spikelike, branches lax and not closely appress-
ed to main axis, 200-350 mm long; lower glume to 3/4 the
length of the upper glume, awned; lemma articulation
between the apex of the lemma and the base of the column;
column present; awns three, laterals well developed; callus

1. 5- 3.0 mm long, tip naked, acuminate, pungent.

Flowering January to April. Deep, heavy sands, often in

disturbed areas. Locally common. Biome: Savanna.
Zimbabwe, Zambia. A polymorphic species which is so
variable that it is often difficult to distinguish between the
subspecies although the extreme variants are distinct.

Description: De Winter 1965 (290). Illustration: Muller
1984 (fig. 39). Voucher: Smith 1640. PRECIS code
9902620-03400.

Aristida stipitata Hack, subsp. spicata (De Winter)

Meld.

Slender perennial; loosely to
densely tufted (erect); to 600 mm
tall. Leaf blades to 150 mm long.

Spikelets 50-100 mm long (in-
cluding awns). Culms much bran-
ched at the upper nodes, lower
internodes glabrous or pubescent,
not woolly-hairy; inflorescence
narrow, dense, spikelike, branches closely appressed to the
main axis, sometimes interrupted towards the base, usually
to 150 mm long; lower glume to 2/3 the length of the upper
glume, awned; lemma articulation between the apex of the
lemma and the base of the column; column well developed;
awns three, laterals well developed; callus 1. 5-3.0 mm
long, tip naked, acuminate, pungent.

Flowering February to May. Deep sandy soils associated
with rocky outcrops. Locally common. Biome: Savanna.
Zambia. A polymorphic species which is so variable that
it is often difficult to distinguish between the subspecies
although the extreme variants are distinct.

Description: De Winter 1965 (290). Illustration: Muller
1984 (fig. 40). Voucher: Acocks 2159. PRECIS code
9902620-03500.

Aristida stipitata Hack, subsp. stipitata

Robust perennial; tufted (e-
rect); to 1 500 mm tall. Leaf blad-
es to 300 mm long; to 4 mm wide.

Spikelets 50-90 mm long (includ-
ing awns). Culms sparsely bran-
ched at the upper nodes, lower
internodes glabrous or pubescent,
not woolly-hairy; inflorescence
narrow, 1 50-300 mm long, dense,
spikelike, the branches closely appressed to the main axis,
sometimes interrupted towards the base; lower glume to 2/3
the length of the upper glume, awned; lemma articulation
between the apex of the lemma and the base of the column;
column present; awns three, laterals well developed; callus

1. 5-3.0 mm long, tip naked, acuminate, pungent.

Flowering December to April. Deep sandy or calcareous
soils. Locally common. Biome: Savanna. Zimbabwe,
Zambia. Domestic use (thatching), or pasture (pioneer grass
of little forage value). A polymorphic species which is so
variable that it is often difficult to distinguish between the
subspecies although the extreme variants are distinct.
Distinguished from A. mollissima subsp. mollissima, which
has the lower internodes woolly.

Description: De Winter 1965 (290). Illustration: Muller
1984 (fig. 37). Voucher: Smook 4310, De Winter 2280.
PRECIS code 9902620-03550.

Aristida stipoides Lam.

(=A. fontismagni
Schweick.) 1.

Annual; loosely tufted; to
1 500 mm tall. Leaf blades to 300
mm long; 3-5 mm wide. Spike-
lets 55-75 mm long (including
awns). Leaf auricles with long
woolly hairs; lower glume lan-
ceolate, 6-9 mm long, to 2/3 the length of the upper glume,
apex acute, mucro or awn absent; lemma articulation
between the apex of the lemma and the base of the column;
column 20-35 mm long; awns three, subequal; callus tip
naked, deeply bifid.

Flowering February to May. Damp sandy soils along
seasonal floodplains, dry river beds, rocky hillsides,
roadsides or old cultivated lands. Locally common. Biome:
Savanna. Zambia, west Africa, Ethiopia to Tanzania.
Resembles A. meridionalis, which is a robust perennial.

Description: De Winter 1965 (286). Illustration: Muller
1984 (fig. 41). Voucher: Smith 1954, Schweickerdt 2129.
PRECIS code 9902620-03600.

Aristida transvaalensis Henr.

Rock three-awns.

Perennial: densely tufted; to
700 mm tall. Leaf blades to 150
mm long. Spikelets 15-30 mm
long (including awns). Plant
branched at all the upper nodes;
lower glume (including awns) 2/3
the length of the upper glume,
awned; lemma without articulation; column of variable
length; central awn sometimes solitary, lateral awns absent
or weakly developed being very much shorter and more
slender; callus tip naked, swollen, obtuse to truncate.

Flowering December to May. Shallow soils in crevices
and pockets on dry rocky outcrops and hillsides. Common.
Biome: Savanna. Endemic. Resembles A. junciformis

55

subsp. junciformis, which has culms branched but not at
every node and lateral awns well developed, and A.
monticola , which has the lower glume mucronate but un-
awned.

Description: De Winter 1965 (263). Voucher: Wells
1866, Smook 3056. PRECIS code 9902620-03700.

Aristida vestita Thunb.

Harde steekgras, large woolly
three-awn.

Perennial; densely tufted; to
850 mm tall. Leaf blades to 240
mm long; to 4 mm wide. Spike-
lets 30-50 mm long (including
awns). Lower culm internodes
pubescent to woolly-hairy, upper
internodes pubescent to glabrous; inflorescence narrowly
oblong to narrowly elliptic, usually asymmetric, to 200 mm
long and 120 mm wide; lower glume to 2/3 the length of
the upper glume, awn or mucro absent; lemma articulation
between the lemma apex and the base of the column;
column 5-7 mm long; awns three, laterals well developed;
callus tip naked, distinctly bifid.

Flowering sporadically, but mostly November to May.
Dry sandy loam or black clay, limestone soils in stony and
rocky veld. Locally common to common. Biome: Savanna
and Nama-Karoo. Northwards to Tanzania. Resembles both
subspecies of A. diffusa and A. engleri, which have the
lower culm intemodes glabrous.

Description: De Winter 1965 (249). Voucher: Smook
3495. PRECIS code 9902620-03800.

Arrhenatherum P. Beauv.

Thorea Rouy, Thoreochloa Holub.

Perennial; caespitose. Culms 300-2000 mm high; herba-
ceous; unbranched above. Leaves without auricles. Leaf
blades linear; flat, or rolled (convolute). Ligule an
unfringed membrane (sometimes puberulent). Plants
bisexual, with bisexual spikelets.

Inflorescence paniculate-, open4, espatheate. Spikelet-
bearing axes persistent.

Spikelets 7-11 mm long ; compressed laterally; disarticu-
lating above the glumes (the florets falling together).
Rachilla prolonged beyond the uppermost female-fertile
floret. Callus short. Glumes two; very unequal ; about
equalling the spikelets to much exceeding the spikelets;
awnless; similar (membranous). Incomplete florets
proximal to the female-fertile florets, or both distal and
proximal to the female-fertile florets. Distal florets when
present merely underdeveloped. Proximal incomplete
florets 1 or rarely absent.

Female-fertile florets 1 (or rarely 2-4). Lemmas decid-
edly firmer than the glumes; 5-9 nerved; entire, or incised;
awnless, or awned. Awns when present 1 ; dorsal; non-genic-
ulate (usually short and slender); much shorter than the
body of the lemma to about as long as the body of the
lemma. Palea present; relatively long. Lodicules 2;
membranous; glabrous. Stamens 3. Ovary hairy. Fruit
small, or medium sized, or large; hilum long-linear; embryo
small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 4 species.
Europe, Mediterranean. Mesophytic to xerophytic; in open
habitats (dry grassland, edges of woods, disturbed ground).
Transvaal and Natal. 1 naturalized species.

Intergeneric hybrids with Avena.

References. 1. Chippindall. 1955. Gr. & Past. 2. Holub.
1980. FI. Europ.

Species treatment by T.M. Sokutu.

Arrhenatherum elatius (L.) Presl

Fig. 21. PI. 13.

Perennial; culms solitary;

500-1400 mm tall. Leaf blades
100-190 mm long; 2-5 mm wide.

Spikelets 7-1 1 mm long. Inflor-
escence usually open, sometimes
contracted, branches filiform,
conspicuously unequal; awn on
lower lemma always conspicuous
and well developed; upper lemma
with less well developed awn (var. bulbosum (Willd.)
Spenner) or as developed as the lower one (var. biaristatum
(Peterm.) Peterm.).

Flowering November to December. Meadows, disturbed
places, roadsides and gardens. Infrequent. Naturalized from
Europe. Biome: Grassland. Planted pasture. The species is
characterized by its conspicuously unequal inflorescence
branches, var. bulbosum (Willd.) Spenner by its bulbous,
corm-like base, var. biaristatum (Peterm.) Peterm. by the
absence of the above. Few specimens were examined, from
which it appears that we may not have records for var.
elatius (L.) Presl.

Description: Holub 1980 (5:216), Chippindall 1955
(81). Illustration: Chippindall 1955 (fig. 52). Voucher:
Huntley 271. PRECIS code 9902000-00100.

56

Arthraxon P. Beauv.

Alectoridia A. Rich., Batratherum Nees, Lasiolytrum
Steud., Lucaea Kunth, Pleuroplitis Trin.

Annual, or perennial; decumbent. Culms 100-1000 mm
high (often trailing); herbaceous; branched above, or un-
branched above. Leaf blades linear-lanceolate to ovate-
lanceolate; cordate. Ligule a fringed membrane (short).
Plants bisexual, with bisexual spikelets. The spikelets of

sexually distinct forms on the same plant (usually); overtly
heteromorphic, or homomorphic.

Inflorescence of spike-like main branches', digitate or
subdigitate (usually subdigitate, rarely a single raceme);
espatheate; not comprising 'partial inflorescences’ and
foliar organs (but the inflorescences terminal and/or
axillary). Spikelet-bearing axes spikes (rarely), or
‘racemes’ (slender); clustered; with very slender rachides\
disarticulating at the joints. ‘Articles’ without a basal
callus-knob.

Spikelets solitary (rarely), or in pairs; consistently in
Tong-and-short’ combinations, or not in distinct Tong-and-
short’ combinations; when paired occurring in pedicellate/
sessile combinations. Pedicels when present free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, or sterile, or vestigial, variable in form,
reduced to a sometimes microscopic pedicel, or totally
suppressed. Female-fertile spikelets 2-6.5 mm long; com-
pressed laterally; falling with the glumes. Glumes two;
more or less equal; awnless; very dissimilar (lower often
coriaceous, rounded on the back; upper less firm, laterally
compressed). Proximal incomplete florets /; epaleate;
sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire, or incised; awned. Awns 1;
median; dorsal; geniculate; much shorter than the body of
the lemma, to much longer than the body of the lemma.
Palea present, or absent; when present very reduced. Lodi-
cules 2; fleshy; glabrous. Stamens 2-3. Ovary glabrous.
Fruit fusiform, or ellipsoid; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 7 species. Tropical
Africa, Madagascar, Mauritius, Indomalayan region to
Japan. Helophytic to mesophytic; in shade, or in open
habitats; glycophytic. Transvaal. 1 indigenous species.

References. 1. Van Welzen. 1981. Blumea 27: 255. 2.
Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Arthraxon lanceolatus (Roxb.) Hochst. var. lanceolatus

Fig. 22. PI. 14.

(=A. prionodes (Steud.)

Dandy) 1.

Perennial; trailing; to 600 mm
tall. Leaf blades 20-70 mm long;

4—20 mm wide (margins with
tubercle-based hairs). Spikelets
(sessile) 5. 0-6. 5 mm long (pedi-
cellate shorter). Lower glume
spiny on keels; female-fertile lemma awned from near base.

Flowering October to April. Riverbanks. Rare and con-
servation status not known. Biome: Forest. Eastern Africa
to India and tropical Asia.

Description: Chippindall 1955 (455), Clayton et al.
1970-1982 (741). Voucher: Stevenson-Hamilton s.n.
PRECIS code 9900670- 00050.

Arundinella Raddi

Acratherum Link, Brandtia Kunth, Calamochloe
Reichenb., Goldbachia Trin., Riedelia Kunth, Thysanachne
Presl.

Annual, or perennial: mostly with tough, erect culms.
Culms 300-1500 mm high; herbaceous; branched above, or
unbranched above. Leaf blades linear; flat, or rolled. Ligule
a fringed membrane (narrow). Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant (i.e., with reduced, sterile spikelets), or
all alike in sexuality.

57

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs; consistently in ‘long-and-
short’ combinations, or not in distinct ‘long-and-short’
combinations. Female-fertile spikelets 1.5-8 mm long;
compressed laterally ; disarticulating above the glumes.
Hairy callus present. Glumes two; very unequal; awned, or
awnless; very dissimilar to similar (membranous to papery,
G1 acute to mucronate, G2 often caudate). Lower glume 3
nerved. Proximal incomplete florets present or rarely
absent, these when present 1; paleate, palea fully developed
(narrow, two keeled); male.

Female-fertile florets I ( rarely 2). Lemmas similar in
texture to the glumes, or decidedly firmer than the glumes
(membranous to thinly coriaceous); not becoming indu-
rated; hairless (scabrid or scabridulous); having the margins
lying flat and exposed on the palea, or having the margins
tucked in onto the palea; with a clear germination flap ; 1-7
nerved; entire, or incised; awnless, or awned. Awns 1
(usually), or 3; median, or median and lateral (via capillary
bristles from the lobes). The median awn different in form
from the laterals (when laterals present); from the sinus;
geniculate; much shorter than the body of the lemma, to
much longer than the body of the lemma. Palea present
(sometimes auriculate at the base, but not keel-winged).

Lodicules 2; fleshy; glabrous. Stamens 3. Ovary glabrous.
Fruit small; hilum short; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional, or unconventional. Organization
of PCR tissue when unconventional Arundinella type.
Biochemical type NADP-ME (A. nepalensis)\ XyMS-.
PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 7, 10, 12, and 14. Panicoideae; Panicodae;
Arundinelleae. 55 species. In warm regions. Helophytic to
mesophytic; in open habitats (marshy places, riverbanks
and rocky slopes); glycophytic. Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 1 in-
digenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by H.M. Anderson.

Arundinella nepalensis Trin.

Fig. 23. PI. 15.

River grass, beesgras, rietgras.

Perennial; tufted; 900-1500
mm tall. Leaf blades 80-300 mm
long; 3-10 mm wide. Spikelets
4-6 mm long. Rhizome creeping,
often covered with short scale-
like leaves, resulting in a plaited
look; panicle dense, 120-300 mm
long; spikelets usually in pairs, brown, sometimes tinged
with green or purple; glumes unequal and acute; lower lem-
ma with a truncate, hairy callus, awns 3-6 mm long.

Flowering December to March. Vleis, riverbanks and
moist grasslands. Locally common. Biome: Savanna and
Grassland. Tropical east Africa to Asia. Domestic use
(thatching), or pasture (natural).

Description: Chippindall 1955 (275). Illustration: Chip-
pindall 1955 (fig. 247). Voucher: Smook 4999. PRECIS
code 9901730-00100.

Arundo L.

Amphidonax Nees, Donacium Fries, Donax P. Beauv.,
Eudonax Fries, Scolochloa Mert. & Koch.

Perennial ( mostly reeds with long canes)\ long-rhizoma-
tous. Culms 2000-6000 mm high, or not applicable
(occasionally pendant, from cliffs); woody and persistent;
branched above (main stems dominant). Leaf blades linear-
lanceolate to lanceolate; flat. Ligule a fringed membrane
(short).

Inflorescence paniculate (plumose); open; espatheate.
Spikelet-bearing axes persistent.

Spikelets 12-18 mm long; compressed laterally; disar-
ticulating above the glumes. Glumes two; more or less
equal; about equalling the spikelets; awnless; similar
(membranous). All florets female-fertile, or with distal in-
complete florets, these merely underdeveloped; proximal
incomplete florets absent.

Female-fertile florets 2-7 . Lemmas less firm than the
glumes, or similar in texture to the glumes (membranous
or hyaline); hairy (villous on the back); 3-9 nerved; entire,
or incised; awnless to awned. Awns when present 1, or 3;
median, or median and lateral. The median awn similar in
form to the laterals (when laterals present); from the sinus;
non-geniculate; much shorter than the body of the lemma.
Palea present; 2-nerved. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Hilum short; pericarp fused;
embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x - 12. Arundinoideae; Arundineae. 3 species.
Tropical and temperate. Helophytic to mesophytic.

58

Botswana, Transvaal, Orange Free State, Natal, and Cape
Province. 1 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Fig. 24. Arundo donax

Arundo donax L.

Fig. 24. PI. 16.

Giant reed, Spaanseriet.

Perennial; rhizomatous; to
3000 mm tall. Leaf blades to 700
mm long; to 80 mm wide. Spike-
lets 8-15 mm long. Robust, not
tufted; leaves deciduous at base
of blade; blades rounded or
caudate at base, tips not sharp;
ligule with fringing hairs shorter than membranous base; in-
florescence 300-600 mm long, compact with ascending
branches; lemmas with long hairs on the back.

Flowering February, or April. Moist disturbed places.
Infrequent. Naturalized from warm regions of the Old
World; escaped from cultivation. Widely cultivated
worldwide. Barrier and ornamental. Easily mistaken for
Phragmites. It may be distinguished most readily by the
very large compact inflorescence. However it seldom
flowers in the highveld, and then is best distinguished by
the combination of leaf characters.

Description; Chippindall 1955 (229). Illustration; Chip-
pindall 1955 (fig. 203). Voucher: McClean 536. PRECIS
code 9902130-00100.

Avena L.

Anelytrum Hack., Preissia Opiz.

Annual ; caespitose to decumbent. Culms 200-2000 mm
high; herbaceous; unbranched above. Leaf blades linear;
flat (usually), or rolled (rarely convolute). Ligule an
unfringed membrane .

Inflorescence paniculate ; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets not in distinct ‘long-and-short’ combinations;
10-45 mm long: compressed laterally; disarticulating above
the glumes, or not disarticulating (cultivated forms).
Glumes two (lanceolate); more or less equal; about
equalling the spikelets, or much exceeding the spikelets
(rarely shorter ); awnless; similar (usually chaffy). Incom-
plete florets distal to the female-fertile florets ; proximal in-
complete florets absent.

Female-fertile florets (l-)2-6. Lemmas similar in
texture to the glumes (rarely), or decidedly firmer than the
glumes (usually coriaceous to crustaceous); 5-9 nerved;
incised; awnless, or awned. Awns when present 1, or 3;
median, or median and lateral. The median awn different
in form from the laterals (when laterals present); dorsal;
geniculate; much longer than the body of the lemma. Palea
present; relatively long, or conspicuous but relatively short,
or very reduced (but large). Lodicules 2; membranous;
glabrous. Stamens 3. Ovary hairy. Fruit medium sized;
hilum long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, .t = 7. Pooideae; Poodae; Aveneae. 27 species.
Europe, Mediterranean, North Africa, western Asia. Meso-
phytic, or xerophytic; in open habitats (mostly in weedy
places); glycophytic. Transvaal, Orange Free State, Natal,
Lesotho, and Cape Province. Naturalized species (4), cult-
ivated ( 1 ).

Intergeneric hybrids with Arrhenatherum.

References. 1. Rocha Afonso. 1980. FI. Europ.

Species treatment by T.M. Sokutu.

1 (0). Lemmas glabrous or sparsely hairy at the base; central
awns straight or weakly geniculate, sometimes

absent 2

Lemmas densely hairy up to awn insertion; central

awns strongly geniculate 3

2(1). Awns with a distinct column A. sativa

Awns lacking a distinct column .... A. byzantina

3(1). Lemma apices awned, awns 4-8 mm long

A. barbata

Lemma apices awnless 4

4(3). Lemma teeth 1. 0-1.5 mm long A. fatua

Lemma teeth to 0.5 mm long A. sterilis

Avena barbata Brot.

PI. 17.

Annual; culms solitary or tuft-
ed; 300-1300 mm tall. Leaf
blades 70-300 mm long; 3-10
mm wide. Spikelets 18-26 mm
long. Leaf blades soft and ex-
panded; spikelets numerous; lem-
mas hairy in the lower half, apices
awned.

Flowering August to Decem-
ber. Waste and/or disturbed places, roadsides on sandy soil.
Locally dominant (disturbed areas in the Cape). Invader
from Europe. Biome: Fynbos and Savanna. Ornamental,
weed, and domestic use (dried flower arrangements).
Variable in plant height and spikelet size, easily
distinguished by awns on its lemma apices.

Description: Adams. & Salter 1950 (69), Rocha Alfonso
1980 (5:206), Stapf 1898-1900 (480), Chippindall 1955
(81). Voucher: Compton 15370. PRECIS code 9901950-
00100.

59

Avena byzantina K. Koch.

Annual; loosely tufted; 500-
1600 mm tall. Leaf blades 1 50—

500 mm long; 3-9 mm wide.

Spikelets 25-35 mm long. Lem-
mas glabrous or sometimes
sparsely hairy at the base; awn
straight or weakly geniculate,
without a distinct column.

Flowering September to Dec-
ember. Waste and disturbed areas and roadsides. Indeter-
minate. Infrequent. Naturalized from Europe. Biome: Fyn-
bos, Savanna and Grassland. Europe. Potential ornamental,
or weed. Very similar to A. sativa and sometimes not easily
distinguished, as the only difference is the distinct column
in the awn of A. sativa. This species seem to grow together
with populations of A. sativa.

Description: Rocha Alfonso J 980 (5:207). Voucher:
Esterhuysen 609. PRECIS code 9901950-00150.

Avena fatua L.

Fig. 25.

Annual; culms solitary or
loosely tufted; 250-700 mm tall.

Leaf blades 50-280 mm long; 3-8
mm wide. Spikelets 18-32 mm
long. Spikelets usually brownish
in colour, especially on the hairs;
lemma apices with teeth usually
1 .0—1 .5 mm long.

Flowering August to Novem-
ber. Disturbed and waste places, roadsides on sandy soil.
Common. Invader from Europe. Biome: Fynbos and Sa-
vanna. Europe, north Africa, western and central Asia,
introduced to Kenya and Zimbabwe. Weed. Not always
easy to distinguish from A. sterilis , but the teeth are usually
shorter in the latter species.

Description: Adams. & Salter 1950 (69), Stapf
1898-1900 (479), Chippindall 1955 (81), Clayton et al.
1970-1982 (82). Illustration: Clayton et al. 1970-1982 (fig.
28). Voucher: Gibbs Russell 3942. PRECIS code
9901950-00200.

Avena sativa L.

Oats.

Annual; culms solitary or
loosely tufted; 350-1 500 mm tall.

Leaf blades 100-400 mm long;

3- 9 mm wide. Spikelets 17-35
mm long. Lemma sparsely hairy
or glabrous, apices emarginate;
awn with a distinct column,
almost straight, sometimes absent.

Flowering September to November. Waste places,
disturbed areas, roadsides. Common (in its habitats). Inva-
der; from Europe. Biome: Fynbos, Savanna and Grassland.
Food and drink (cereal crop). This species can be easily
confused with A. byzantina K. Koch, but can be
distinguished by the distinct column of the awns. Three
subspecies can be recognized: subsp. macrantha (lemma
sparsely hairy), subsp. sativa (lemma glabrous, unawned)
and subsp. praegravis (lemma glabrous, awned).

Description: Adams. & Salter 1950 (69), Stapf
1898-1900 (478), Chippindall 1955 (81). Voucher: Crook
802. PRECIS code 9901950-00300.

Avena sterilis L.

Annual; culms solitary or
loosely tufted; 500-1450 mm tall.

Leaf blades 200-500 mm long;

4- 15 mm wide. Spikelets 20-46
mm long. Lemmas with short
rigid hairs on proximal 2/3,
apices with teeth to 0.5 mm long,
with long hairs on the lower 1/2.

Flowering September to Nov-
ember. Waste places, disturbed areas, mainly on sandy soil.
Common (in its habitats). Invader from Europe. Biome:
Fynbos. Weed. The distinguishing characters do not always
occur together. Sometimes the rigid proximal hairs are
missing, but the short teeth are always present. Very much
like A. fatua. Two subspecies, subsp. sterilis (ligule 5-6
mm long, florets up to 5) and subsp. ludoviciana (ligule 3
mm long, florets up to 3). Some specimens of this species
have previously been wrongly referred to as A. strigosa.

Description: Rocha Alfonso 1980 (5:208), Stapf
1898-1900 (479), Clayton et al. 1970-1982 (84). Voucher:
Bolus 24908. PRECIS code 9901950-00400.

Axonopus P. Beauv.

Anastrophus Schlecht., Cabrera Lag., Lappogopsis
Steud.

Annual (rarely), or perennial; long-stoloniferous
(sometimes mat-forming), or caespitose. Culms 150-1000
mm high (or more?); herbaceous. Leaf blades linear-
lanceolate to ovate-lanceolate; flat, or folded. Ligule an
unfringed membrane.

Inflorescence of spike-like main branches', digitate or
subdigitate (rarely), or non-digitate; espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary; biseriate; adaxial', compressed dorsi-
ventrally; falling with the glumes. Glumes one per spikelet
(membranous)', awnless. Proximal incomplete florets 1;
epaleate; sterile.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes; smooth to striate; becoming indurated, or not
becoming indurated (papery to crustaceous); hairless;

60

having the margins tucked in onto the palea; with a clear
germination flap; 4 nerved; entire; awnless. Palea present;
relatively long. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small, ellipsoid; hilum short; embryo
large.

Photosynthetic pathway. C4; NADP-ME (1 species);
XyMS-. PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Panicodae; Paniceae. 110
species. Tropical South America. Helophytic to meso-
phytic; in open habitats (savanna, forest clearings, moist
and weedy places); glycophytic. Transvaal, Swaziland,
Natal and Cape Province. 1 naturalized species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

Fig. 26. Axonopus affinis

Axonopus affinis Chase

Fig. 26. PI. 18.

(-A. compressus sensu
Chippind., non (Swartz)

Beauv.) 1.

Carpet grass.

Perennial; stoloniferous; 250-
600 mm tall. Leaf blades to 200
mm long; to 8 mm wide. Spike-
lets 2 mm long; 1 mm wide.

Strongly stoloniferous, forming dense swards; culm nodes
glabrous; leaf blades rounded at tip; female-fertile (upper)
lemma as long as spikelet.

Flowering December to May. Moist and disturbed
ground. Infrequent. Naturalized from tropical America.
Biome: Savanna. Global. Cultivated pasture and erosion
control. The rounded tip of the leaf blade distinguishes this
from Digitaria species with a similar inflorescence.

Description: Clayton et al. 1970-1982 (446)..

Illustration: Chippindall & Crook 1976 (130). Voucher:
Smook 5526. PRECIS code 9901050-00100.

Bambusa Schreber

Arundarbor Kuntze, Bonia Balansa, Criciuma
Soderstrom & Londoho, Dendrocalamopsis (Chia & Fung)
Keng f., Eremocaulon Soderstrom & Londoho, Guadua
Kunth, Ischurochloa Buse, Leleba Nakai, Lingnania
McClure, Tetragonocalamus Nakai — cf. Clayton and
Renvoize (1986). Soderstrom and Ellis (1987) refer
Criciuma , Eremocaulon and Guadua to their subtribe
Guaduinae, along with Olmeca , and place
Tetragonocalamus in the Arundinariinae, but revised
generic descriptions adequate for the present purpose are
not available.

Perennial. Culms 5000-35000 mm high (rarely to only
2000 cm)\ woody and persistent. Culms reaching 150 mm
in diameter (in ‘large’ species). Culms branched above.
Leaf blades pseudopetiolate; disarticulating from the
sheaths. Ligule an unfringed membrane to a fringed
membrane.

Inflorescence paniculate-, spatheate (with or without
foliage leaves). Spikelet-bearing axes persistent.

Spikelets 10-80 mm long; compressed laterally to not
noticeably compressed; disarticulating above the glumes.
Glumes two; more or less equal; decidedly shorter than the
adjacent lemmas; awnless; similar. Proximal incomplete
florets present or absent, these 1-3 (? — fewer than 4) when
present; sterile.

Female-fertile florets 1-20 (‘many’). Lemmas entire;
awnless; 9-22 nerved. Palea present; relatively long; with
several nerves (about 6-16). Lodicules 3; membranous;
ciliate. Stamens 6. Ovary hairy; with a conspicuous apical
appendage. The appendage broadly conical, fleshy. Stigmas
3 (usually?). Hilum long-linear; embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells; with fusoids. Midrib vascularization complex.

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Bambusodae; Bambuseae.
About 120 species. Tropical and subtropical Asia, Africa,
America. Transvaal, Natal, and Cape Province. 1
naturalized species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

61

Bambusa balcooa Roxb. ex Roxb.

PI. 19.

Bamboo; rhizomatous;

15000-21000 mm tall. Leaf
blades to 150 mm long; to 40 mm
wide. Spikelets 7-16 mm long.

Grows in dense clumps, the culms
arching gracefully at the top.

Flowering rare and sporadic.

Streambanks and forest margins.

Naturalized from India.

Occasionally escaped from cultivation, and distinguished
from indigenous bamboos by its great height.

Description: Chippindall 1955 (31). Voucher: Forbes &
MacClean 26173. PRECIS code 9904710-00100.

Bewsia Goossens

Perennial; caespitose (with short, creeping rhizomes).
Culms 260-930 mm high; herbaceous; unbranched above.
Leaf blades linear to linear-lanceolate; flat, or rolled (the
margins becoming involute under water stress). Ligule an
unfringed membrane (minutely ciliolate only). The spikelets
all alike in sexuality.

Inflorescence of spike-like main branches (oppressed to
the central axis)', with about 10-15 primary inflorescence
branches', espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; 5.5-9 mm long; compressed
laterally (strongly so); disarticulating above the glumes; not
disarticulating between the florets. Glumes two; more or
less equal; about equalling the spikelets (a little shorter to
a little longer); awnless; similar. Incomplete florets distal
to the female-fertile florets, merely underdeveloped;
proximal incomplete florets absent.

Female-fertile florets 2-6. Lemmas similar in texture to
the glumes (membranous); without a germination flap; 3
nerved; entire, or incised (minutely notched); awned. Awns
1; median; dorsal; non-geniculate; much shorter than the
body of the lemma to about as long as the body of the
lemma (1-4 mm). Palea present; relatively long. Lodicules
2; fleshy (long, narrow); glabrous. Stamens 3. Ovary
glabrous. Fruit small (about 2 mm long); linear (to oblong);
pericarp fused.

Photosynthetic pathway and related features. C4;
XyMS-t- (the ms thick-walled, sometimes double). PCR
sheath outlines even. PCR sheath extensions absent. PCR
cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. Southern tropical and
South Africa. Mesophytic (in grassveld, often on sandy
soil); in open habitats. Namibia, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 1 in-
digenous species.

References. 1. Clayton. 1970. FWTA. 2. Clayton et al.

1974. FTEA. 3. Goossens. 1941. SA Jl. Sci. 37: 183-191.

Species treatment by M. Koekemoer.

Fig. 27. PI. 20.

Bewsia biflora (Hack.) Goossens

(=Diplachne biflora Hack, ex
Schinz) 1.

Blousaadgras.

Perennial; shortly rhizomatous
and tufted; 260-930 mm tall. Leaf
blades 100-400 mm long; 1-5
mm wide. Spikelets 5. 5-9.0 mm

long. Spikelets 2— 4-flowered; lemma dorsally awned; awn
1-8 mm long, arising 1-2 mm from the tip.

Flowering November to April. Rocky hillsides or plains,

mixed bushveld to open flats. Common. Biome: Savanna
and Grassland. Tropical east Africa. Ornamental (in grass
gardens). The dorsally awned lemmas in Bewsia differ from
other Eragrostideae, where the awn arises from the tip of
the lemma.

Description: Chippindall & Crook 1976 (176), Goossens
1941 (183), Stapf 1898-1900 (593), Chippindall 1955
(121), Clayton et al. 1970-1982 (286). Illustration: Chip-
pindall 1955 (fig. 92), Clayton et al. 1970-1982 (fig. 78).
Voucher: Smook 946. PRECIS code 9903442-00100.

62

Bothriochloa Kuntze

Amphilophis Nash, Gymnandropogon (Nees) Duthie.
Sometimes included in Dichanthium.

Perennial; long-rhizomatous, or long-stoloniferous, or
caespitose, or decumbent. Culms 150-2000 mm high; her-
baceous; branched above, or unbranched above. Leaf blades
linear; flat. Ligule an unfringed membrane to a fringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets of sexually distinct forms on the same plant ;

overtly heteromorphic (the pedicellate spikelets smaller,
awnless), or homomorphic; all in heterogamous combina-
tions.

Inflorescence of spike-like main branches (many-jointed
‘racemes' ), or paniculate (rarely: the lower ‘racemes'
sometimes branched again at the base)', digitate or subdigi-
tate (the racemes often almost palmate, towards the culm
tips), or non-digitate; spatheate, or espatheate; not com-
prising ‘partial inflorescences’ and foliar organs. Spikelet-
bearing axes ‘racemes’ (with many — more than 8 —
sessile spikelets); solitary, or paired, or clustered; with very
slender rachides\ disarticulating at the joints. The pedicels
and internodes of the rachis with a longitudinal, translucent
furrow (often villous). ‘Articles’ without a basal callus-
knob.

Spikelets in pairs (with a terminal triplet); consistently
in ‘long-and-short’ combinations; these pedicellate/sessile.
Pedicels free of the rachis. The sessile spikelets hermaphro-
dite. The pedicellate spikelets male-only, or sterile,
awnless. Female-fertile spikelets compressed dorsiven-
trally; falling with the glumes (and with the joint). Glumes
two; more or less equal; awnless; very dissimilar (lower
bicarinate, often with a pit on the back; upper narrower,
naviculate). Proximal incomplete florets I ; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (reduced to a hyaline stipe); entire; awned. Awns
1; median; apical; geniculate; much longer than the body
of the lemma. Palea present, or absent; when present very
reduced. Lodicules 2; fleshy; glabrous. Stamens 1-3. Ovary
glabrous. Fruit small; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 35 species. Warm regions. Meso-
phytic; in open habitats (grassy places); glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, and Cape Province. 3 indigenous species.

Intergeneric hybrids with Capillipedium, Dichanthium.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Lower glumes lacking a pit, with a few long sparse

hairs on the lower half B. radicans

Glumes with a deep pit, usually glabrous 2

2(1). Inflorescence axis longer than the racemes; racemes
more than 20; hairs along sides of pedicels and on

callus usually shorter than 1 mm B. bladhii

Inflorescence axis shorter than the racemes; racemes
3-20; hairs along sides of pedicels and on callus
1-3 mm long B. insculpta

Bothriochloa bladhii (Retz.) S.T. Blake

(=B. glabra (Roxb.) A.

Camus) 1; ( =B . insculpta (A.

Rich.) A. Camus var. vegetior
(Hack.) C.E. Hubb.) 1.

Blouklosgras, purple plume
grass.

Perennial; tufted; 600-1800
mm tall. Leaf blades 100-550 mm long; 2-12 mm wide.
Spikelets (sessile) 3-4 mm long. Inflorescence axis longer
than racemes; racemes more than 20, pedicel and callus
hairs usually shorter than 1 mm; lower glumes pitted.

Flowering December to June. Riverbanks and vleis.
Common. Biome; Savanna and Nama-Karoo. Old World
tropics. Hybridizes readily with related species.

Description: Chippindall 1955 (483), Clayton et al.
1970-1982 (719). Voucher: Schoenfelder 95. PRECIS code
9900630-00100.

63

Bothriochloa insculpta (A. Rich.) A. Camus

Fig. 28. PI. 21.

(=B. pertusa auctt., non (L.)

A. Camus) 1.

Pinhole grass, klosgras,
stippelgras.

Perennial; sometimes stolon-
iferous; to 1500 mm tall. Leaf
blades 40-300 mm long; 2-8 mm
wide. Spikelets (sessile) 4. 5-5.0 mm long. Inflorescence
axis shorter than racemes; racemes 3-20, pedicel callus
hairs 1-3 mm long; lower glumes pitted.

Flowering October to June. Grassland and hillsides,
often in overgrazed places. Common. Biome: Savanna and
Grassland. Throughout Africa.

Description: Chippindall 1955 (483), Clayton et al.
1970-1982 (720). Voucher: Liebenberg 4382. PRECIS
code 9900630-00150.

Bothriochloa radicans (Lehm.) A. Camus

Perennial; often stoloniferous;

300-700 mm tall. Leaf blades
60-200 mm long; 2-6 mm wide.

Spikelets (sessile) 3-5 mm long.

Inflorescence axes shorter than
racemes, racemes 5-16, lower
glume of sessile spikelets
sparsely hairy, glumes not pitted.

Flowering October to April.

Rocky hillsides. Common. Biome: Savanna and Grassland.
North to Ethiopia, introduced to tropical America.

Description: Chippindall 1955 (482), Clayton et al.
1970-1982 (721). Illustration: Chippindall 1955 (fig. 395).
Voucher: De Winter 2824. PRECIS code 9900630-00500.

Brachiaria (Trin.) Griseb.

Pseudobrachiaria Launert.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 70-2000 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear to ovate-lanceolate; flat, or folded, or
rolled. Ligule an unfringed membrane, or a fringed
membrane, or a fringe of hairs. Plants bisexual, with
bisexual spikelets.

Inflorescence of spike-like main branches ; espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs (or occasionally in
fascicles); biseriate; not in distinct ‘long-and-short’ combi-
nations; broadly elliptic, plump, more or less obtuse;
awnless, muticous\ adaxial (or orientation ambiguous)-, not
noticeably compressed (rarely), or compressed dorsiven-
trally, falling with the glumes. Glumes two ; very unequal,
or more or less equal (rarely); awnless; very dissimilar (the
upper similar to LI). Proximal incomplete florets 1 ; paleate,
or epaleate; male, or sterile.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (crustaceous to subcoriaceous); striate, or
rugose (and rarely smooth ); becoming indurated, or not
becoming indurated; hairless (smooth or tuberculate);
having the margins tucked in onto the palea; with a clear
germination flap; 3-5 nerved; entire; awnless, pointed or
apiculate but not mucronate. Palea present (the tip not
reflexed); relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Photosynthetic pathway. C4; PCK (12 species, including
Pseudobrachiaria)', XyMS+. PCR cell chloroplasts
centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 7 and 9. Panicoideae; Panicodae; Paniceae.
About 100 species. Warm regions. Helophytic, or meso-
phytic, or xerophytic; mostly in open habitats, or in shade
(diverse habitats, from semidesert to swamps). Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho and Cape Province. Indigenous species (19),
naturalized species (1).

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA. 3. Scholz. 1978. Willdenowia
8:383.

Species treatment by M. Koekemoer.

64

1(0). Inflorescences with pedicels of unequal lengths, at
least some spikelets borne on long slender pedicels
exceeding 2 mm; spikelets mostly more than their

own length apart 2

Inflorescences with pedicels of equal lengths,
spikelets almost sessile or borne on pedicels shorter
than 1 mm; spikelets overlapping for at least 1/4

their length 4

2(1). Leaf blades with a cordate, pseudopetiolate base;
racemes usually fewer than 5, lax and widely
spaced; a coastal, bushland or forest grass that

rarely occurs inland B. chusqueoides

Leaf blades linear or rounded at the base; racemes
more than 5, stiff and arranged in a broadly ovate

panicle; wooded grassland species 3

3(2). Spikelets 2.0-3. 5 mm long; upper lemma finely

rugose B. deflexa

Spikelets 3. 5-4. 2 mm long; upper lemma coarsely

rugose B. grossa

4(1). Racemes not conspicuously secund; spikelets
clustered; racemes mostly appressed to the central

axis or spreading slightly 5

Racemes very conspicuously secund; spikelets
usually arranged in one or two rows (rarely four);

racemes rarely appressed 6

5(4). Plant covered with soft, short, velvety, white hairs;

rachilla extension short B. glomerata

Plant with very dense, long, golden hairs all over;

rachilla extension absent B. psammophila

6(4). Lower glumes distinctly darker at the base, tightly
clasping a short intemode that separates the lower
and upper glume; lower glumes very variable but
at least some are long acuminate and 1/2-2/3 the
spikelet length; upper glume and lower lemma also

acuminate B. nigropedata

Lower glumes rarely with a distinctive colour
difference, not clasping the short internode between
the glumes; lower glumes truncate, rounded (or
acute when scale-like), to 2/3 the spikelet length;
upper glume and lower lemma usually truncate or

rounded, but sometimes mucronate 7

7(6). Lower glume 2/3 the spikelet length, 7-9-nerved,
usually glabrous; cross-veins present between

nerves of upper glume and lower lemma 8

Lower glume less than half the spikelet length, mostly
1-3-nerved (5-nerved in B. xantholeuca, 7-nerved
in B. brizantha)', cross-veins absent on upper glume

and lower lemma 11

8(7). Upper and lower glume separated by a short

internode; spikelets 4. 5-7.0 mm long 9

Upper and lower glume not separated; spikelets

2. 8^1.5 mm long 10

9(8). Racemes 4-12; plants densely tufted with a creeping

rhizome; culms erect B. dictyoneura

Racemes 1-4; plants stoloniferous and rooting at the
nodes; culms often decumbent . . B. humidicola

10(8). Leaf blades filiform, usually up to half the length

of the culms, rarely hairy B. subulifolia

Leaf blades flat or sometimes convolute, usually
much less than half the length of the culms,

usually hairy B. bovonei

11(7). Lower glume 7-nerved, separated from the upper
glume by a short internode; spikelets 4-6 mm
long, arranged in a single row (occasionally two

rows near the base) B. brizantha

Lower glume 1-5-nerved, not separated from the
upper glume; spikelets 1. 5-5.0 mm long,

arranged in one, two or four rows 12

12(11). Lower glume 3-5-nerved, 1/2 the spikelet length,

not scale-like 13

Lower glume 1 -nerved or unobtrusively nerved,
usually 1/4 the spikelet length (rarely to 1/2 the
length), scale-like 15

13(12). Plants annual; spikelets 2. 7-4.0 mm long; leaf
blades flat, broadly linear to narrowly lanceolate,

4—10 mm wide, velvety pubescent

B. xantholeuca

Plants perennial; spikelets 4—5 mm long; leaf blades
convolute and wiry, 1-3 mm wide, glabrous . .

14

14(13). Spikelets glabrous; occurs in Namibia and

Botswana B. dura var. dura

Spikelets densely pilose; known only from Witsand

(Hay district) B. dura var. pilosa

15(12). Spikelets arranged in four rows, sparsely hairy with
distinct tufts of erect, stiff hairs near the apex on
either side of the upper glume and lower lemma,
usually between the first and second nerve . . .

B. marlothii

Spikelets arranged in two rows, without distinct

tufts of hairs 16

16(15). Rachis of racemes flat, ribbon-like, broadly winged,

1.0-1. 9 mm wide; spikelets glabrous

B. arrecta

Rachis of racemes solid, triquetrous or crescentric,
sometimes very narrowly winged, less than 1 mm
wide; spikelets sparsely or densely hairy ... 17
17(16). Lower lemma and palea extending into a short stout
mucro, up to 1 mm long; spikelets with dense,
white or purple, long hairs concentrated at the

apex B. serrata

Lower lemma acute or rounded; palea rounded at
the tip; spikelets sparsely pilose or covered in

dense silky hairs all over 18

18(17). Spikelets very densely hairy; hairs long, white and
silky; rachis very delicate and usually bare for the

lowest part; panicle mostly simple

B. schoenfelderi

Spikelets sparsely pilose; hairs short; rachis
delicate or firm, with or without spikelets on the
lowest part; panicle simple or compound ... 19
19(18). Panicle compound with racemes attached to
primary branches (at least in the lower part);
lowest part of rachis bare; spikelets bearded at

the apex B. malacodes

Panicle simple with racemes attached to the central
axis; spikelets covering the whole length of the
rachis; spikelets not bearded at the apex ... 20
20(19). Spikelets longer than 3 mm, distinctly flattened on
the inner side; central axis and rachis firm . . .

B. advena

Spikelets shorter than 3 mm, not distinctly
flattened; central axis and rachis very delicate
B. eruciformis

Brachiaria advena Vickery

Perennial, or annual; very
loosely tufted, or stoloniferous
(occasionally); 200-800 mm tall.

Leaf blades 50-130 mm long; 2-6
mm wide. Spikelets 3.0-3. 8 mm
long. Culms erect or decumbent
and rooting at the nodes, some-
times straggly; panicle simple;
racemes secund, 10-30 mm long;
spikelets arranged in two rows, sparsely pilose, distinctly
flattened on the inner side, pedicels equal.

Flowering December to March. Usually in damp,
disturbed areas on black clayey soil, often in mealie or
sunflower fields. Infrequent to locally common. Biome: Sa-
vanna and Grassland. Naturalized from Australia. Weed (of

65

cultivation). Resembles B. eruciformis, which has a panicle
with a delicate appearance and shorter spikelets that are not
distinctly flattened on the one side, and B. malacodes,
which has a compound panicle.

Description: Chippindall 1955 (377). Voucher: Smook
4686. PRECIS code 9901040-00100.

Brachiaria arrecta (Dur. & Schinz) Stent

( =B . latifolia Stapf) 2.

Perennial; hydrophyte, or sto-
loniferous, or tufted (prostrate
and rooting at the nodes);

500-1300 mm tall. Leaf blades
50-250 mm long; 5-15 mm wide.

Spikelets 3. 0-4. 3 mm long.

Racemes secund, 10-50 mm
long; rachis flat and ribbon-like, broadly winged, 1.0- 1.9
mm wide; spikelets glabrous, arranged in two rows,
pedicels equal; lower glume 1-nerved, less than 1/2 the
spikelet length.

Flowering December to June. In shallow water of river
floodplains or vleis, but also extending to areas around
rivers and lakes, often in the shade and usually on wet soils.
Locally common. Biome: Savanna and Grassland. Tropical
east Africa. Introduced to tropical America. The flat
ribbon-like rachis distinguishes this from other Brachiaria
species with spikelets arranged in two rows and 1-nerved
lower glumes. Some specimens have previously been
wrongly identified as B. rugulosa.

Description: Stapf 1898-1900 (393), Chippindall 1955
(374), Clayton et al. 1970-1982 (585). Voucher: Smook
1920, De Winter & Marais 4912. PRECIS code
9901040-00200.

Brachiaria bovonei (Chiov.) Robyns

Wiry signal grass.

Perennial; densely tufted;

250-1000 mm tall. Leaf blades
30-300 mm long; 3-6 mm wide.

Spikelets 3. 2^L 5 mm long. Leaf
blades flat or sometimes
convolute, usually much less than
1/2 the length of the culms,
usually hairy; racemes secund, 10-30(-50) mm long; spike-
lets arranged in two rows, pedicels equal; lower glume 2/3
the spikelet length, 7-nerved; cross-veins present on upper
glume and lower lemma.

Flowering October to January. In wet, marshy or damp
areas around vleis, dams or streams in open veld or on
mountain slopes, usually on sandy soils. Infrequent to local-
ly common. Biome: Savanna and Grassland. Southern
tropical Africa. Very similar to B. suhulifolia, which has
filiform leaf blades that can be up to 1/2 the culm length.

Description: Chippindall & Crook 1976 (129), Clayton
et al. 1970-1982 (582). Voucher; Liebenberg 2805.
PRECIS code 9901040-00250.

Brachiaria brizantha (A. Rich.) Stapf

PI. 22.

Broodsinjaalgras; common
signal grass.

Perennial; loosely tufted
(often robust); 300-2000 mm tall.

Leaf blades 100-400 mm long;

7-20 mm wide. Spikelets 4-6 mm
long. Racemes secund, 25-100
mm long; spikelets arranged in a
single row or occasionally with two rows near the base,
pedicels equal; lower glume 7-nerved, less than 1/2 the
spikelet length, separated from the upper glume by a short
intemode.

Flowering October to May. Prefers undisturbed areas
near streams, especially under trees in open woodland,

usually in sandy or rich soils. Common. Biome: Savanna,
Grassland, and Nama-Karoo. Tropical Africa. Palatable
pasture (good forage value). Distinguished from other
Brachiaria species by the 7-nerved lower glume that is
separated from the upper glume by a short internode.

Description: Chippindall & Crook 1976 (125), Stapf
1920 (531), Launert 1970 (160:40), Stapf 1 898-1900 (386),
Chippindall 1955 (371), Clayton et al. 1970-1982 (587).
Voucher: De Winter 3913. PRECIS code 9901040-00300.

Brachiaria chusqueoides (Hack.) Clayton

(-Panicum chusqueoides
Hack.) 2.

Annual; tufted (scandent or
creeping); 300-750 mm tall. Leaf
blades 30-120 mm long; 3-10
mm wide. Spikelets 3-5 mm long.

Leaf blades cordate and
pseudopetiolate at the base;
racemes usually fewer than 5, lax and widely spaced, 15-70
mm long; spikelets more than their own length apart;
pedicels of unequal lengths.

Flowering October to April. Forest undergrowth in
disturbed or more open places, frequently in coastal dune
forest, on deep sand or humiferous soil. Common. Biome:
Savanna. Northwards into tropical east Africa. Resembles
B. grossa , which lacks pseudopetiolate leaf blade bases, has
more racemes and grows in Namibia, Botswana and
northern Transvaal, and similar to B. deflexa, which usually
has smaller spikelets and lacks pseudopetioles.

Description: Clayton et al. 1970-1982 (590). Voucher:
Anderson 37. PRECIS code 9901040-00350.

Brachiaria deflexa (Schumach.) C.E. Hubb. ex Robyns

Fig. 30. PI. 23.

(=Pseudobrachiaria deflexa
(Schumach.) Launert) 2.

False signal grass, bastersin-
jaalgras.

Annual; loosely tufted (culms
often weak and ascending,
solitary or branched); 150-700
mm tall. Leaf blades 40-180(-250) mm long; 4-22 mm
wide. Spikelets 2. 0-3. 4 mm long. Panicle broadly ovate,
branches rigid, simple or compound; racemes 7-15, 20-100
mm long, pedicels unequal, the longer one up to 15 mm
long; upper lemma finely rugose.

Flowering December to June. Shady places in open
woodland or forest margins, often ruderal in disturbed
areas. Common. Biome: Savanna, Grassland, Nama-Karoo,
and Desert. Northwards to Senegal and Yemen with a few
records from India. Formerly placed in a separate genus
P seudobrachiaria , but because of its very close relationship
with B. grossa , which has larger spikelets and a coarsely
rugose upper lemma, and with B. chusqueoides , which has
pseudopetiolate leaf blades, this species is retained in
Brachiaria pending further research.

Description: Chippindall & Crook 1976 (122), Muller
1984 (212), Chippindall 1955 (378), Clayton et al.
1970-1982 (598). Illustration: Chippindall 1955 (fig. 323).
Voucher: Smook 1138. PRECIS code 9901040-00380.

Brachiaria dictyoneura (Fig. & De Not.) Stapf

Perennial; densely tufted and
rhizomatous; 300-1200 mm tall.

Leaf blades 50-300 mm long;

3— 1 0(— 30) mm wide. Spikelets
5-7 mm long. Racemes 4-12,
secund, 10-80 mm long; spikelets
arranged in two rows, pedicels
equal; lower glume more than 2/3
the spikelet length, 7-9-nerved,

66

separated from upper glume by a short internode; upper
glume and lower lemma with cross-veins between the
nerves.

Flowering November to March. Usually in bush or
mixed mopane veld or along roadsides in damp ditches. In-
frequent. Biome: Savanna. Tropical Africa to Ethiopia.
Closely related to B. humidicola, which has fewer racemes,
is stoloniferous and has culms often decumbent and rooting
at the nodes.

Description: Stapf 1920 (512), Launert 1970 (160:39),
Chippindall 1955 (372), Clayton et al. 1970-1982 (582).
Voucher: Anderson 51. PRECIS code 9901040-00400.

Brachiaria dura Stapf var. dura

Perennial; rhizomatous (rhi-
zome short, oblique); 400-1500
mm tall. Leaf blades 100-350
mm long, convolute and wiry; 1-3
mm wide. Spikelets 4-5 mm long.

Racemes secund, (l-)2, 90-120
mm long; spikelets glabrous, in
one or two rows, pedicels equal;
lower glume 1/2 the spikelet
length, 4-nerved.

Flowering December to May. On dunes or sandy soil
along dry rivers and floodplains, often in the shade. Locally
common. Biome: Savanna. Northwards to Guinea.

Distinguished from var. pilosa by spikelet vestiture and
distribution. Similar to B. xantholeuca , which is annual, has
smaller spikelets and broader leaf blades.

Description: Stapf 1920 (531), Launert 1970 (160:40).
Voucher: Maguire 2206. PRECIS code 9901040-00500.

Brachiaria dura Stapf var. pilosa J.G. Anders.

Perennial; rhizomatous (rhi-
zome usually deeply buried);

500-1300 mm tall. Leaf blades
100-350 mm long, convolute and
wiry ; 1-3 mm wide. Spikelets 4-5
mm long. Racemes ( 1— )2(— 3),
secund, 90-120 mm long; spike-
lets densely pilose, in one or two
rows, pedicels equal; lower
glume 1/2 the spikelet length, 3-4-nerved.

Flowering December to April. At Witsand on white sand
dunes. Rare. Locally common (Witsand). Biome: Savanna.
Distinguished from var. dura by spikelet vestiture and
distribution. Similar to B. xantholeuca, which is annual, has
smaller spikelets and broader leaf blades.

Description: Anderson 1961 Kirkia 1 (104). Voucher:
Leistner 1372. PRECIS code 9901040-00600.

Brachiaria eruciformis (J.E. Sm.) Griseb.

Litjiesinjaalgras; sweet signal
grass.

Annual; loosely tufted (some-
times straggly or procumbent);

100-500(-1000) mm tall. Leaf
blades 20-150 mm long; 2-6 mm
wide. Spikelets 1.7-2. 7 mm long.

Culms erect or decumbent and
rooting at the nodes; racemes secund, 1 0— 25(— 30) mm long;
spikelets sparsely pilose, arranged in two rows, pedicels
equal.

Flowering November to May. In moist places on clay
or black turf and in disturbed areas. Common. Biome: Sa-
vanna, Grassland, and Nama-Karoo. Northwards to the
Mediterranean and then eastwards to India, naturalized in
U.S.A. Indicator (waterlogged soils), or weed (in gardens
and cultivations). Resembles B. advena , which has longer
spikelets that are distinctly flattened on the inner side and
has panicles with firm, stout central axes and rachises.

Description: Chippindall & Crook 1976 (123), Launert
1970 (160:42), Chippindall 1955 (376), Clayton et al.
1970-1982 (590). Voucher: Leistner 1243. PRECIS code
9901040-00700.

Brachiaria glomerata (Hack.) A. Camus

Annual; loosely tufted (de-
cumbent, sometimes rooting from
the lower nodes); 100-300(-600)
mm tall. Leaf blades 50-130 mm
long; 5-10(-18)mm wide. Spike-
lets 2-A mm long. Plants covered
with soft, white, velvety hairs;
racemes not conspicuously
secund, appressed to the central
axis, 20-30 mm long; spikelets densely clustered around the
rachis, pedicels equal but inconspicuous; rachilla extension
short and stalk-like.

Flowering December to June. On red sand dunes or
sandy patches on granite outcrops, also in dry water
courses. Locally common. Biome: Savanna, Nama-Karoo,
and Desert. ?Endemic. Closely related to B. psammophila,
which is golden-hairy and lacks a rachilla extension. Giess
13422 and a few other specimens might represent another
taxon with larger, loosely arranged spikelets that combine
the short velvety hairs and rachilla extention of B.
glomerata with the robust habit of B psammophila.

Description: Launert 1970 (160:40), Stapf 1898-1900
(393), Chippindall 1955 (379). Illustration: Chippindall
1955 (fig. 324). Voucher: Jensen 249; Giess 13422.
PRECIS code 9901040-00900.

Brachiaria grossa Stapf

Annual; tufted (with few basal
leaves); 300- 1 000(— 1500) mm
tall. Leaf blades 50-300 mm
long; 4-1 5(— 20) mm wide. Spike-
lets 3.0 — 4. 2 mm long. Leaf blades
oblique and rounded at the base;
panicle broadly ovate; racemes
5-12, rigid, 30-100 mm long;
spikelets spaced, appearing
loosely continuous; pedicels of unequal lengths; upper
lemma coarsely rugose.

Flowering January to April. In sandy pockets of soil on
granite outcrops or on rocky mountain slopes, also around
pans or rivers, occasionally in the shade. Infrequent and lo-
cally common. Biome: Savanna. North to Tanzania. Pasture
(seldom cultivated), or weed (in lucerne but not very
common). Very closely related to B. chusqueoides, which
has cordate, pseudopetiolate leaves and grows in Natal and
the Cape, and to B. deflexa, which is a smaller plant with
smaller spikelets.

Description: Stapf 1920 (547), Chippindall 1955 (379),
Clayton et al. 1970-1982 (597). Voucher: Giess, Volk &
Bleissner 5712. PRECIS code 9901040-01000.

Brachiaria humidicola (Rendle) Schweick.

Creeping signal grass; kruip-
sinjaalgras.

Stoloniferous (culms procum-
bent except for flowering parts);

400-1100 mm tall. Leaf blades
40-250 mm long; 3-16 mm wide.

Spikelets 4. 5-6.0 mm long.

Racemes 1-4, widely spaced,
secund, 25-55 mm long; spikelets in 1-2 rows, pedicels
equal; lower glume more than 2/3 the spikelet length, 7-
nerved, separated from the upper glume by a short
intemode; upper glume and lower Jemma with cross-veins
between the nerves.

67

Flowering December to May. Favours wet areas such as
vlei edges or seasonally swampy grassland but also extends
into woodlands, usually on sandy soils. Infrequent to locally
common. Biome: Savanna and Grassland. Tropical Africa
to Sudan and Ethiopia. Erosion control (roadsides). Closely
related to B. dictyoneura, which has more racemes and is
densely tufted with a creeping rhizome and erect culms.

Description: Chippindall & Crook 1976 (126), Launert
1970 (160:39), Chippindall 1955 (372), Clayton et al.
1970-1982 (583). Illustration: Chippindall 1955 (fig. 319).
Voucher: Smith 2636. PRECIS code 9901040-01 100.

Brachiaria malacodes (Mez & K. Schum.) Scholz

(=B. poaeoides Stapf) 3.

Annual; very loosely tufted
(culms usually erect but often de-
cumbent and rooting at the nodes,
few basal leaves); 200-850 mm
tall. Leaf blades 50-180 mm
long; 4-11 mm wide. Spikelets

2.0-3. 7 mm long. Panicle
compound, branching at least in the lower part, branches
filiform, lax and bare for about half their length; racemes
5-25 mm long, secund; spikelets sparsely pilose, bearded
at the apex, arranged in two rows.

Flowering February to May. In forests or open mopane
woodland on sand or black clay in vleis, often in seasonally
wet depressions. Locally common. Biome: Savanna.
Angola. Valuable pasture (Namibia). Related to B. advena
andB. eruciformis, which have unbranched panicles.

Description: Stapf 1919 (554), Chippindall 1955 (377).
Illustration: Chippindall 1955 (fig. 322). Voucher: Smook
5086. PRECIS code 9901040-01170.

lemma also acuminate.

Flowering November to April. Open veld or bush on
rocky slopes or among rocks, usually on sandy or well-
drained soils. Common (usually scattered but sometimes
forming dense stands). Biome: Savanna and Nama-Karoo.
Southern tropical Africa with interrupted northern
extensions. Palatable pasture (with good forage value).
Fairly easily distinguished from other Brachiaria species by
the tightly clasping lower glume that is dark coloured below
and often long-acuminate.

Description: Chippindall & Crook 1976 (128), Stapf
1920 (535), Launert 1970 (160:42), Stapf 1898-1900 (388),
Chippindall 1955 (374), Clayton et al. 1970-1982 (587).
Illustration: Chippindall 1955 (fig. 320). Voucher: Smook
4871, Story 6127. PRECIS code 9901040-01300.

Brachiaria psammophila (Welw. ex Rendle) Launert

Annual; tufted (erect or de-
cumbent with few basal leaves);

250-400 mm tall. Leaf blades
40-130 mm long; 7-13 mm wide.

Spikelets 3. 0-4. 5 mm long. Plant
covered with very dense, long
golden-yellow hairs; racemes not
conspicuously secund, 10-30 mm
long, mostly appressed to the
central axis; spikelets clustered, lacking a conspicuous
rhachilla extension; pedicels equal.

Flowering November, March, and April. On sand dunes
or in dry river beds. Rare and conservation status not
known. Biome: Savanna and Desert. Angola. Closely
related toB. glomerata, which is covered with short velvety
hairs and has spikelets with a conspicuous rhachilla
extension.

Description: Launert 1970 (160:41). Voucher:

Merxmuller & Giess 3066 1 . PRECIS code 990 1 040-0 1 500.

Brachiaria marlothii (Hack.) Stent

Brachiaria schoenfelderi C.E. Hubb. & Schweick.

Usually annual; stoloniferous,
or tufted (decumbent and rooting
from the lower nodes); 70-500
mm tall. Leaf blades 10-140 mm
long; 1-5 mm wide. Spikelets

2.0-2. 5 mm long. Racemes
secund, 25-50 mm long; spikelets
arranged in four rows, sparsely
hairy on the nerves, with very
distinct tufts of hairs in the upper half on either side of the
upper glume and lower lemma; pedicels equal.

Flowering December to May. Most frequently in
disturbed or heavily grazed areas on shallow sand or loam
near dams or in seasonally moist spots. Common. Biome:
Savanna, Grassland, and Nama-Karoo. Pasture (good
fodder for sheep), or weed (can be a nuisance in lawns and
gardens).

Description: Launert 1970 (160:41), Stapf 1898-1900
(390), Chippindall 1955 (376). Voucher: Smook 2882.
PRECIS code 9901040-01200.

Brachiaria nigropedata (Fical. & Hiern) Stapf

Wurmsinjaalgras; spotted
signal grass.

Perennial; densely tufted and
rhizomatous (rhizome long and
creeping); 300-1200 mm tall.

Leaf blades to 300 mm long; 5-9
mm wide. Spikelets 3-4 mm long.

Racemes secund, 1 5— 30( — 40) mm
long; spikelets in two rows, pedicels equal; lower glume
acuminate but very variable in the same inflorescence,
distinctly darker coloured at the base and clasping the short
internode between the two glumes; upper glume and lower

Annual, or perennial; occasi-
onally stoloniferous, or tufted
(culms branching at the base,
often decumbent and rooting at
the nodes); 300-800 mm tall.

Leaf blades 30— 120(— 1 50) mm
long; 3-8 mm wide. Spikelets

2.0-3.4 mm long. Racemes se-
cund, 10-35 mm long, often
incurved, lower racemes often bare in the lower part; rachis
delicate and lax; spikelets very densely pilose, arranged in
2 rows; pedicels equal.

Flowering February to May. In gravelly and black vlei
soil in depressions, in mopane veld or bushveld. Locally
common (plants usually scattered amongst other grasses).
Biome: Savanna. Distinguished from B. malacodes, B.
advena and B. eruciformis by its very densely hairy spike-
lets.

Description: Launert 1970 (160:41), Chippindall 1955
(378). Voucher: Smook 5119. PRECIS code 9901040-
01600.

Brachiaria serrata (Thunb.) Stapf

(=B. serrata (Thunb.) Stapf
var. serrata ) 2; (=B. serrata
(Thunb.) Stapf var. gossypina
(A. Rich.) Stapf) 2.

Red top grass; rooisinjaalgras.

Perennial; tufted (densely or
loosely), or rhizomatous; 300-
750 mm tall. Leaf blades 50-250 mm long; 2-10 mm wide.
Spikelets 2. 3-4. 5 mm long. Vegetative parts very variable;
leaves can be in a dense tuft at the base or cauline, leaving

Fig. 29. PI. 24.

68

the base bare; culms single or profusely branched, racemes
secund, 10-25 mm long; spikelets with dense, long, silky,
white or purple hairs concentrated at the apex; pedicels
equal; lower lemma and palea extending into a short stout
mucro up to 1 mm long.

Flowering October to May. On shallow sandy soil,
usually in rocky areas or on mountain slopes but extending
into open bush, grassland and occasionally to vlei edges.
Common. Biome: Fynbos, Savanna, and Grassland.
Northwards into tropical Africa. Pasture (average forage
value). The mucronate lower lemma and palea, serrate leaf
margins and concentration of hairs on the spikelet apex
distinguish this species from B. arrecta, B. dura and B.
xantholeuca, which all also have 1 -nerved lower glumes

Fig. 30. Brachiaria deflexa

and spikelets arranged in two rows. Previously a var.
gossypina was recognized, which has a distinctly different
habit and leaves, but further study is needed to establish if
this is a distinct taxon.

Description: Chippindall & Crook 1976 (124), Stapf
1920 (537), Stapf 1898-1900 (388), Chippindall 1955
(375), Clayton et al. 1970-1982 (588). Illustration: Chip-
pindall 1955 (pi. 12(11)). Voucher: Smook 4849. PRECIS
code 9901040-01700.

Brachiaria subulifolia (Mez) Clayton

(=B. filifolia Stapf) 2.

Perennial; rhizomatous (rhi-
zome oblique), or tufted (erect);

200-1000 mm tall. Leaf blades
50-200 mm long; filiform,

0. 7-1.0 mm wide. Spikelets
2. 8-4.0 mm long. Leaf blades
usually up to half the length of the
culms, usually glabrous; racemes secund, 10-30 mm long;
spikelets arranged in 2 rows; pedicels equal; lower glume
2/3 the spikelet length, 7-nerved, cross-veins present on
upper glume and lower lemma.

Flowering September to November. Frequently in damp
or seepage areas on sandy soils. Rare and conservation sta-
tus not known. Infrequent. Biome: Savanna and Grassland.
Northwards into east Africa. Very similar to B. bovonei,
which has flat or convolute leaf blades that are hairy and
much shorter than the culms.

Description: Chippindall 1955 (373), Clayton et al.
1970-1982 (582). Voucher: De Winter & Codd 164.
PRECIS code 9901040-01850.

Brachiaria xantholeuca (Schinz) Stapf

Annual; tufted (culms decum-
bent and branching at the lower
nodes); 200-600 mm tall. Leaf
blades 30-150 mm long; 4—10
mm wide. Spikelets 2. 7-4.0 mm
long. Leaf blades broadly linear
to narrowly lanceolate, velvety
pubescent; racemes secund,

20-70 mm long; spikelets
arranged in 2 rows; pedicels equal; lower glume 3-nerved,
1/2 the spikelet length.

Flowering November to March. Among trees, usually
near water in sandy loam or clayey soils, often in
overgrazed and disturbed places. Locally common. Biome:
Savanna. Tropical Africa. Weed (in some areas). Closely
related to B. dura , which is perennial with larger spikelets
and narrower leaf blades.

Description: Stapf 1920 (541), Launert 1970 (160:42),
Clayton et al. 1970-1982 (597). Voucher: Smook 4778.
PRECIS code 9901040-01900.

Brachyachne (Benth.) Stapf

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 80-700 mm high; herbaceous. Leaf blades linear;
flat, or rolled (involute and filiform). Ligule a fringed
membrane to a fringe of hairs.

Inflorescence of spike-like main branches', digitate or
subdigitate; espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; subsessile', compressed
laterally; disarticulating above the glumes. Hairy callus
present. Glumes two; more or less equal; awnless; very dis-
similar (thinly leathery, lower curved, upper straight). All
florets female-fertile, or a single distal incomplete floret

69

also present; proximal incomplete florets absent.

Female -fertile florets 1. Lemmas less firm than the
glumes to similar in texture to the glumes (membranous to
hyaline); 3 nerved; entire, or incised; awnless, or mucronate
(rarely). Palea present; relatively long to conspicuous but
relatively short. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small; ellipsoid; hilum short; pericarp
fused; embryo large.

Fig. 3 1 . Brachyachne patentiflora

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven, or even. PCR sheath
extensions absent. PCR cell chloroplasts centrifugal/
peripheral, or centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 10 species. Africa, Australia. Helo-
phytic to mesophytic; in open habitats (seasonal swamps
and moist rock crevices); glycophytic. Botswana. 1
indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Brachyachne patentiflora (Stent & Rattray) C.E. Hubb.

Fig. 31. PI. 25.

Perennial; tufted; 100-500
mm tall. Leaf blades 30-120 mm
long; about 1 mm wide. Spikelets
3. 0^1. 4 mm long. Old leaf bases
persistent and breaking into
fibres; spikes slender, solitary, or
sometimes paired; glumes not
strongly keeled.

Flowering December to Jan-
uary. Seasonal swamps on waterlogged clayey soil and
moist crevices on rocky outcrops. Infrequent. Biome: Sa-
vanna. Central tropical Africa. Similar in habit to
Microchloa caffra, which has a solitary spike and strongly
keeled glumes.

Description: Chippindall & Crook 1976 (237), Clayton
et al. 1970-1982 (311). Voucher: Smith 4081. PRECIS
code 9902970-00200.

Brachychloa S.M. Phillips

Annual, or perennial; long-stoloniferous. Culms to 500
mm high (usually less); herbaceous; branched above
(sometimes), or unbranched above (usually). Leaf blades
linear to lanceolate; flat. Ligule a fringed membrane .

Inflorescence of spike-like main branches', open (with
the branches appressed in B. schiemanniana and spreading
in B. fragilis ); non-digitate\ espatheate. Spikelet-bearing
axes disarticulating, or persistent; when disarticulating
falling entire.

Spikelets solitary; biseriate; with short-pedicellate
pedicellate spikelets', 3.5-7 mm long; compressed laterally;
disarticulating above the glumes; disarticulating between
the florets. Glumes two; more or less equal; markedly
shorter than the spikelets; decidedly shorter than the adja-
cent lemmas', awnless; similar. Incomplete florets distal to
the female-fertile florets, awnless; proximal incomplete
florets absent.

Female-fertile florets 3-7. Lemmas similar in texture to
the glumes (membranous); without a germination flap; 3-7
nerved (5 to 7 nerved in B. schiemanniana)', incised; very
shortly mucronate (from between the lobes). Palea present;
relatively long. Lodicules 2; fleshy (?); glabrous. Stamens
3. Ovary glabrous. Fruit small (0.8 mm long); hilum short;
pericarp free.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven and even. PCR sheath
extensions present. Maximum number of extension cells 1.
PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Southern Mozambique,
Natal. Mesophytic to xerophytic; in shade and in open
habitats; glycophytic (in coastal forests on sandy soil).
Natal. 2 indigenous species.

References. 1. Phillips. 1982. Kew Bull. 37:133.

Species treatment by M. Koekemoer.

70

1(0). Plants annual; inflorescence with long and deciduous

spikes; lemmas 3-nerved B. fragilis

Plants perennial; inflorescence with short, persistent
spikes (often in a loose head); lemmas 5-7-nerved

B. schiemanniana

Brachychloa fragilis S.M. Phillips

Annual; tufted (culms decum-
bent); 250-500 mm tall. Leaf
blades 50-100 mm long; 2-6 mm
wide. Spikelets 4-5 mm long.

Spikes in inflorescence longer
than 30 mm, deciduous; lemma 3-
nerved.

Sandy soil on coastal dunes.

Rare. Biome: Savanna. Mozam-
bique (Maputo). Only three known specimens, none at PRE.
The holotype, Pooley 1650, was collected at Ulukondo in
Natal and is now housed at Kew.

Description: Phillips 1982 (145 & 159). PRECIS code
9902864-00100.

Fig. 32. Brachychloa schiemanniana

Brachychloa schiemanniana (Schweick.) S.M. Phillips

( =Heterocarpha
schiemanniana Schweick.) 1.

Perennial; stoloniferous;

150-300 mm tall. Leaf blades
40-80 mm long; 5-8 mm wide.

Spikelets 4-7 mm long. Spikes in
inflorescence 15-40 mm long,
persistent; spikelets 6— 8(— 1 0)-
flowered; lemma 5-7-nerved.

Flowering February to May. Sandy soil, dunes and forest
margins. Rare and conservation status not known. Biome:
Savanna. Mozambique.

Description: Phillips 1982 (145). Voucher: Schweikerdt
1908. PRECIS code 9902864-00200.

Fig. 32. PI. 26.

Brachypodium P. Beauv.

Brevipodium A. & D. Love, Trachynia Link, Tragus
Panzer.

Annual, or perennial; long-rhizomatous to caespitose.
Culms 20-2000 mm high; herbaceous; unbranched above.
Leaf blades linear; flat, or rolled (convolute). Ligule an
unfringed membrane.

Inflorescence a single raceme , or paniculate (rarely)',
open; espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; distichous; 13-40 mm long', com-
pressed laterally; disarticulating above the glumes. Glumes
two; very unequal, or more or less equal; markedly shorter
than the spikelets; awned, or awnless; similar (lanceolate).
Lower glume 5-7 nerved. Incomplete florets distal to the
female-fertile florets, merely underdeveloped, awned, or
awnless; proximal incomplete florets absent.

Female-fertile florets 8-22. Lemmas similar in texture
to the glumes; 7 nerved; entire; awned. Awns 1; median;
apical; non-geniculate; much shorter than the body of the
lemma to about as long as the body of the lemma. Palea
present; relatively long. Lodicules 2; membranous; ciliate.
Stamens 3. Ovary hairy. Fruit medium sized; hilum long-
linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 5, 7, 9, and 10. Pooideae; Triticodae; Brachy-
podieae. 16 species. Temperate, and tropical mountains.
Mesophytic; in shade and in open habitats. Transvaal,
Orange Free State, Natal, Lesotho, and Cape Province. In-
digenous species (2), naturalized species (1).

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA.

Species treatment by M. Koekemoer.

1(0). Lemma awns 10-25 mm long; plants annual

B. distachyon

Lemma awns 4-8 mm long; plants perennial .... 2
2(1). Culms erect, racemes straight, with \-4 spikelets'
crowded near the apex; leaves mainly basal, rigid,
erect, lanceolate; a mountain grassland species . .

B. bolusii

Culms straggling, racemes usually flexuous, with
(3-)5-9 spikelets along the rachis; leaves mostly
cauline, soft, spreading, linear; a forest species that
occasionally extends to thickets and bushland . .
B. flexum

71

Fig. 33. Brachypodium flexum

Brachypodium bolusii Stapf

Perennial; densely tufted;

15(M-50(-700) mm tall. Leaf
blades 20-70(-170) mm long; to
4 mm wide. Spikelets to 30 mm
long. Leaves basal, rigid, erect,
lanceolate; racemes to 55 mm
long, straight, with 1-4 spikelets
crowded near the apex; lemma
awns 4—8 mm long.

Flowering November to March. In mountain grassland.
Locally common. Biome; Savanna and Grassland.
Endemic. Very similar to B. flexum, which usually has
straggling culms, flexuous racemes, spikelets (3— )5— 9 and
leaves cauline. Intermediates between these two species are
common and very difficult to place.

Description: Stapf 1898-1900 (737), Chippindall 1955
(68). Illustration: Chippindall 1955 (fig. 40). Voucher:
Edwards 646. PRECIS code 9904320-00100.

Brachypodium distachyon (L.) Beauv.

Slender to fairly robust an-
nual; tufted (culms often
decumbent and branching near
the base); 100-500(-700) mm
tall. Leaf blades 20-90 mm long;
to 5 mm wide. Spikelets
10-35(-40) mm long. Leaves
mostly cauline, rigid or soft,
young leaves erect, old leaves
curly; racemes ( 1 0— )30— 90(— 1 00) mm long, with 2-6 spike-
lets; lemma awn 10-25 mm long.

Flowering sporadically throughout the year, but mainly
September to January. In disturbed places such as gardens,
excavations, roadsides and waste places, occasionally in
native vegetation. Locally common. Naturalized from the
Mediterranean. Biome: Fynbos and Succulent Karoo.
Europe and the Mediterranean. Weed.

Description: Smith 1980 FI. Europ. (5:189), Stapf
1898-1900 (735), Hitchcock & Chase 1950 (57), Chippin-
dall 1955 (68). Voucher: Smook 3711. PRECIS code
9904320-00200.

Brachypodium flexum Nees

Fig. 33. PI. 27.

Perennial; culms straggling,
slender, wiry, sometimes decum-
bent and rooting at the lower
nodes; 300-900 mm tall. Leaf
blades 50-170 mm long; 2-8 mm
wide. Spikelets 1 2 — 44 mm long.

Leaves mainly cauline, linear; ra-
cemes 60-120 mm long, usually
flexuous, with (3— )5— 9 spike-
lets spread along the rachis; lemma awn 4—8 mm long.

Flowering October to April. In moist shady places of
forests, usually near streams, occasionally in thickets and
bushland. Locally common. Biome: Fynbos, Savanna, and
Grassland. Endemic. Very similar to B. bolusii , which has
leaves tufted at the base, culms erect and racemes straight
with 1-4 spikelets.

Description: Clayton 1972 FTWA (371), Stapf

1898-1900 (736), Chippindall 1955 (68), Clayton et al.
1970-1982 (71). Illustration: Chippindall 1955 (fig. 39),
Clayton et al. 1970-1982 (fig. 24). Voucher: Davidse 6787.
PRECIS code 9904320-00300.

72

Briza L.

Chascolytrum Desv. ,Chondrachyrum Nees ,Tremularia
Fabric.

Annual, or perennial; long-rhizomatous, or caespitose.
Culms 50-1000 mm high; herbaceous; unbranched above.
Leaf blades linear to linear-lanceolate; flat. Ligule an
unfringed membrane.

Inflorescence paniculate; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets 2.5-25 mm long; compressed laterally; disar-
ticulating above the glumes. Glumes two; more or less
equal; markedly shorter than the spikelets; awnless; similar
(broad & cordate; thin & papery). Incomplete florets florets
distal to the female-fertile florets, merely underdeveloped,
awnless; proximal incomplete florets absent.

Female-fertile florets 4-20. Lemmas similar in texture
to the glumes; 7-15 nerved; as broad as long, gibbous and
umbonate, cordate at base\ entire, or incised (obtuse,
cuspidate, bidentate or mucronate); awnless, or mucronate
(the mucro less than 1.5 mm). Palea present; conspicuous
but relatively short. Lodicules 2; membranous; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short, or
long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, jr = 5 and 7. Pooideae; Poodae; Poeae. 16 species.
North temperate, South America. Mesophytic; mostly in
open habitats (dry to moist soils). Transvaal, Natal. Orange
Free State and Cape Province. 3 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1971. FZ. 3. Matthei. 1975. Willdenowia 8: 79. 4. Clayton
& Renvoize. 1986. Gen. Gram. 5. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

Fig. 34. Briza minor

1(0). Plants perennial; lemmas acuminate

B. subaristatum

Plants annual; lemmas obtuse to obscurely acuminate

2

2(1). Panicle appearing secund, with 3-12 nodding
spikelets; spikelets longer than wide, 8-25 mm

long, 8-15 mm wide B. maxima

Panicle open, with more than 20 spikelets; spikelets
as wide as, or slightly wider than long, 3-5 mm
long, 3-6 mm wide B. minor

Briza maxima L.

PI. 28.

Big quaking grass, groot-
bewertjiegras.

Annual; loosely tufted (culms
erect, simple or branched);

100-600(-1000) mm tall. Leaf
blades 70-250 mm long; 3-8 mm
wide. Spikelets 8-25 mm long;

8-15 mm wide. Panicle open,
branches single or in fascicles of 2-4; spikelets 3-12,
longer than wide, 7-20-flowered, nodding, solitary at the
tips of slender branches; glumes green, brown or purple,
5-9-nerved; lemmas straw-coloured, sometimes variegated
with purple.

Flowering July to December. Mostly on well-drained
soils in disturbed areas, especially on roadsides or on the
margins of irrigated lands, cultivated in gardens. Locally
common. Naturalized from the Mediterranean region.
Biome: Fynbos, Savanna, and Grassland. Mediterranean
region, naturalized in many warm temperate countries. Do-
mestic use (for dried flower arrangements), or ornamental
(in gardens), or weed (roadsides).

Description: Chippindall & Crook 1976 (202), Linder
(43), Stapf 1898-1900 (708), Hitchcock & Chase 1950
(137), Chippindall 1955 (49), Clayton et al. 1970-1982
(53). Illustration: Chippindall 1955 (fig. 16), Clayton et al.
1970-1982 (fig. 19). Voucher: Loxton 205. PRECIS code
9904040-00100.

Briza minor L.

Little quaking grass, klein-
bewertjiegras.

Annual; loosely tufted (culms
soft with dark nodes, often
branched near base); 100-600
(-700) mm tall. Leaf blades
40-220 mm long; 3-9 mm wide.

Spikelets 3-5 mm long; 3-6 mm
wide. Panicle open, spreading, branches single or in
fascicles of 2^4; spikelets numerous, almost as wide as or
slightly wider than long, 3-8-flowered; glumes 3-5-
nerved; glumes and lemmas green or tinged with purple,
margins distinctly lighter in colour.

Flowering September to December. Usually found in
moist shady and disturbed places around streams and vleis
on loam or clayey soils. Infrequent to locally common. Nat-
uralized from the Mediterranean region. Biome: Fynbos,
Savanna, and Grassland. Mediterranean region, naturalized
in many warm temperate countries. Ornamental (in gardens
and in dried flower arrangements), or weed (roadsides).

Description: Chippindall & Crook 1976 (203), Linder
(42), Stapf 1898-1900 (709), Hitchcock & Chase 1950
(137), Chippindall 1955 (49), Clayton et al. 1970-1982
(53). Illustration: Chippindall 1955 (fig. 17), Hitchcock &
Chase 1950 (fig. 266). Voucher: Smook 3686. PRECIS
code 9904040-00200.

Fig. 34.

73

Briza subaristatum Lam.

(=B. triloba Nees) 1;

(=Chascolytrum subaristatum
(Lam.) Desv.) 4.

Perennial; tufted; 300-600
mm tall. Leaf blades 80-200 mm
long, rolled; 1-2 mm wide.

Spikelets 4-5 mm long; 2. 0-3. 5
mm wide. Basal sheaths per-
sisting as fibres; panicle contracted; spikelets ovoid, 6-10-
flowered; glumes mucronate, 5-7-nerved, cordate at the
base.

Flowering October to December. In moist cultivated
areas and on roadsides. Rare. Naturalized from South
America. Biome: Fynbos. South and Central America.

Description: Chippindall 1955 (47). Voucher:

Liebenberg 4221. PRECIS code 9904040-00300.

Bromus L.

Aechmorpha Steud., Anisantha Koch, Avenaria
Fabrich., Bromopsis (Dumort.) Fourr., Ceratochloa P.
Beauv., Forasaccus Bub., Genea (Dumort.) Dumort.,
Libertia Lejeun t, Michelaria Dumort., Nevskiella Krecz &
Vved., Serrafalcus Pari., Stenofestuca (Honda) Nakai,
Triniusa Steud., Trisetobromus Nevski.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 30-1900 mm
high; herbaceous; unbranched above. Sheath margins
joined. Leaf blades linear; usually flat, or rolled (somewhat
involute, or convolute). Ligule an unfringed membrane.

Inflorescence a single raceme (rarely), or paniculate;
open, or contracted; espatheate. Spikelet-bearing axes
persistent.

Spikelets (5-) 10-70 mm long; compressed laterally; dis-
articulating above the glumes. Glumes two; very unequal
(usually), or more or less equal (rarely); decidedly shorter
than the adjacent lemmas; awnless; similar (persistent). In-
complete florets distal to the female-fertile florets, merely
underdeveloped, usually awned, or awnless; proximal in-
complete florets absent.

Female-fertile florets 3-30 (rarely 1-2). Lemmas similar
in texture to the glumes; 5-15 nerved; incised (usually), or
entire; awnless, or mucronate, or awned. Awns when
present 1; median; from the sinus, or dorsal; non-genicu-
late; much shorter than the body of the lemma, to much
longer than the body of the lemma. Palea present; relatively
long to conspicuous but relatively short. Lodicules 2;
fleshy, or membranous; glabrous. Stamens 1-3. Ovary
hairy; with a conspicuous apical appendage ( the styles
lateral ). Fruit medium sized; hilum long-linear; embryo
small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Triticodae; Bromeae. About 150
species. North temperate, tropical mountains, South
America. Mesophytic, or xerophytic; in shade and in open
habitats. Namibia, Transvaal, Orange Free State, Natal,
Lesotho, and Cape Province. Indigenous species (6),
naturalized species (9).

Supposed intergeneric hybrid with Festuca : X

Bromofestuca Prodan.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA. 3. Pinto-Escobar. 1976. Caldasia 11: 9-16. 4.
Linder. 1986. Bothalia 16: 61. 5. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Lemma awns stiff and straight, 30-70 mm long . . 2
Lemma awns fine, straight or spreading, 0.2-25.0 mm

long 3

2(1). Callus of lemma rounded; spikelets usually open,
showing the rachis between the lemmas; panicle

usually lax and spreading B. diandrus

Callus of lemma pointed; spikelets usually tight, very
rarely showing the rachis; panicle usually dense and
erect B. rigidus

74

3(1). Plants perennial; anthers 2-8 mm long 4

Plants annual (or short-lived perennial in B.

catharticus ); anthers 0.5-1. 5 mm long 8

4(3). Old leaf sheaths flimsy, curly, not fibrous; lemma

awns 0.5-5 .0 mm long 5

Old leaf sheaths firm, erect, fibrous; lemma awns

3-15 mm long 6

5(4). Lemma awns shorter than 2 mm; plants rhizomatous

B. inermis

Lemma awns longer than 3 mm; plants tufted

B. leptoclados

6(4). Lemmas glabrous; spikelets (including awns) 35-55

mm long B. natalensis

Lemmas villous; spikelets (including awns) 20^45

mm long 7

7(6). Pedicels glabrous or sparsely hairy, shorter than the

spikelets B. speciosus

Pedicels villous, longer than the spikelets

B. firmior

8(3). Upper glume 3-nerved; lower glume 1-nerved ... 9
Upper glume 5-9-nerved; lower glume 3-9-nerved

(occasionally 1-nerved in B. pectinatus) 11

9(8). Culms hairy below the inflorescence . . . B. rubens
Culms glabrous below the inflorescence 10

10(9). Panicles with spikelets more or less secund; lower
pedicels with about four spikelets; upper glume

7-12 mm long; lemmas 8-13 mm long

B. tectorum

Panicles with spikelets not secund; lower pedicels
with one or two spikelets; upper glume 14-20

mm long; lemmas 12-15 mm long

B. madritensis

11(8). Glumes and lemmas sharply keeled, tips minutely
bifid or acute, occasionally extending into an awn

shorter than 3 mm; spikelets compressed

B. catharticus

Glumes and lemmas rounded on the back, tips
distinctly bifid, awns 3-18 mm long, from the
sinus between the lobes; spikelets more or less

terete 12

12(1 1). Panicle open with at least some of the pedicels

longer than the spikelets 13

Panicle contracted, with all the pedicels shorter than

the spikelets 14

13(12). Awns equalling or shorter than the lemmas; lemma
tips rounded to subacute .... B. commutatus
Awns longer than the lemmas; lemma tips acute to

subacuminate B. pectinatus

14(12). Spikelets 18-25 mm long; awns flattened, twisted

and spreading B. alopecurus

Spikelets 7-15 mm long; awns not flattened,

occasionally twisted and spreading 15

15(14). Spikelets densely villous; awns somewhat
spreading and twisting; plants growing in small

cushions, to 200 mm tall

B. hordeaceus subsp. ferronii

Spikelets villous to scabrid; awns straight and erect;
plants growing in erect tussocks, to 600 mm tall
B. hordeaceus subsp. molliformis

Bromus alopecurus Poir.

Annual; tufted; 150-300 mm
tall. Leaf blades 30-70 mm long;

1-3 mm wide. Spikelets 18-23
mm long; 2-3 mm wide. Panicle
contracted, pedicels absent or
shorter than the spikelets; lower
glume 5-nerved; lemmas 9-12
mm long, awns 8-15 mm long,
flattened, twisted and patent.

Flowering October. Roadsides and waste ground. Rare.
Naturalized from the Mediterranean. Biome: Fynbos.
Central and eastern Mediterranean basin; western Asia.
Known from a single collection at Caledon, Du Toit 1823,

which is in the Stellenbosch herbarium.

Description: Bor 1985 (1808), Smith 1980 FI. Europ.
(5:188). PRECIS code 9904280-00025.

Bromus catharticus Vahl

(=B. unioloides H.B.K.) 3;

( =B . willdenowii Kunth) 3.

Reddingsgras, rescue grass

Short-lived perennial, or ;
nual; tufted; 150-1000 mm t:

Leaf blades 5-8 mm wf
Spikelets 20-35 mm long; 5-8 mm wide. Panicle lax,
pedicels longer than the spikelets; spikelets compressed;
glumes and lemmas prominently keeled, acute; glumes 3-9-
nerved; lemmas 15-25 mm long, minutely bifid, lobes
shorter than 0.5 mm, awns shorter than 3 mm.

Flowering October to April. Usually in moist to wet
places, often in the shade, in disturbed and natural veld.
Common. Naturalized, different strains from Europe,
Australia and America. Biome: Fynbos, Grassland, and
Nama-Karoo. Worldwide. Very good annual winter pasture,
or erosion control, or weed (in disturbed and moist places).
The great variability in this species can be attributed to the
large range of habitats and growth conditions in which it
appears.

Description: Chippindall & Crook 1976 (213), Stapf
1898-1900 (734), Chippindall 1955 (63), Clayton et al.
1970-1982 (67). Illustration: Chippindall 1955 (fig. 35).
Voucher: Smook 3879. PRECIS code 9904280-00050.

Bromus commutatus Schrad.

Hairy chess.

Annual; tufted; 150-900 mm
tall. Leaf blades 60-150 mm
long; 3-6 mm wide. Spikelets
10-25 mm long; 4-7 mm wide.

Panicle lax, pedicels (at least
some of them) longer than the
spikelets; spikelets almost linear
at maturity; lower glume 3-5-nerved, upper 5-9-nerved;
lemmas 7-9 mm long, awns 3-8 mm long.

Flowering September to December. In disturbed, wet
places. Infrequent. Biome: Fynbos, Grassland and Succu-
lent Karoo. Endemic. Pasture.

Description: Stapf 1898-1900 (728), Chippindall 1955
(66). Voucher: Theron 341. PRECIS code 9904280-00100.

Bromus diandrus Roth

Langnaaldbromus, predikant-
luis, ripgut brome.

Annual; loosely tufted; 300-
1 100 mm tall. Leaf blades 50^100
mm long; 3-8 mm wide. Spike-
lets 30-90 mm long (including
awns); 3-8 mm wide. Panicle
usually lax and spreading, ped-
icels glabrous, scabrid or villous; spikelets open, wedge-
shaped; lower glume 1-nerved, upper 3-nerved; lemmas
12-22 mm long, awns stiff and straight, 30-70 mm long.

Flowering September to January (occasionally in
March). In disturbed and weedy places. Locally common.
Naturalized from Europe. Biome: Fynbos and Nama-Karoo.
Mediterranean region eastwards to central Asia. Introduced
to temperate countries. Serious weed (of cultivated and
disturbed areas, especially in the winter rainfall areas of the
Cape). Two distinct species, B. diandrus and B. rigidus, are

75

recognized in Europe. In South Africa these species
hybridize to form a complete range of intermediates which
can be named B. diandrus agg.

Description: Bor 1985 (1800), Chippindall & Crook
1976 (214), Smith 1980 FI. Europ. (5:183), Chippindall
1955 (67), Clayton et al. 1970-1982 (67). Illustration:
Chippindall 1955 (fig. 38). Voucher: Von Breitenbach 43.
PRECIS code 9904280-00200.

Bromus firmior (Nees) Stapf

(=B . firmior (Nees) Stapf var.
firmior) 5; {-B . firmior (Nees)

Stapf var. leiorhachis Stapf) 5.

Perennial; tufted; 500-1500
mm tall. Leaf blades 200^-00
mm long; 4-8 mm wide. Spike-
lets 15-45 mm long; 4-8 mm
wide. Panicle open, much ex-
serted, pedicels sparsely to densely pilose, longer than the
spikelets; spikelets green to purple; lower glume 1-3-
nerved, upper 3-5-nerved; lemmas villous, 1 0-20 mm long,
awns 3-12 mm long.

Flowering November to March. At high altitudes on
moist grassy slopes of the Drakensberg. Locally common.
Biome: Grassland. Endemic. Very closely related to B.
natalensis; which has glabrous spikelets and to B. speciosus,
which has glabrous pedicels.

Description: Stapf 1898-1900 (733), Chippindall 1955
(64). Voucher: Killick 1629. PRECIS code 9904280-
00350.

Bromus hordeaceus L. subsp. ferronii (Mabille) P.M.

Sm.

Annual; tufted (culms decum-
bent-erect); to 200 mm tall. Leaf
blades 50-150 mm long; 3-^1 mm
wide. Spikelets 7-15 mm long;

3-5 mm wide. The densely
villous spikelets, spreading and
twisted awns and plant height
distinguish this subsp. from
subsp. molliformis.

Weedy places. Rare. Naturalized from France and
Britain. Biome: Savanna. Europe. Known from a single
collection, Paterson 2284, housed at BOL.

Description: Smith 1980 FI. Europ. (5:187). PRECIS
code 9904280-00420.

Bromus hordeaceus L. subsp. molliformis (J. Lloyd)

Maire & Weiller

(=6. molliformis Lloyd) 5.

Soft brome.

Annual; tufted; to 600 mm
tall. Panicle contracted, often
elliptical, pedicels pilose to
villous, shorter than the spikelets;
spikelets densely villous; lower
glume 3-5-nerved, upper 7-nerved; lemmas 6-9 mm long,
awns 3-7 mm long, scabrid, erect or somewhat spreading.

Flowering October to February. Cultivated lands and
other disturbed areas. Locally common. Naturalized from
Europe. Biome: Fynbos, Grassland, and Nama-Karoo.
Europe. Weed (of cultivation).

Description: Smith 1980 FI. Europ. (5:187), Stapf
1898-1900 (91), Chippindall 1955 (66). Voucher: Crook
2345. PRECIS code 9904280-00430.

Bromus inermis Leyss.

Smooth brome.

Perennial; rhizomatous;

500-1000 mm tall. Leaf blades
200-500 mm long; 4-7 mm wide.

Spikelets 10-20 mm long. Pani-
cle open, pedicels glabrous to
scabrid; spikelets narrowly
oblong; lower glume 1 -nerved,
upper 3-nerved; lemmas 6.0-8. 5 mm long, awns fine, to 1
mm long.

Flowering November to April. Disturbed or weedy
places. Rare. Naturalized from Europe. Biome: Savanna.
Europe. Planted for cultivated pasture. A very variable
species.

Description: Smith 1980 FI. Europ. (5: 184), Chippindall
1955 (65). Voucher: Galpin 7944. PRECIS code
9904280-00500.

Bromus leptoclados Nees

Mountain brome grass.

Perennial; tufted; (200-)500-
1500 mm tall. Leaf blades
100-300 mm long; 5-13 mm
wide. Spikelets 10-30 mm long;

2-6 mm wide. Panicle open, pedi-
cels scabrid; spikelets open or
with florets closely arranged;
lower glume 1-3-nerved, upper 3-5-nerved; lemmas 8-12
mm long, awns up to 5 mm long.

Flowering October to February. Usually in moist, shady
places along rivers and streams. Locally common. Biome:
Fynbos, Grassland, and Nama-Karoo. Montane areas of
tropical Africa. Palatable natural pasture.

Description: Chippindall & Crook 1976 (215), Stapf
1898-1900 (731), Chippindall 1955 (65), Clayton et al.
1970-1982 (68). Illustration: Clayton et al. 1970-1982 (fig.
23). Voucher: Acocks 18671. PRECIS code

9904280-00850.

Bromus madritensis L.

Spanish brome.

Slender annual; tufted; 1 20—

350(-600) mm tall. Leaf blades
80-200 mm long; 2-5 mm wide.

Spikelets 10-25 mm long; 4-5
mm wide. Panicle dense or some-
what lax, lower pedicels with 1-2
spikelets, pedicels shorter than
the spikelets; lower glume 1 -nerved, upper 3-nerved;
lemmas 12-15 mm long, awns erect to recurved, 15-20 mm
long.

Flowering October. Weedy places. Rare. Biome: Fyn-
bos. Mediterranean region eastwards to Afghanistan.
Naturalized worldwide.

Description: Bor 1985 (1805). Voucher: Parker 4917.
PRECIS code 9904280-01000.

Bromus natalensis Stapf

(=B. natalensis Stapf var.
lasiophilus Stapf) 5; (=B.
speciosus sensu Compton, non
Nees) 5.

Perennial; rhizomatous; 500-
1200 mm tall. Leaf blades 70-
150 mm long; 4-7 mm wide.

Spikelets 35-55 mm long; 5-10

76

mm wide. Panicle open, pedicels sparsely villous to
echinate; spikelets glabrous or finely scabrid; lower glume
3-nerved, upper 5-7-nerved; lemmas 15-18 mm long, awns
6-15 mm long.

Flowering October to January. Rocky hillsides. Infre-
quent. Biome: Grassland. Endemic.

Description: Stapf 1898-1900 (732), Chippindall 1955
(64). Illustration: Chippindall 1955 (fig. 36). Voucher:
Codd 8142. PRECIS code 9904280-01000.

from Europe. Biome: Desert. Mediterranean region

eastwards to central Asia, introduced to North America.
Known from a single collection at Oranjemund. Specimen
at BOL.

Description: Bor 1985 (1807), Hitchcock & Chase 1950
(54). Illustration: Hitchcock & Chase 1950 (fig. 51).
PRECIS code 9904280-01150.

Bromus speciosus Nees

Bromus pectinatus Thunb.

(=B. japonicus sensu
Chippind., non Thunb. var.
japonicus ) 5; ( =B . japonicus
sensu Chippind., non Thunb.
var. velutinus (Nocc.) Aschers.

& Graebn.) 5.

Annual; tufted; 100-800 mm
tall. Leaf blades 50-300 mm
long; 2-8 mm wide. Spikelets 10-30 mm long; 3-6 mm
wide. Panicle open, pedicels longer than the spikelets;
spikelets laterally compressed; lower glume 1 -3-nerved,
upper 5-7-nerved; lemmas 7-14 mm long, tips acute to
subacuminate, awns 6-18 mm long.

Flowering August to February. In disturbed and eroded
areas. Locally common. Biome: Fynbos, Savanna, Grass-
land, and Nama-Karoo. Temperate and montane regions of
Africa and the Middle East.

Description: Chippindall 1955 (68). Voucher: Repton
2633. PRECIS code 9904280-01 115.

Bromus rigidus Roth

Ripgut brome.

Annual; loosely tufted; 400-
700 mm tall. Leaf blades 50-150
mm long; 3-7 mm wide. Spike-
lets 15-25 mm long; 2-6 mm
wide. Panicle contracted, pedicels
hispid to villous; spikelets nar-
row, ovate to linear; lower
glume 1 -nerved, upper 3-nerved; lemmas 20-25 mm long,
awns 30-50 mm long.

Flowering September to October. Disturbed and weedy
places. Infrequent. Naturalized from the Mediterranean
basin. Biome: Fynbos. Northern America and the
Mediterranean region. Weed. Distinct in Europe, but forms
a complete range of intermediates with B. diandrus in South
Africa. These intermediates are best identified as B.
diandrus agg.

Description: Bor 1985 (1802), Smith 1980 FI. Europ.
(5:183), Hitchcock & Chase 1950 (53). Illustration: Hitch-
cock & Chase 1950 (fig. 47). Voucher: Cleghom 3106.
PRECIS code 9904280-01125.

Bromus rubens L.

Red brome, foxtail chess.

Annual; tufted; 150-450 mm
tall. Leaf blades 60-120 mm
long; 3-5 mm wide. Spikelets
21-28 mm long. Culms hairy
below the inflorescence; panicle
erect, compact and ovoid, lower
branches fascicled; spikelets
somewhat wedge-shaped, often reddish; lower glume 1-
nerved, upper 3-nerved; lemmas 13-15 mm long, awns
18-22 mm long.

Disturbed places and waste ground. Rare. Naturalized

Purple brome.

Perennial; tufted; 300-600
mm tall. Leaf blades 80-200 mm
long; 2—4 mm wide. Spikelets
15-50 mm long; 3-6 mm wide.

Panicle open, pedicels glabrous
or sparsely scabrid, almost as
long as the spikelets; spikelets
linear, purplish; lower glume 1-3-nerved, upper 3-5-
nerved; lemmas 10-15 mm long, awns 3-6 mm long.

Flowering December to March. On steep, moist
mountain slopes, occasionally in the shade and along
streams. Infrequent. Biome: Grassland. Endemic. Very
closely related to B.firmior, which has pedicels villous, and
to B. natalensis , which has lemmas 35-55 mm long and
glabrous.

Description: Stapf 1898-1900 (733), Chippindall 1955
(64). Voucher: Dyer 253. PRECIS code 9904280-01200.

Bromus tectorum L.

Annual; tufted; 100-250 mm
tall. Leaf blades 60-100 mm
long; 2-4 mm wide. Spikelets
8-15 mm long; 2—4 mm wide.

Culms glabrous below the
inflorescence; panicle with spike-
lets more or less secund, lower
pedicels with about four spike-
lets; lower glume 1 -nerved, upper
3-nerved; lemmas 8-13 mm long, awns 8-20 mm long.

Flowering August to October. Sandy soil on roadsides.
Rare. Naturalized from the Mediterranean. Biome: Succu-
lent Karoo. Mediterranean region.

Description: Bor 1985 (1811). Voucher: Rosch & Le
Roux 634. PRECIS code 9904280-01300.

Calamagrostis Adans.

Achaeta Foum., Amagris Raf., Ancistrochloa Honda,
Anisachne Keng, Athernotus Dulac, Aulacolepis Hack.,
Chamaecalamus Meyen, Cinnagrostis Griseb.,
Neoaulacolepis Rauschert, Pteropodium Steud.,
Sclerodeuxia Pilger.

Perennial (some species reedlike ); long-rhizomatous, or
long-stoloniferous, or caespitose, or decumbent. Culms
100-2000 mm high; herbaceous; unbranched above. Leaf
blades linear; flat (usually), or rolled (convolute). Ligule an
unfringed membrane (sometimes erose-ciliate).

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 3-7(-8) mm long; compressed laterally; disar-
ticulating above the glumes. Hairy callus present ( the hairs
more than 0.5 mm long, often about as long as, and
surrounding, the lemma). Glumes two; more or less equal;
much exceeding the spikelets; awnless; similar. All florets
female -fertile ; proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline)-, 3-5 nerved; incised; awned. Awns 1 ;
median; from the sinus, or dorsal; non-geniculate, or genic-
ulate; much shorter than the body of the lemma to much
longer than the body of the lemma. Paiea present; relatively

77

^ Sli

long. Lodicules 2; membranous; glabrous. Stamens 3.
Ovary glabrous. Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. About 230
species. Temperate. Mostly helophytic to mesophytic; in
shade or in open habitats; glycophytic, or maritime-
arenicolous to halophytic (rarely — but C. epigeios x
Ammophila arenaria (X Ammocalamagrostis ) is a valuable
sand binder). Transvaal and Cape Province. 1 indigenous
species.

Intergeneric hybrids with Agrostis. C. epigeios
hybridizes with Ammophila arenaria (X

Ammocalamagrostis P. Fourn., a useful sand stabilizer. See
alsoX Calamophila O. Schwartz).

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA.

Species treatment by G.E. Gibbs Russell.

Calamagrostis epigeios (L.) Roth var. capensis Stapf

Fig. 36, PI. 31.

Erect perennial; tufted and

rhizomatous (rhizome creeping);

600-1200 mm tall. Leaf blades to
450 mm long; to 10 mm wide.

Spikelets 5. 5-8.0 mm long. Pani-
cle narrow, to 250 mm long, light
brown; florets with conspicuous
long white callus hairs.

Flowering January to May.

Vleis. Rare (not collected in the eastern Cape mountains
since 1954). To east Africa. Possibly introduced from the
Cape mountains to the Transvaal. The typical variety occurs
in temperate Europe and Asia and has smaller spikelets.

Description: Stapf 1898-1900 (551), Chippindall 1955
(94), Clayton et al. 1970-1982 (103). Illustration: Chippin-
dall 1955 (fig. 66), Clayton et al. 1970-1982 (fig. 35).
Voucher: Codd 2733. PRECIS code 9902460-00100.

Catalepis Stapf & Stent

Perennial; caespitose. Culms 50—400 mm high; herba-
ceous; unbranched above. Leaf blades linear; to I mm wide\
folded, or rolled (rarely flat). Ligule a fringe of hairs.

Inflorescence of spike-like main branches, or panicu-
late; contracted (very much so: the lateral branches short,
sometimes reduced to 4 or 5 spikelets); non-digitate;
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; not secund; 4-5 mm long; compressed
laterally; falling with the glumes (seeming to disarticulate
at base of pedicel). Glumes two; relatively large (i.e., the
upper glumes); very unequal; much exceeding the spikelets 1
(i.e. the upper); awnless; very dissimilar (the lower reduced
to a small subulate scale, the upper lanceolate). All florets
female-fertile; proximal incomplete florets absent.

Female-fertile florets l ( lanceolate ). Lemmas similar in
texture to the glumes (thin); without a germination flap; 3
nerved; entire; awnless. Palea present (broad); relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions present. Maximum number of extension cells 1.
PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. South Africa. Mesophytic
(locally abundant in mountain grassland); in open habitats.
Transvaal, Orange Free State, Natal, Lesotho, and Cape
Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past.

Fig. 36. Calamagrostis epigeios var. capensis

Species treatment by M. Koekemoer.

78

Gause grass.

Fig. 37. PI. 32.

Creeping perennial; rhizomat-
ous and tufted; 100^100 mm tall.

Leaf blades 10-1 50 mm long; 1-2
mm wide. Spikelets 4-5 mm long.

Leaves fine and curly with age;
panicle spike-like, 15-30 mm
long; lower glume reduced to a
small scale; upper glume 1 -nerved.

Flowering January to March. Sometimes in shallow
sandy soil, but more often on black clay in vleis. Locally
common. Biome: Grassland. Highly palatable natural pas-
ture, or erosion control (pioneer), or weed (roadsides).

Description: Chippindall 1955 (207). Illustration: Chip-
pindall 1955 (plate 4). Voucher: Schweickerdt 1760.
PRECIS code 9902942-00100.

Catapodium Link

Scleropoa Griseb. , Synaphe Dulac. Sometimes included
in Desmazeria.

Annual ; caespitose (or culms solitary). Culms 100-500
mm high; herbaceous; unbranched above. Leaf blades
linear; flat, or rolled (convolute when dry). Ligule an
unfringed membrane.

Inflorescence a single raceme, or paniculate (rigid,
spikelike)\ open, or contracted (the branches with small
adaxial pulvini); espatheate. Spikelet-bearing axes
persistent.

Spikelets secund (oppressed to one side of the axis)', 4-9
mm long; compressed laterally; disarticulating above the
glumes. Glumes present; two: more or less equal; markedly
shorter than the spikelets; awnless; carinate', lower
lanceolate, upper ovate. Incomplete florets distal to the
female-fertile florets, merely underdeveloped; proximal in-
complete florets absent.

Female-fertile florets 3-12. Lemmas similar in texture
to the glumes; 5 nerved; entire; awnless. Palea present; rela-
tively long. Lodicules 2; membranous; glabrous. Stamens
3. Ovary glabrous. Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. 2 species. Europe,
Mediterranean. Mesophytic to xerophytic (in dry
microhabitats); in open habitats; maritime-arenicolous, or
halophytic, or glycophytic. Cape Province. 1 naturalized
species.

References. 1. Stace. 1980. FI. Europ. 2. Linder. Unpubl.
ms, FSA.

Species treatment by M. Koekemoer.

79

Catapodium rigidum (L.) C.E. Hubb.

Fig. 38. PI. 33.

(- Desmazeria rigida (L.)

C.E. Hubb.) 1; ( =Scleropoa
rigida (L.) Griseb.) 1.

Fern grass.

Slender annual; tufted (culms
erect or geniculate, fasciculate,
rarely solitary); 100-250(-450)
mm tall. Leaf blades 30-150 mm long; 1 .0— 2.0(— 3.5) mm
wide. Spikelets 5— 7(— 1 0) mm long. Panicle narrow, 50-100
mm long, branches stiff, alternate, simple, bearing 2-5
spikelets; spikelets 6-10-flowered, more or less secund,
borne on trigonous pedicels; glumes more or less equal,
1.8-2. 2 mm long; florets widely spaced on the fragile
rhachilla, awnless.

Flowering October to December. Mostly on waste land
in parks, gardens and disturbed places in moist shady areas,
also in rock crevices amongst other Fynbos species. Infre-
quent to locally common. Naturalized from Mediterranean
region. Atlantic islands, Europe and Mediterranean region
eastwards to Iran. Naturalized in Australia, New Zealand,
Tasmania and temperate north and south America. Common
garden and roadside weed.

Description: Bor 1985 (1721), Chippindall & Crook
1976 (224), Linder (61), Stapf 1898-1900 (718), Hitchcock
& Chase 1950 (77), Chippindall 1955 (50). Illustration:
Chippindall 1955 (fig. 20), Hitchcock & Chase 1950 (fig.
108). Voucher: Hugo 1954. PRECIS code 9904200-00100.

Cenchrus L.

Echinaria Fabric., Nastus Lunell, Raram Adans.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms

50-1000(-1500) mm high; herbaceous; branched above.
Leaf blades linear, or linear-lanceolate; flat, or folded.
Ligule a fringed membrane to a fringe of hairs. The
spikelets of sexually distinct forms on the same plant.

Inflorescence a false spike, with clusters of spikelets on
reduced axes (spikelets in prickly glomerules (burrs)
composed of coalescing spines representing modified
branchlets ); espatheate. Spikelet-bearing axes disarticu-
lating (but the main axis persistent); falling entire (i.e., the
burrs falling).

Spikelets with ‘involucres’ of ‘bristles' (the bristles
coalescing: contrast Pennisetum). The ‘bristles' nearly
always spiny, markedly coalescent basally (not spiny,
merely ciliate, in C. ciliaris). Female-fertile spikelets com-
pressed dorsiventrally; falling with the glumes (i.e., in the
burrs). Glumes present; two; very unequal; awnless; very
dissimilar, or similar (hyaline or membranous). Proximal
incomplete florets /; paleate, or epaleate (rarely), palea
fully developed, or reduced (rarely); male, or sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes (firmly
membranous, dull, papery or coriaceous); smooth; not
becoming indurated; hairless; having the margins lying flat
and exposed on the palea; with a clear germination flap; 3-7
nerved; entire; awnless. Palea present; relatively long.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Photosynthetic pathway. C4; NADP-ME (pauciflorus ,
incertusf XyMS-. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 12. Panicoideae; Panicodae; Paniceae.
22 species. Tropical and warm temperate. Mesophytic to
xerophytic (‘sand-burrs’); in shade, or in open habitats
(grassland, bush and weedy places); maritime-arenicolous,
or glycophytic. Namibia, Botswana, Transvaal, Orange
Free State, Swaziland, Natal, and Cape Province. Indige-
nous species (1), naturalized species (3).

References. 1. Chippindall. 1955. Gr. & Past. 2. Delisle.
1963. Iowa St. Jour. Sci. 37: 259.

Species treatment by H.M. Anderson.

Fig. 39 . Cenchrus ciliaris

1(0). Perennial; bristles 5-10 mm long, joined only at the
base, forming a small inconspicuous disc below the

spikelet cluster C. ciliaris

Annual; bristles and/or spines 2-5 mm long, joined
above the base to form a hard spiny involucre

around the spikelet cluster 2

2(1). Involucre with two distinct clefts; spines all similar,

flattened and spreading at base; no bristles

C. incertus

Involucre without distinct clefts; spines in two whorls,

inner spines connate, outer bristle-like 3

3(2). Involucre with a distinct ovate disc at base; inner
spines with 1-3 shallow grooves .... C. biflorus
Involucre with an inconspicuous disc at base; none of
the spines with grooves C. brownii

Cenchrus biflorus Roxb.

Annual; tufted; to 800 mm
tall. Leaf blades 60-350 mm
long; 4-10 mm wide. Spikelets 5
mm long; 4 mm wide. Inflores-
cence a false spike 20-100 mm
long; spikelet involucre in two
whorls, inner whorl with connate
spines which are plumose on their
inner surface and with 1-3

80

shallow grooves on their outer surface, outer whorl bristle-
like and short, 40-60 spines; base of burr with a distinct
ovoid disc.

Flowering February to June. Mainly sandy soil. Infre-
quent. Invader from tropical America. Biome: Savanna and
Grassland. Pantropical weed.

Description: De Lisle 1963 (333). Illustration: De Lisle
1963 (fig. 2 1 (I-L). Voucher: De Winter9194. PRECIS code
9901400-00100.

Description: Chippindall 1955 (452). Illustration: Chip-
pindall 1955 (fig. 375). Voucher: Fellingham 244. PRECIS
code 9901400-00400.

Centropodia Reichenb.

Asthenatherum Nevski.

Cenchrus brownii Roem. & Schult.

Fine-bristled burgrass.

Annual; tufted; 300-900 mm
tall. Leaf blades 80-150 mm
long; 8-10 mm wide. Spikelets
4-6 mm long; 4-6 mm wide. In-
florescence a false spike, 30-100
mm long; spikelet involucre in
two whorls, inner one with con-
nate spines which are plumose on their inner surface, the
outer one with spines which are bristle-like and short,
40-80 spines; burr with an inconspicuous disc at base.

Flowering January to June. Mainly sandy soil. Infre-
quent. Invader from tropical America. Biome: Savanna and
Grassland. Pantropical weed.

Description: Chippindall 1955 (451). Illustration: Chip-
pindall 1955 (375). Voucher: Smook 1906. PRECIS code
9901400-00200.

Cenchrus ciliaris L.

Buffelsgras.

Fig. 39. PI. 34.

Perennial; tufted; 600-1000
mm tall. Leaf blades 100-250
mm long; 4-8 mm wide. Spike-
lets 4—5 mm long; 3 mm wide. In-
florescence a bristly false spike,

40-120 mm long, straw or purple;
bristles mostly 5-10 mm long,

inner bristles slender and plumose, outer bristles slender
and scabrid, all the bristles are joined at base below the
spikelet cluster to form a small inconspicuous disc.

Flowering August to April. Common in hot dry areas,
especially on sandy soils, widespread elsewhere. Common.
Biome: Savanna, Grassland, and Nama-Karoo. Mainly
Africa, India and other hot drier areas of the world. Pasture
(cultivated). Variable species, with many cultivars
available. May be confused with Pennisetum foermerianum
which has an interrupted panicle, with bristles shorter and
plumose and with Enneapogon cenchroides, which has 9-
lobed lemmas with long awns and no bristles at the base
of the spikelet.

Description: Farming in South Africa leaflet 1 14 1983
A cultivation guide, Chippindall 1955 (451). Illustration:
Chippindall 1955 (fig. 374). Voucher: Smook 2822.
PRECIS code 9901400-00300.

Cenchrus incertus M.A. Curtis

(=C. pauciflorus Benth.) 2.

Annual; tufted; 100-400 mm
tall. Leaf blades 60-120 mm
long; 3-4 mm wide. Spikelets 5-7
mm long; 3-4 mm wide. Inflores-
cence open or compact 20-80 mm
long; burrs ovoid to globose with
clefts on two sides, with 8—40
spines of variable shape and size, the inner spines are more
regular with a plumose inner surface, while the outer spines
are connate and spreading in all directions.

Flowering January to March. Mainly sandy soil.
Infrequent. Invader from tropical America. Biome: Fynbos,
Savanna, and Grassland. Pantropical weed.

Annual, or perennial (with glaucous stems and leaves);
caespitose to decumbent. Culms 30-1500 mm high; herba-
ceous; unbranched above (but often branched near the
base). Leaf blades linear-lanceolate', flat, or rolled
(convolute). Ligule a fringe of hairs.

Inflorescence paniculate', contracted; espatheate (but
panicles enclosed by spathe-like upper leaf sheaths).
Spikelet-bearing axes persistent.

Spikelets 7-10 mm long; compressed laterally; disartic-
ulating above the glumes. Callus long. Glumes two; more
or less equal; much exceeding the spikelets4, awnless; similar
(papery). All florets female-fertile, or with distal incom-
plete florets, these merely underdeveloped, awned;
proximal incomplete florets absent.

Female-fertile florets 2-5. Lemmas similar in texture to
the glumes (papery); hairy (hairs in 6 to 8 bristle-tipped

81

tufts, in transverse rows, with a transverse row of tufts level
with the base of the awn, as well as longitudinal rows of
hairs); without a germination flap; 7-11 nerved; incised;
awned. Awns 1, or 3; median, or median and lateral (by
small straight extensions from the lobes). The median awn
different in form from the laterals (when laterals present);
from the sinus; geniculate; much shorter than the body of
the lemma to about as long as the body of the lemma. Palea
present; relatively long (almost equalling the lemma);
2-nerved. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Hilum short; pericarp fused; embryo large.

Photosynthetic pathway. C4; biochemichal type and
ultrastructure need investigating, in view of the peculiarity
of other C4 arundinoids and especially in view of the
variation in PCR sheath form; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x - 12. Arundinoideae; Danthonieae. 4 species.
North Africa, South and South West Africa and Middle
East. Xerophytic; in open habitats. Namibia, Botswana, and
Cape Province. 2 indigenous species.

References. 1. Cope. 1982. Kew Bull. 37: 657. 2.
Conert. 1962. Senck. Biol. 43: 239-266. 3. Ellis. 1984.
Bothalia 15: 153-159.

Species treatment by N.P. Barker.

1(0). Central awn of lemmas 3-5 mm long; panicle 30-120
mm long; spikelets 3(^l)-flowered; plants to 750

mm tall C. glauca

Central awn of lemmas 10-16 mm long; panicle
150-270 mm long; spikelets 4—6-flowered; plants
600-1500 mm tall C. mossamedensis

Centropodia glauca (Nees) T.A. Cope

Fig. 40. PI. 35.

( =Asthenatherum forskahlei
auctt., non (Vahl) Nevski) 1;

( =Asthenatherum glaucum
(Nees) Nevski) 1; (=Danthonia
glauca Nees) 1.

Weakly perennial, or annual;
tufted; 200-750 mm tall. Leaf
blades to 1 10 mm long; 8-10 mm
wide. Spikelets 7.5-10.0 mm long. Lower leaf sheaths loose
and densely hairy; panicle 30-120 mm long; spikelets
3(-4)-flowered, lower 2 bisexual, upper usually male;
glumes 6.5-10.0 mm long; central lemma awn 3-5 mm
long.

Flowering September to May. Loose, sandy substrates,
almost exclusively on dunes. Common (at foot of sand
dunes). Biome: Savannna, Nama-Karoo, Succulent Karoo
and Desert. Endemic. Tends to be annual on dunes and
perennial in gravel flats between dunes. Two varieties have
been described but are not recognised here because the
character upon which they are separated (leaf indumentum)
is variable with habitat, an observation supported by the
work of Ellis (1984).

Description: Conert 1962 (252), Launert 1970 ( 160:35),
Stapf 1898-1900 (534), Chippindall 1955 (246).

Illustration: Conert 1962 fig. 4—6, Chippindall 1955 (fig.
218.). Voucher: Ellis 4337. PRECIS code 9902035-00100.

Centropodia mossamedensis (Rendle) T.A. Cope

(=Asthenatherum
mossamedense (Rendle) Conert)
1; (=Danthonia mossamedensis
Rendle) 1.

Perennial; rhizomatous and
tufted; 600-1500 mm tall. Leaf
blades to 200 mm long; to 7 mm
wide. Spikelets 18-24 mm long.

Rhizome woody, bulbous, covered in hairy scales; lower
sheaths usually absent, but if present then not hairy; panicle
150-270 mm long; spikelets 4-6-flowered; glumes 17-24
mm long: central lemma awn 10-16 mm long.

Flowering March to June. Riverbeds and drainage lines.
Locally common. Biome: Nama-Karoo and Desert.
Anatomically similar to C. glauca (Ellis 1984).

Description: Conert 1962 (254), Launert 1970 (160:36).
Chippindall 1955 (246.). Illustration: Conert 1962 fig. 7-8.
Voucher: Oliver, Muller & Steenkamp 6711. PRECIS code
9902035-00200.

Chaetobromus Nees

Perennial; long-rhizomatous (sometimes), orcaespitose,
or decumbent. Culms 150-750 mm high; herbaceous;
branched above (but not profusely). Plants unarmed. Leaf
blades linear to linear-lanceolate; flat, or folded. Ligule a
fringe of hairs.

Inflorescence paniculate ( rarely racemose, in
depauperate plants)-, open, or contracted (sometimes with
few spikelets); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 10-17 mm long; compressed laterally;
falling with the glumes (the hairs on the persistent pedicel
allowing the spikelet to move in only one direction ). Callus
long. Glumes two; more or less equal; about equalling the
spikelets, or much exceeding the spikelets; awnless; similar
(subherbaceous, with scarious margins). Incomplete florets
distal to the female-fertile florets, merely underdeveloped;
proximal incomplete florets absent.

Fig. 41. Chaetobromus dregeanus

82

Female-fertile florets (2-)3-4(-6). Lemmas less firm
than the glumes to similar in texture to the glumes
(membranous); hairy (mostly), or hairless (LI ); 7-9 nerved;
incised; awned (but the LI sometimes with a reduced awn
or awnless). Awns l, or 3 ; median, or median and lateral
(the lateral lemma lobes sometimes bristle-tipped). The
median awn different in form from the laterals (when
laterals present); from the sinus (mostly), or apical
(sometimes, in the LI); geniculate (and twisted below);
much longer than the body of the lemma. Palea present; rel-
atively long; 2-nerved. Lodicules fleshy; glabrous. Stamens
3. Ovary glabrous. Hilum long-linear; pericarp fused.

Photosynthetic pathway. C3 (probably — though the
lateral cell count is low between all but a few bundles, at
least in C. dregeanus)', XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 2-3 species. Southern Africa. Xerophytic;
commonly maritime-arenicolous, or glycophytic (generally
on sandy soil). Namibia and Cape Province. 2 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Ellis.
1988. Bothalia. 18: 195-209.

This genus appears to be a polyploid series (2n = 12-72)
and is in a taxonomically chaotic state because of much
intra-taxon variation. It is divisible into two groups, based
on size differences in the floral parts, possibly related to
differences in chromosome numbers. (Spies, Du Plessis &
Barker, in prep.). The species accepted here are therefore
possibly artificial, and only two of the four published names
are used.

Species treatment by N.P. Barker.

1(0). Some or all basal leaves densely pubescent, hairs silky
and appressed; glumes 9- 1 2 mm long; lemma backs
of upper florets densely pubescent, hairs short,
erect; central awn of lemma of basal floret usually

not geniculate C. involucratus

Basal leaves glabrous or sparsely pubescent, hairs
then not appressed; glumes 12-17 mm long; lemma
backs of upper florets glabrous or pubescent, hairs
then lax and not very dense; central awn of lemma
of basal floret usually geniculate . . C. dregeanus

Chaetobromus dregeanus Nees

Fig. 41 . PI. 36.

Perennial; stoloniferous, or
tufted; to 600 mm tall. Leaf
blades to 270 mm long; 5 mm
wide (occasionally wider). Spike-
lets 12-18 mm long (excluding
awns); to 10 mm wide. Leaves
glabrous or sometimes sparsely
pubescent; spikelets 2-4-flower-
ed; glumes 12-17 mm long;
lemma back of basal floret glabrous or sparsely hairy,
lemma backs of upper florets glabrous or pubescent, the
hairs then lax; basal lemma body 4—5 mm long, lemma
lobes usually absent; body of upper lemmas 2. 8-5. 5 mm
long with lobes attenuating into bristles; central awn of bas-
al and upper florets usually geniculate.

Flowering August to November (occasionally later).
Sandy areas and rocky hillsides in low rainfall areas. Com-
mon (Strandveld, Namaqualand coastal belt and
Namaqualand Broken Veld, occasionally in Fynbos).
Biome: Fynbos, Nama-Karoo, Succulent Karoo, and Desert.
Endemic. Natural pasture. Ellis (1988) describes four
anatomical forms within the genus, three of which are in-
cluded under this concept of C. dregeanus.

Description: Stapf 1898-1900 (538), Chippindall 1955
(373). Illustration; Chippindall 1955 (fig. 246). Voucher:
Goldblatt 2558. PRECIS code 9902060-00100.

Chaetobromus involucratus (Schrad.) Nees

Perennial; stoloniferous, or
tufted; to 300 mm tall. Leaf
blades to 120 mm long; to 6 mm
wide. Spikelets 9-14 mm long
(excluding awns); 10 mm wide.

Some or all basal leaves and
sheaths covered in long, silky,
appressed hairs; spikelets 3-A-
flowered; glumes 9-12 mm long;
lemma back of basal floret glabrous, lemma backs of upper
florets densely covered in short, erect hairs; basal lemma
body 2. 5-4.0 mm long, lemma lobes absent; lemma body
of upper florets 1. 8-3.0 mm long, lobes attenuating into
bristles; central awn of basal floret usually not geniculate.

Flowering August to October (and occasionally later).
Coastal areas of the northern Cape. Locally common (Port
Nolloth). Biome: Succulent Karoo and Desert. Endemic.
Ellis (1988) considers the anatomical form corresponding
to this species to be anatomically distinct because of the
presence of silky macrohairs, which however are not visible
on all leaves of the plant, often appearing only on some of
the most basal leaves.

Description: Stapf 1898-1900 (537), Chippindall 1955
(274). Voucher: De Winter 9549. PRECIS code 9902060-
00200.

Chloris O. Swartz

Actinochloris Steud., Agrostomia Cer v.,Apogon Steud.,
Chloridopsis Hack., Chloropsis Kuntze, Chlorostis Raf.,
Geopogon Steud .. Heterolepis Boiss ..Leptochloris Kuntze,
Phacellaria Steud., Trichloris Benth.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 100-3000 mm
high; herbaceous. Leaf blades linear, flat, or folded, or
rolled. Ligule a fringed membrane to a fringe of hairs.

Inflorescence of spike-like main branches', digitate or
subdigitate (except C. roxburghana)\ espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary, or in pairs; biseriate; not in distinct
‘long-and-short’ combinations; 1.8-5. 5 mm long; com-
pressed laterally, disarticulating above the glumes (the
glumes usually persistent). Hairy callus present (usually
minute). Glumes two; very unequal; decidedly shorter than
the adjacent lemmas, or long relative to the adjacent
lemmas; awnless; similar to very dissimilar (narrow,
membranous, or the lower sometimes subulate). Incomplete
florets 2-5, distal to the female-fertile florets, merely under-
developed; proximal incomplete florets absent.

Female-fertile florets I (rarely 2). Lemmas similar in
texture to the glumes, or decidedly firmer than the glumes
(membranous or cartilaginous); 1-7 nerved; entire
(truncate), or incised; awned. Awns 1 (usually), or 3;
median, or median and lateral (rarely, Trichloris). The
median awn similar in form to the laterals (when laterals
present); from the sinus, or apical; non-geniculate. Palea
present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; ellipsoid (to
lanceolate); not noticeably compressed (subterete), or
trigonous ; hilum short; pericarp fused; embryo large (1/2
to 2/3 the grain length).

Photosynthetic pathway and related features. C4; PCK
(6 species); XyMS+. PCR sheath outlines uneven. PCR
sheath extensions usually absent, or present (in C. virgata).
Maximum number of extension cells in C. virgata 2. PCR
cell chloroplasts ovoid; with well developed grana;
centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato.
About 55 species. Tropical and warm temperate. Meso-
phytic, or xerophytic; in open habitats (diverse habitats,

83

mostly in short grassland on poor soil or disturbed ground).
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. Indigenous
species (7), naturalized species (1).

Intergeneric hybrids with Cynodon — X Cynochloris
Clifford & Everist: several species involved.

References. 1. Clayton et al. 1974. FTEA. 2. Renvoize.
1977. Kew Bull. 31:844. 3. Clayton. 1982. Kew Bull. 37:
419.

Species treatment by M. Koekemoer.

1 (0). Spikes numerous, arranged on a long central axis; axis

60-180 mm long C. roxburghiana

Spikes fewer than 20, digitate or subdigitate 2

2(1). Upper glumes with a distinct, dense rim of silky white
hairs on the margins; spikes stout and shorter than

30 mm C. flabellata

Upper glumes scabrid or glabrous on the margins;
spikes slender and usually longer than 30 mm . 3
3(2). Leaf blades distinctly or slightly rounded at the tips

4

Leaf blades acuminate or tapering to a fine point . 6
4(3). Leaf blades blunt and broadly rounded at the tips;
lemma awns 1 1-27 mm long .... C. pycnothrix
Leaf blades tapering to a fine rounded tip; lemma

awns 6-1 1 mm long 5

5(4). Spikes 6-10, long, slender, flexuous, spreading
almost horizontally, 70-150 mm long; plants

flowering in winter C. truncata

Spikes usually 4-6, rather coarse, firm, never
horizontally spreading, 30-80 mm long; plants

flowering in summer C. mossambicensis

6(3). Spikelets distant, more or less their own length apart;
spikes slender, flexuous; lowest spikes almost

horizontally spreading C. diluta

Spikelets imbricate, overlapping for most of their
length; spikes firm, usually erect but not spreading
more than 45 degrees from the central axis .... 7
7(6). Lemma tips concealed by a tuft of stiff, erect hairs;
awns 5-15 mm long, at least four times the length
of the body; spikelets delicate; uppermost leaf
sheaths often inflated around the young
inflorescences; plants not more than 900 mm tall

C. virgata

Lemma tips truncate; awn 2-10 mm long, shorter than
to as long as the body; spikelets rather coarse;
uppermost leaf sheath not inflated; plants 500-2200
mm tall C. gayana

Chloris diluta Renvoize

Perennial; rhizomatous and
stoloniferous; 300-1000 mm tall.

Leaf blades 100-300 mm long;

4-7 mm wide. Spikelets 3—4 mm
long. Culms wiry, usually erect;
leaf blades acuminate; spikes
4-6, 50-80 mm long; spikelets
more or less their own length
apart; lower glume 2. 0-2. 5
mm long; lowest lemma 3.5 mm long, with awn 2-8 mm
long.

Flowering March to June. In scrub forest on river banks.
Rare. Biome: Savanna. Zimbabwe. Similar to Chloris
pycnothrix, which has leaf blade tips broadly rounded and
lemma awns 1 1-27 mm long, and C. truncata, which has
leaf tips finely rounded and spikes 6-10.

Description: Renvoize 1977 (844). Voucher: Scheepers
1138. PRECIS code 9903010-00050.

Chloris flabellata (Hack.) Launert

Swardforming perennial; sto-
loniferous (stolons stout and
woody); 200-450 mm tall. Leaf
blades 50-80 mm long; 3-5 mm
wide. Spikelets 2. 5-3. 5 mm long.

Spikes stout, curved inwards,

1 0-30 mm long; lower glume 1 .5
mm long; upper glume with a
dense rim of white silky hairs;
lemma 2.5 mm long.

Flowering December to April. Confined to coastal
regions on saline marshes or flats, also in sandy, muddy
places and edges of reed beds. Rare. Biome: Desert. Coastal

84

regions of southern Angola. Erosion control. The silky edge
of the upper glume is very characteristic and does not occur
in any other southern African species of this genus.

Description: Launert 1970 Mitt. Bot. Munch. 8.
Voucher: Tinley 1626. PRECIS code 9903010-00100.

Chloris gayana Kunth

Rhodes grass.

Perennial; stoloniferous and
tufted; 500-1200 mm tall. Leaf
blades 250-500 mm long; 3-9
mm wide. Spikelets 3-5 mm long.

Basal leaf sheaths strongly
keeled; spikes 7-20, 40-150 mm
long; spikelets overlapping for
most of their length; lower glume 1.5-2. 5 mm long; lowest
lemma 2. 5-3. 5 mm long, awn 1-10 mm long, shorter than
or as long as the body of the lemma.

Flowering November to May. Riverine woodland to
open veld on well drained soils. Common. Naturalized from
India. Biome: Fynbos, Savanna, and Grassland. Tropical
Africa to China; cultivated pasture in Australia, New
Zealand and North America. Pasture (planted as forage).
Very similar to C. virgata, which is larger and has a tuft
of stiff, erect hairs on the lemma tips.

Description: Chippindall & Crook 1976 (2), Stapf
1898-1900 (642), Hitchcock & Chase 1950 (502), Chippin-
dall 1955 (197), Clayton et al. 1970-1982 (346).
Illustration: Chippindall 1955 (plate 7), Hitchcock & Chase
1950 (fig. 1065). Voucher: Scheepers 941. PRECIS code
9903010-00200.

Chloris mossambicensis K. Schum.

(= Tetrapogon

mossambicensis (K. Schum.)

Chippind. ex Fisher) 1.

Robust perennial; rhizomatous
and stoloniferous; 150-800 mm
tall. Leaf blades 100-350 mm
long; 3-6 mm wide. Spikelets 2-\
mm long. Basal leaf sheaths
strongly keeled; spikes usually 4-5, yellowish, 30-80 mm
long; spikelets coarse; lower glume 1 .7-2.0 mm long; upper
glume 2.75-3.50(-4.0) mm long; lowest lemma 2-3(-4)
mm long, awns two, 4-1 1 mm long.

Flowering October to April. Along rivers or on
seasonally flooded pans on clayey, waterlogged and turf
soils. Infrequent. Biome: Savanna and Grassland. Southern
tropical Africa.

Description: Clayton et al. 1970-1982 (341).

Illustration: Clayton et al. 1970-1982 (fig. 96(5)). Voucher:
Bredenkamp 1525. PRECIS code 9903010-00250.

Chloris pycnothrix Trin.

Fig. 42.

Spiderweb chloris. l

Usually annual, or perennial; \ J '>xj
tufted; 150-500 mm tall. Leaf (
blades 20-100 mm long; 3-5 mm \
wide. Spikelets 2-3 mm long.

Leaves with rounded and blunt \ Jr

tips; spikes 4-9, delicate, narrow, V— - —

40-100 mm long, horizontally

spreading; lower glume 1 .5—3.0 mm long; lowest lemma
2. 5-3. 2 mm long, tips acuminate to acute, awns 1 1-27 mm
long.

Flowering September to May. In cultivated lands,
disturbed areas and on roadsides in shallow stony soils.
Common. Biome: Savanna and Grassland. Tropical Africa,
tropical South America. Erosion control (pioneer), or weed
(in disturbed places). Similar to C. diluta , in which the

spikelets are distant, about their own length apart and the
lemma awn 2-8 mm long, and C. truncata, which has leaf
blades with fine round tips and lemma awns 6-1 1 mm
long.

Description: Chippindall & Crook 1976 (3), Stapf
1898-1900 (641), Chippindall 1955 (198), Clayton et al.
1970-1982 (340). Illustration: Chippindall 1955 (fig. 173).
Voucher: Smook 5593. PRECIS code 9903010-00350.

Chloris roxburghiana Schult.

(=C. myriostachya
Hochst.) 1.

Perennial; rhizomatous and
tufted; 700-1250 mm tall. Leaf
blades 100-400 mm long; 2-10
mm wide. Spikelets 1-3 mm long.

Panicle dense, axis 40-100 mm
long; spikes numerous, 30-80
mm long; spikelets 3^4- flowered; lower glume 1.0-1. 5 mm
long; upper glume 2. 0-2. 8 mm long; lowest lemma 1. 5-2.0
mm long, awns 8-17 mm long.

Flowering November to May. Dry, sandy or stony soil
on river banks, in open veld or disturbed places. Locally
common. Biome: Savanna. Central tropical Africa and
southern India. The inflorescence is strikingly different
from other southern African Chloris species, which have
digitate or subdigitate racemes.

Description: Chippindall 1955 (196), Clayton et al.
1970-1982 (338). Illustration: Chippindall 1955 (fig. 171).
Voucher: Godfrey SH 1729. PRECIS code 9903010-00500.

Chloris truncata R. Br.

Perennial; stoloniferous and
tufted; 250-450 mm tall. Leaf
blades 30-200 mm long; 2-3 mm
wide. Spikelets 2 mm long.

Spikes 6-10, 80-150 mm long,
flexuous, spreading horizontally;
spikelets overlapping about half
their length; awns 6-12 mm long.

Flowering June to July.

Disturbed places and lucerne paddocks. Infrequent. Nat-
uralized, or invader (possibly) from Australia. Biome; Fyn-
bos. Ornamental (occasionally cultivated in grass gardens),
or weed (in lucerne). Introduced weed that seems to be
spreading in the winter rainfall area. Similar to C.
pycnothrix , which has longer awns, and C. diluta, which has
spikelets about their own length apart.

Description: Hitchcock & Chase 1950 (509). Voucher:
P.C.V. du Toil 2172. PRECIS code 9903010-00550.

Chloris virgata Swartz

PI. 37.

Feathered chloris, klossiegras.

Usually annual, or perennial;
tufted; 300-750 mm tall. Leaf
blades 100-300 mm long; 2-6
mm wide. Spikelets 3. 0-3. 5 mm
long. Upper leaf sheaths inflated;
spikes 7-15, 20-80 mm long,
erect, silky-feathery; lower glume
1.5-2. 5 mm long; upper glume 2. 5^4. 5 mm long; lowest
lemma 2. 5-4.0 mm long, with a crown of spreading hairs
1. 5^4.0 mm long at the apex; awns 2-15 mm long.

Flowering December to June. Disturbed places on a
variety of soil types. Common. Biome: Savanna, Grassland,
Nama-Karoo, Succulent Karoo, and Desert. Worldwide in
tropical and temperate countries. Pasture (good grazing and
hay), or weed (disturbed areas). Very similar to C. gayana,
which lacks the tuft of hairs on the lemma tips.

Description: Chippindall & Crook 1976 (4), Stapf
1898-1900 (641), Hitchcock & Chase 1950 (504), Chippin-

85

dall 1955 (197), Clayton et al. 1970-1982 (343).
Illustration: Chippindall 1955 (fig. 172), Clayton et al.
1970-1982 (fig. 97). Voucher: Smook 5122. PRECIS code
9903010-00600.

Chrysopogon Trin.

Centrophorum Trin., Chalcoelytrum Lunell, Pollinia
Spreng., Raphis Lour., Trianthium Desv.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 150-1500 mm
high; herbaceous; usually unbranched above. Ligule a
fringed membrane (short), or a fringe of hairs. Plants
bisexual, with bisexual spikelets; with hermaphrodite
florets. The spikelets of sexually distinct forms on the same
plant ; overtly heteromorphic (pedicellate spikelets flattened
dorsally, awnless or not: often the sessile spikelet pallid or
yellowish, the pedicellate spikelet purple).

Inflorescence paniculate; open ( with whorls of slender,
persistent branches); espatheate; not comprising ‘partial
inflorescences' and foliar organs. Spikelet-bearing axes
very much reduced (usually to a single joint and the
terminal triad, but sometimes with a long-pedicel! short-
pedicel pair below)\ with very slender rachides; disarticu-
lating at the joints (beneath the triad, and beneath the pairs
when present). ‘Articles’ without a basal callus-knob.

Spikelets in triplets, or in triplets and in pairs; consis-
tently in Tong-and-short’ combinations; these pedicellate/
sessile. Pedicels free of the rachis. The sessile spikelets her-
maphrodite. The pedicellate spikelets male-only or sterile,
on slender flat pedicels, dorsally compressed, awned or
awnless, LI empty, L2 usually with a male floret. Female-
fertile spikelets 5—8.5 mm long; compressed laterally,
falling with the glumes. Glumes two; more or less equal;
awned and awnless (G2 often awned), or awnless. Lower
glume convex on the back (or keeled upwards, sometimes
with spinulose margins). Proximal incomplete florets T,
epaleate; sterile.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline); entire, or incised; awned. Awns 1;
median; from the sinus, or apical; geniculate; much shorter
than the body of the lemma, to much longer than the body
of the lemma (?). Palea present, or absent; when present
conspicuous but relatively short, or very reduced. Lodicules
2; fleshy; glabrous. Stamens 3. Ovary glabrous; hilum
short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 25 species. Tropical and
subtropical. Mesophytic, or xerophytic (from rainforest to
subdesert); in open habitats (on poor soils, often in
disturbed ground); glycophytic. Botswana, Transvaal, and
Cape Province. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Chrysopogon serrulatus Trin.

(=C. montanus Trin. var.
tremulus (Hack.) Stapf) 1.

Krulgras, golden beard grass.

Perennial; sometimes rhizo-
matous and tufted; to 1000 mm
tall. Leaf blades to 300 mm long;

2-10 mm wide. Spikelets (sessile
and pedicellate) 5-8 mm long (but sessile laterally
compressed, pedicellate dorsally compressed). Inflores-
cence branches long, whorled, bare below, terminated by
a triad of spikelets.

Flowering December to April. Rocky hillsides, stony
soils. Biome: Savanna, Grassland, and Nama-Karoo.
Eastern Africa to tropical Asia.

Description: Chippindall 1955 (468), Clayton et al.
1970-1982 (736). Illustration: Chippindall 1955 (fig. 384).
Voucher: Brueckner 121. PRECIS code 9900500-00200.

Cladoraphis Franch.

A segregate of Eragrostis.

Perennial; long-rhizomatous, or long-rhizomatous and
long-stoloniferous (occasionally). Culms 200-800 mm
high-, woody and persistent ; branched above. Plants con-
spicuously armed (with pungent tipped leaf blades and
inflorescence axes). Leaf blades linear-lanceolate to
lanceolate; becoming rolled; hard, woody, needle-like.
Ligule a fringe of hairs. The spikelets all alike in sexuality.

Inflorescence paniculate (with distant branches, or
reduced to a single branch or cluster); open (but the lateral
branches compact); espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary; not two-ranked; 7-16 mm long; com-
pressed laterally; disarticulating above the glumes and
between the florets (tardily). Glumes two; more or less
equal; markedly shorter than the spikelets; awnless; similar.
Incomplete florets distal to the female-fertile florets, merely
underdeveloped, awnless; proximal incomplete florets
absent.

Female-fertile florets 3-16. Lemmas similar in texture
to the glumes; without a germination flap; 3 nerved; entire;
awnless (muticous). Palea present; relatively long
(equalling the lemmas or slightly shorter). Lodicules 2;
fleshy. Stamens 3. Ovary glabrous. Fruit small (1.3 to 2 mm
long); hilum short; pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven (owing to the adaxial
extensions). PCR sheath extensions present (with most

Fig. 43. PI. 38.

86

bundles). Maximum number of extension cells 4-5. PCR
cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Southern Africa. Xero-
phytic; in open habitats; maritime-arenicolous (C.
cyperoides on beach dunes), or glycophytic (C. spinosa on
desert dunes and sandy beds of dry watercourses). Namibia
and Cape Province. 2 indigenous species.

References. 1. Phillips. 1982. Kew Bull. 37:133.

1(0). Inflorescences with primary branches less than their
own length apart; primary branches spiny; spikelets
3-18-flowered, arranged on short intervals, usually
almost perpendicular to the branches . C. spinosa
Inflorescence with primary branches more than their
own length(and often more than twice their length)
apart; primary branches not always developed to a
spine; spikelets 4-9(-20)-flowered, usually

clustered and appressed to the branches

C. cyperoides

Cladoraphis cyperoides (Thunb.) S.M. Phillips

(-Eragrostis cyperoides
(Thunb.) Beauv.) 1 .

Sedge-stemmed love grass,
steekriet.

Spiny, bushy perennial; tuft-
ed; 200-800 mm tall. Leaf blades
20-110 mm long; 4-9 mm wide.

Spikelets 4—8 mm long; 3-5 mm wide. Panicle branches
more than their own length apart (often more than twice
their length), not always produced as a spine, up to 80 mm
long but usually much shorter; spikelets usually clustered
and appressed to the branches, 4—9(-20)-flowered.

Flowering August to May (peak flowering from August
to October). Deep loose sand, coastal dunes or on the edges
of fresh or saltwater lagoons. Locally common (coast).
Biome: Fynbos, Succulent Karoo, and Desert. Endemic,
probably cultivated in Oregon USA. Occasionally grazed
pasture and erosion control (stabilizing windblown dunes).

Description: Stapf 1898-1900 (611), Hitchcock &
Chase 1950 (168), Chippindall 1955 (184). Voucher:
Goldblatt 4244. PRECIS code 9902865-00100.

Cladoraphis spinosa (L. f.) S.M. Phillips

Fig. 44. PI. 39.

(= Eragrostis spinosa (L. f.)

Trin.) 1.

Spiny love grass, volstruis-
doring.

Spiny, bushy perennial; tuft-
ed; 200-600 mm tall. Leaf blades
10-60 mm long; 4-9 mm wide.

Spikelets 6-18 mm long; 3-4 mm wide. Panicle branches
less than their own length apart, spine-tipped, up to 50 mm
long; spikelets usually almost perpendicular to the
branches, arranged at short intervals, 3-18-flowered.

Flowering August to May (peak flowering from August
to October). Well-drained, deep, loose sand on dunes and
river margins. Locally common. Biome: Fynbos, Nama-
Karoo, Succulent Karoo, and Desert. Endemic.
Occasionally grazed pasture, or erosion control (stabilizing
windblown sand dunes), or indicator (overgrazed veld).

Description: Stapf 1898-1900 (612), Chippindall 1955
(183). Illustration: Chippindall 1955 (fig. 158). Voucher:
Ward 163. PRECIS code 9902865-00200.

Cleistachne Benth.

Annual. Culms 600-2500 mm high; herbaceous; usually
unbranched above (sometimes with stilt roots). Leaf blades
linear; rolled. Ligule an unfringed membrane (scarious).
Plants bisexual, with bisexual spikelets.

Inflorescence paniculate ; large, terminal, linear to
lanceolate; espatheate; not comprising ‘partial inflores-
cences’ and foliar organs. Spikelet-bearing axes ‘racemes'
(long, narrow, with many joints); with very slender
rachides; persistent. ‘Articles’ densely long-hairy, or
somewhat hairy (rachis and pedicels with grey or brown
hairs).

Spikelets solitary (perhaps representing ‘racemes’
reduced to single spikelets: cf. Sorghum , Sorghastrum.
Female-fertile spikelets 4-5 mm long (rarely 3 or 6 mm);
compressed dorsiventrally, falling with the glumes
(disarticulating from apex of pedicel). Glumes two; more
or less equal ; awnless; similar (leathery, with inrolled
margins). Proximal incomplete florets /; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); incised; awned. Awns 1; median; from

87

the sinus; geniculate; much longer than the body of the
lemma. Palea present (but small); conspicuous but rela-
tively short. Lodicules 2; fleshy; ciliate. Stamens 3. Ovary
glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 1 species. Tropical Africa, India.
Helophytic to mesophytic; in open habitats (riverbanks and
old farmland); glycophytic. .Transvaal. 1 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Fig. 45. Cleistachne sorghoides

Cleistachne sorghoides Benth.

Fig. 45. PI. 40.

Coarse, robust annual; to 2500
mm tall. Leaf blades to 1000 mm
long; to 14 mm wide. Spikelets
4-5 mm long. Spikelets all alike,
pedicellate, not paired, dark and
glossy at maturity.

Flowering February to April.

Riverbanks and vleis. Conserva-
tion status not known. Biome: Sa-
vanna and Grassland. Through eastern tropical Africa to
India.

Description: Chippindall 1955 (468). Illustration: Chip-
pindall 1955 (fig. 383), Clayton et al. 1970-1982 (fig. 170).
Voucher: Vermeulen April 1952. PRECIS code 9900480-
00100.

Coelachyrum Hochst. & Nees

Sometimes includes Cypholepis Chiov.

Perennial; caespitose (densely). Culms 300-1000 mm
high; herbaceous. Leaf blades linear; usually flat. Ligule a
fringed membrane.

Inflorescence of spike-like main branches', non-digitate;
distant, erect. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; short pedicellate', 5-10 mm
long; compressed laterally; disarticulating above the
glumes; disarticulating between the florets. Glumes two;
more or less equal; long relative to the adjacent lemmas',
awnless; similar (lanceolate, membranous). Incomplete
florets distal to the female-fertile florets, male; proximal in-
complete florets absent.

Female-fertile florets 7-10. Lemmas decidedly firmer
than the glumes (membranous, becoming cartilaginous
below); hairy (pilose with club-shaped hairs on the lower
back); 3 nerved; entire; awnless. Palea present; conspicuous

Fig. 46. Coelachyrum yemenicum

but relatively short (about half lemma length). Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous (?). Fruit small
(1.2-1. 4 mm); hilum short; pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. Northeast to southeast and
southern Africa. Xerophytic; in open habitats. Namibia,
Botswana, Transvaal, and Cape Province. 1 indigenous
species.

References. 1. Phillips. 1982. Kew Bull. 37; 133.

Species treatment by M. Koekemoer.

Coelachyrum yemenicum (Schweinf.) S.M. Phillips

(-Cypholepis yemenica
(Schweinf.) Chiov.) 1.

Perennial; slender, densely
tufted (culms erect or geniculate);

310-630 mm tall. Leaf blades
70-320 mm long; 2. 5-5. 5 mm
wide. Spikelets 5-10 mm long.

Leaf sheaths keeled; racemes
2-8, far apart; spikelets 7-12-flowered; lemma with club-
shaped hairs near the base.

Flowering February to June. Calcareous pans, shallow
limestone, often in light shade. Infrequent. Biome; Savanna
and Nama-Karoo. Eastern Africa to Yemen.

Description: Chippindall 1955 (121), Clayton et al.
1970-1982 (248). Illustration: Chippindall 1955 (fig. 94),
Clayton et al. 1970-1982 (fig. 69). Voucher: Paton 3156.
PRECIS code 9903360-00100.

Fig. 46. PI. 41.

Coelorhachis Brongn.

Apogonia Nutt, Cycloteria Stapf.

Perennial; mostly robust, tall, forming clumps. Culms
700-4000 mm high\ herbaceous (coriaceous); branched
above. Leaf blades linear; flat (or rarely filiform). Ligule a
fringed membrane to a fringe of hairs. Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant ; overtly heteromorphic, or homomorphic.

Inflorescence of spike-like main branches, or paniculate
(oflong-peduncled, spikelike ‘racemes' , solitary at culm or
branchlet apices, often in ‘false panicles' ); spatheate ; a
complex of ‘partial inflorescences’ and intervening foliar
organs (the unit consisting of ‘raceme’, its peduncle,
subtending leaf and next internode (peduncle of the unit)).
Spikelet-bearing axes spike-like\ solitary, or clustered
(fascicled); with substantial rachides; disarticulating at the
joints. ‘Articles’ non-linear (concave, clavate, shorter than
the sessile spikelet); with a basal callus-knob.

Spikelets in pairs; consistently in Tong-and-short' com-
binations; these pedicellate/sessile. Pedicels free of the
rachis (but closely contiguous). The sessile spikelets her-
maphrodite. The pedicellate spikelets hermaphrodite
(rarely), or male-only, or sterile. Female-fertile spikelets
3-4.5 mm long; compressed dorsiventrally; falling with the
glumes. Glumes two; more or less equal; awnless; very dis-
similar (lower two-keeled and two-winged above, upper 1-
keeled and wingless). Proximal incomplete florets 7;
epaleate, or paleate, palea reduced; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present; relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Andropogonodae; Andropo-
goneae; Rottboelliinae. About 20 species. Mainly tropical.

Helophytic to mesophytic; in open habitats (grassland and
savanna, often on damp soils); glycophytic. Natal and Cape
Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Veldkamp. 1986. Blumea 31: 281.

Species could be transferred to Mnesithea Kunth.

Species treatment by G.E. Gibbs Russell.

Coelorhachis capensis Stapf

Fig. 47. PI. 42.

Perennial; tufted; to 700 mm
tall. Leaf blades 3-8 mm wide.

Spikelets (sessile) 4.5-5 mm long
(pedicellate smaller, sometimes
much reduced). Raceme narrowly
cylindrical, culm-like, with
sunken spikelets.

Flowering September to
March. Grassveld. Infrequent.

Mozambique.

Description: Chippindall 1955 (523). Illustration: Chip-
pindall 1955 (fig. 417). Voucher: Sim 2733. PRECIS code

89

Coix L.

Lacryma Medik ,Lacryma-jobi Ort., Lacrymaria Fabric.,
Sphaerium Kuntze.

Annual to perennial; stems erect or straggling, prop-
roots from the lower nodes. Culms 700-4000 mm high; her-
baceous; branched above. Leaf blades lanceolate; flat.
Ligule an unfringed membrane to a fringed membrane.
Plants monoecious with all the fertile spikelets unisexual.
The spikelets of sexually distinct forms on the same plant;
overtly heteromorphic.

Inflorescence paniculate (but peculiar — see below);
spatheate; a complex of ‘partial inflorescences’ and inter-
vening foliar organs (the partial inflorescences of peculiar
form, on flattened peduncles, in leafy panicles). Spikelet-
bearing axes very much reduced ( the female ‘raceme’
usually represented by three spikelets, enclosed in a
globose, hardened involucre or utricle 6-12 mm long,
separated from the male raceme by a prophyll at its base.
Male raceme exserted on a peduncle through the apex of
the involucre ); disarticulating; falling entire (within its
involucre).

Spikelets consistently in Tong-and-short’ combinations;

these pedicellate/sessile (in both male and female racemes).
The sessile spikelets of the female racemes female-only.
The pedicellate spikelets of the female racemes female-
only, or sterile. Female-fertile spikelets falling with the
glumes. Glumes two; more or less equal; awnless; very dis-
similar (both beaked; the lower subglobose, hyaline below,
subcartilaginous above; the upper narrower, strongly
keeled, subhyaline). Proximal incomplete florets 1;
epaleate; sterile. Male spikelets in pairs or triads, several
per disarticulating male raceme; dorsally compressed, with
two florets, both male or the lower sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes, or similar in texture to the glumes (similar to the
upper glume, but less strongly keeled; very thin and hyaline
beneath the beak); entire; mucronate (beaked). Palea
present; conspicuous but relatively short (broad, beaked).
Stamens 0 (or 3 staminodes). Ovary glabrous. Fruit medium
sized; hilum short (circular or elliptical, quite large);
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5. Panicoideae; Andropogonodae; Maydeae. 5
species. Tropical Asia. Helophytic to mesophytic; in shade,
or in open habitats (forest margins and swamps);
glycophytic. Transvaal, Natal, and Cape Province. 1
naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Coix lacryma-jobi L.

Fig. 48. PI. 43.

Job’s tears.

Annual; 900-1200 mm tall.

Leaf blades 100-500 mm long;

2-7 mm wide. Spikelets 7-10 mm
long (unisexual). Inflorescence
involucre is hard, whitish and
beadlike.

Flowering August to April.

Damp places. Infrequent. Naturalized (cultivated in warm
areas worldwide); originally from the East Indies. Domestic
use (beads), or weed (ruderal).

Description: Chippindall 1955 (504), Clayton et al.
1970-1982 (857). Illustration; Chippindall 1955 (fig. 419),
Clayton et al. 1970-1982 (fig. 205). Voucher: Smook 1298.
PRECIS code 9900020-00100.

Colpodium Trin.

Including Paracolpodium, Keniochloa.

Perennial; long-rhizomatous, or long-stoloniferous, or
caespitose, or decumbent. Culms 100-300 mm high; herba-
ceous; unbranched above. Sheath margins joined to free.
Leaf blades flat. Ligule an unfringed membrane .

Inflorescence paniculate ; open\ espatheate. Spikelet-
bearing axes persistent.

Spikelets 2-8 mm long; compressed laterally to not
noticeably compressed; disarticulating above the glumes.
Hairy callus absent. Glumes two; relatively large; more or
less equal; decidedly shorter than the adjacent lemmas, or
long relative to the adjacent lemmas', awnless; similar. All
florets female-fertile, or distal incomplete florets also
present, merely underdeveloped, awnless', proximal incom-
plete florets absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes; 3-5 nerved;
incised; awnless. Palea present; relatively long. Lodicules
2; membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
medium sized; hilum short', embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 2. Pooideae; Poodae; Poeae. Sensu stricto, 3
species (?). North temperate. Natal and Lesotho. 1 indige-
nous species.

References. 1. Clayton. 1970. FTEA. 2. Linder. Unpubl.
ms, FSA.

Species treatment by M. Koekemoer.

90

Colpodium hedbergii (Meld.) Tzvel.

Fig. 49. PI. 44.

( =Catabrosia aquatica auctt.,
non (L.) Beauv.) 1 .

Perennial; hydrophyte, stolon-
iferous, and tufted; 100-250
(-300) mm tall. Leaf blades
20— 60(— 1 50) mm long; 3-5 mm
wide. Spikelets 2. 5-4.0 mm long.

Leaf blades strongly keeled,
folded when young; panicle 40-120 mm long with
spreading branches which have the lower part bare; spike-
lets 1 -flowered; glumes subequal, slightly longer than the
lemma.

Flowering December to March. Wet places, in streams
and sedge meadows at high altitudes, 2900-4000 m. Rare.
Locally common. Biome: Grassland. Kenya. The South

African specimens do not fit satisfactorily into this species
and Hedberg (pers. comm.) regards our specimens as
representing a new taxon.

Description: Linder (57), Clayton et al. 1970-1982 (51 ).
Voucher: Killick 4414. PRECIS code 9904100-00100.

Cortaderia Stapf

Moore a Lemaire.

Perennial; caespitose (mostly large, tussocky). Culms
1000-4000 mm high. Leaf blades disarticulating from the
sheaths (the sheaths disintegrating or rolling). Ligule a
fringe of hairs. Plants bisexual, with bisexual spikelets, or
dioecious (being exclusively gynodioecious). The spikelets
all alike in sexuality (i.e., on the same plant).

Inflorescence paniculate; open', espatheate. Spikelet-
bearing axes persistent.

Spikelets 10-18 mm long; compressed laterally; disar-
ticulating above the glumes. Callus long. Glumes two; more
or less equal; about equalling the spikelets; awnless;
similar. Incomplete florets distal to the female-fertile
florets, merely underdeveloped; proximal incomplete
florets absent.

Female-fertile florets 2-3 (-5). Lemmas similar in
texture to the glumes to decidedly firmer than the glumes
(membranous); hairy (hairs in tufts, or not in tufts; in trans-
verse rows, or not in transverse rows); 3 nerved; entire, or
incised; awned. Awns 1 , or 3; median, or median and lateral
(via the lateral lobes). The median awn similar in form to
the laterals (or somewhat more flattened, when laterals
present); from the sinus, or apical; non-geniculate to genic-
ulate. Palea present; relatively long; 2-nerved. Lodicules 2;
fleshy; ciliate. Stamens 3, or 0 (in female plants of
dioecious species). Ovary glabrous. Hilum long-linear;
pericarp fused; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, a = 9. Arundinoideae; Danthonieae. 24 species.
New Zealand. South America. Mesophytic to xerophytic;
in open habitats (on hillsides, among scrub and in weedy
places). Transvaal, Natal, and Cape Province. 2 naturalized
species.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Robinson. 1984. S. Afr. J. Bot. 3: 343.

Species treatment by E.R. Robinson & G.E. Gibbs
Russell.

1(0). Leaves bright green, tips not setaceous, blades flat or
slightly V-shaped in cross-section, reaching only to
1/2 culm height; glume veins and rachillas purple

C.jubata

Leaves glaucous, tips setaceous, blades often V-
shaped in cross section, usually reaching more than
2/3 culm height; glume veins and rachillas white
C. selloana

Cortaderia jubata (Lem.) Stapf

Perennial; tufted (densely); to
3000 mm tall. Leaf blades
1000-1500 mm long; 9-12 mm
wide. Robust, leaves mostly bas-
al, to half the height of the culm,
bright green, with cutting margins
and midrib, flat or only slightly
V-shaped in cross section, tips
not setaceous; panicle 600-800
mm long; ( 1— )3— 5 florets per spikelet, all florets are female

91

Fig. 50. Cortaderia selloana

only; glumes approximately equal, 9-13 mm long; glume
veins, rachises and rachillas purple; lemmas 8-1 1 (— 15) mm
long.

Flowering November to February (rarely to March).
Disturbed places. Invader from high Andes of South
America. Erosion control (on mine dumps), or ornamental
(widely cultivated), or weed. Reproduces by agamospermy.

PRECIS code 9902110-00050.

Cortaderia selloana (Schult.) Aschers. & Graebn.

Fig. 50. PI. 45.

Perennial; densely tufted; to
4000 mm tall. Leaf blades
800-1800 mm long; 8-10 mm
wide.. Robust, leaves mainly bas-
al, glaucous, to 2/3 culm height;
blades with cutting margins and
midrib, usually markedly Y--
shaped in cross section, tips
setaceous; panicle 400-600
(-700) mm long; spikelets either female or hermaphrodite

(but then functionally male); female-fertile spikelets with
(5— )6(— 7) florets per spikelet, hermaphrodite spikelets with
( 1— )3( — 4) florets; glumes of equal length, 8-15 mm long;
glume veins, rachises and rachilla colourless; lemmas of
female-fertile florets 10-14 mm long, those of
hermaphrodite florets 12-15 mm long.

Flowering February and April. Seasonally wet habitats.
Invader (in the PWV area) from low lying riverbanks in
South America. Ornamental and weed (spreading in the
southern Transvaal).

Description: Chippindall 1955 (230). Illustration: Chip-
pindall 1955 (fig. 204). PRECIS code 99021 10- 00100.

Corynephorus P. Beauv.

Including Anachortus.

Annual, or perennial; caespitose. Culms 100-600 mm
high; herbaceous; unbranched above. Leaf blades linear;
folded, or rolled. Ligule an unfringed membrane.

Inflorescence paniculate; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 3-5 mm long; compressed laterally; disarticu-
lating above the glumes. Glumes two; more or less equal;
about equalling the spikelets; awnless; similar (lanceolate).
All florets female-fertile; proximal incomplete florets
absent.

Female-fertile florets 2. Lemmas similar in texture to the
glumes; 1 nerved; incised; awned. Awns 1; median ( with

92

a clavate apex enclosed by the glumes, and with a ring of
minute hairs distal to the twisted lower half)', dorsal; genic-
ulate; much longer than the body of the lemma. Palea
present; relatively long. Lodicules 2; membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit small; hilum
short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 5 species.
Europe, Mediterranean. Xerophytic; in open habitats (in
sandy places, often coastal). Cape Province. 1 naturalized
species.

References. 1. Tutin. 1980. FI. Europ.

Species treatment by M. Koekemoer.

Corynephorus fasciculatus Boiss. & Reut.

Fig. 51. PI. 46.

Annual; culms solitary or
loosely tufted; 200-550 mm tall.
Leafblades50-120mmlong; 1-2
mm wide. Spikelets about 3 mm
long. Hairs at the base of the flo-
ret not longer than 1/4 the lemma
length; lemma awn basal, usually
about as long as the glumes,
divided into a lower dark-
coloured column and a clavate upper limb, with a ring of
fleshy hairs at the junction.

Flowering October to November. Sandy soils in
disturbed places. Rare. Naturalized from Europe. Biome:
Fynbos. Portugal to the western Mediterranean. Similar to
the European species, C. divaricatus, which has larger
spikelets, the awn distinctly shorter than the glumes and
longer hairs at the base of the floret.

Description: Tutin 1980 (5: 231). Voucher; Van
Rensburg 139. PRECIS Code 9901880-00100.

Craspedorhachis Benth.

Perennial; often stoloniferous. Culms to 1000 mm high;
herbaceous. Ligule a fringed membrane to a fringe of hairs.

Inflorescence of spike-like main branches (several
slender spikes, usually on a long axis)-, digitate or subdigi-
tate, or non-digitate; espatheate. Spikelet-bearing axes
spikes', disarticulating (in C. africana, the peduncle bears
a cupular disarticulation zone at the point of origin of the
spikes, which seem to fall together), or persistent (?);
falling entire.

Spikelets solitary; biseriate; 3 mm long', compressed dor-
siventrally, disarticulating above the glumes. Glumes two;
more or less equal (long); awnless; very dissimilar (both
long, membranous, the lower asymmetrically 1 -keeled, the
upper flat-backed, infolded with two keels). All florets
female-fertile; proximal incomplete florets absent.

Female -fertile florets 1 . Lemmas less firm than the
glumes (hyaline); without a germination flap; 3 nerved;
awnless to mucronate. Palea present; relatively long. Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small (c. 1 mm); obovoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 5-6 species. Tropical Africa, North
and South America. Mesophytic to xerophytic; in open
habitats (sandy savanna); glycophytic. Namibia and
Botswana. 2 indigenous species.

References. 1. Launert. 1970. FSWA. 2. Clayton &
Renvoize. 1986. Gen. Gram.

Species treatment by L. Smook.

1(0). Lemma and palea glabrous C. rhodesiana

Lemma and palea with long hairs C. africana

Craspedorhachis africana Benth.

PI. 47.

Perennial; shortly rhizomatous
and tufted (densely, erect); to
1200 mm tall. Leaf blades to 200
mm long; to 5.5 mm wide. Spike-
lets 3-A mm long. Racemes gen-
erally shorter than 90 mm; lemma
and palea with long hairs.

Flowering January to April. In
sandy soils. Rare (in southern
Africa). Biome: Savanna. Zimbabwe, Mozambique,

Zambia, Madagascar. Closely related to C. rhodesiana,
which has short spikelets and glabrous lemmas.

Description: Chippindall & Crook 1976 (209). Voucher:
Story 6452 (in K). PRECIS code 9903090-00050.

Fig. 52. Craspedorhachis rhodesiana

Craspedorhachis rhodesiana Rendle

Fig. 52.

Perennial; shortly rhizomatous
and tufted (densely and erect); to
1200 mm tall. Leaf blades to 200
mm long; to 2.5 mm wide. Spike-
lets 2. 5-3. 2 mm long. Racemes
generally longer than 90 mm;
lemma and palea glabrous.

Flowering December, Feb-
ruary and March. Sandy soils or

93

sandy loam along pan edges, in and along dry river beds
and on sand dunes. Infrequent. Biome: Savanna. Angola,
Zimbabwe, Mozambique, Zambia. Closely related to C.
africana, which has slightly larger spikelets and lemmas
with long hairs.

Description: Chippindall & Crook 1976 (209), Launert
1970 (160:49). Voucher: Wild & Drummond 7043. PRECIS
code 9903090-00100.

Ctenium Panzer

Aplocera Raf., Campuloa Desv., Campulosus Desv.,
Monathera Raf .,Monocera Elliott, Triatherus Raf.

Perennial (rarely annual ); caespitose (densely). Culms
400-1000 mm high; herbaceous. Leaf blades linear; flat, or
rolled (convolute). Ligule a fringed membrane ( very short).
Plants bisexual, with bisexual spikelets. The spikelets all
alike in sexuality.

Inflorescence a single spike, or of spike-like main
branches ; spikes pectinate, usually curved; non-digitate, or
digitate or subdigitate; espatheate. Spikelet-bearing axes
spikes ; persistent.

Spikelets solitary; biseriate (along the midrib of the
rachis); 4-9 mm long; adaxial ; compressed laterally ; disar-
ticulating above the glumes; not disarticulating between the
florets. Glumes present; two; very unequal; awned (G2
shortly awn-tipped, and with a spreading awn from the
middle of its back); very dissimilar (G2 larger, firmer,
awned). Incomplete florets both distal and proximal to the
female-fertile florets; distal incomplete florets merely un-
derdeveloped (male or barren); proximal incomplete florets
2; male, or sterile. The proximal lemmas awned (from just
below tip).

Female-fertile florets 1. Lemmas less firm than the
glumes (i.e., than G2 — membranous); 3 nerved; entire;
awned. Awns 1; median; dorsal; non-geniculate; about as
long as the body of the lemma to much longer than the body
of the lemma. Palea present; relatively long. Lodicules 2;
fleshy; glabrous. Stamens 3 (2 in male florets). Ovary
glabrous. Fruit ellipsoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR cell chloroplasts
centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 9. Chloridoideae; Chlorideae sensu lato. 20
species. Tropical and subtropical America and Africa. In
open habitats (savanna). Transvaal, Swaziland, Natal,
Lesotho, and Cape Province. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Ctenium concinnum Nees

Fig. 53. PI. 48.

Sickle grass.

Wiry perennial; tufted; 400-
700 mm tall. Leaf blades 100-
300 mm long; 2-5 mm wide.

Spikelets 5-7 mm long. Inflores-
cence a one-sided spike, 50-170
mm long, sickle-shaped to cork-
screw - like at maturity; upper
glume tubercled; lemmas awned from below apex on
nerves, with a spreading awn on the back; female-fertile
lemma 4. 0-4. 5 mm long; awns 4. 5-5. 5 mm long.

Flowering December to April. Open veld on sandy or
sometimes moist soils. Locally common. Biome: Savanna
and Grassland. To central tropical Africa.

Fig. 53. Ctenium concinnum

Description: Stapf 1898-1900 (638), Chippindall 1955
(192), Clayton et al. 1970-1982 (325). Illustration: Chip-
pindall 1955 (fig. 167). Voucher: Du Toit 2374. PRECIS
code 9902990-00100.

94

Cymbopogon Spreng.

Cymbanthelia Anderss., Gymnanthelia Schweinf.

Perennial (rarely annual); caespitose. Culms 150-3000
mm high; herbaceous; usually unbranched above. The
shoots aromatic. Leaf blades linear (from broadly so, to
filiform); flat, or folded. Ligule an unfringed membrane to
a fringed membrane . Plants bisexual, with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant ; overtly heteromorphic (the pedicellate spikelets
not depressed abaxially, awnless); in both homogamous and
heterogamous combinations (lowermost pair of lowest
raceme, or of each raceme, homogamous and imperfect).

Inflorescence paniculate (decompound, leafy)', spathe-
ate; a complex of ‘partial inflorescences’ and intervening
foliar organs. Spikelet-bearing axes ‘racemes’ (short,
spikelike, each pair with a spatheole); paired (connate at
base, often widely spreading ordeflexed); with very slender
rachides; disarticulating at the joints.

Spikelets in pairs (or with a terminal triplet); not secund;
consistently in ‘long-and-short’ combinations; these

Fig. 54. Cymbopogon excavatus

pedicellate/sessile. Pedicels free of the rachis. The sessile
spikelets hermaphrodite. The pedicellate spikelets male-
only (usually), or sterile; never depressed or canaliculate on
the back; only LI present, hyaline, 2-nerved, its floret
usually male but occasionally sterile or suppressed. Female-
fertile spikelets 3-7 mm long; compressed laterally, or not
noticeably compressed, or compressed dorsiventrally;
falling with the glumes. Glumes two; more or less equal;
awnless; very dissimilar (lower bicarinate, upper
naviculate). Proximal incomplete florets I ; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline to firm-stipitate beneath the awn); incised
(bifid or apically bilobed); awnless, or awned. Awns when
present 1; from the sinus; geniculate; much shorter than the
body of the lemma, to much longer than the body of the
lemma. Palea absent. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5, or 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. About 40 species.
Tropical and subtropical Africa and Asia, Australia. Meso-
phytic to xerophytic; in open habitats (savanna);
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 6 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

The southern African species are greatly in need of
revision, especially in relation to the tropical African
species.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf blades rounded to cordate at base; lower glume
of sessile spikelet with a deep narrow median
groove in the lower half, appearing as a rib inside;
sessile spikelets 3. 5-5.0 mm long; raceme bases
often much swollen: lowest node of old culms

exposed, not clothed by leaf sheaths

C. excavatus

Leaf blades parallel-sided at base; lower glume of
sessile spikelet deeply concave to flat on back;
sessile spikelets (4.5-)5.0-6.0 mm long; raceme
bases not swollen; lowest node of culms clothed by

leaf sheaths 2

2(1). Lower glume of sessile spikelet wingless or with a
narrow wing (0.1 -0.3 mm wide), back usually
deeply concave (rarely only shallowly concave);
leaf blades 2-4 mm across, usually folded and

appearing setaceous C. plurinodis

Lower glume of sessile spikelet with a wing 0.3-0. 7
mm wide, back shallowly concave to flat; leaf

blades of various widths 3

3(2). Ligules 5-12 mm long, usually acutely pointed and
papery; leaf blades to 2 mm across, usually folded

and appearing setaceous C. dieterlenii

Ligules to 10 mm long, usually truncate or rounded
and firm-textured; leaf blades (2 — )3 mm or more

across, usually flat 4

4(3). Culms robust, 1200-2400 mm tall; leaf blades
500-700 mm long, 5-10 mm across; racemes
appearing glabrous because pedicels are hairy only
along the sides (rarely appearing hairy in specimens

from Natal sea dunes) C. validus

Culms slender, 450-1200 mm tall; leaf blades
300-500 mm long, 3-6 mm across; racemes hairy

or nearly glabrous 5

5(4). Racemes appearing very hairy, rachises and pedicels

with long hairs on the sides and backs

C. marginatus

Racemes appearing nearly glabrous, rachises and
pedicels with hairs only along sides, backs glabrous
C. prolixus

95

Cymbopogon dieterlenii Stapf ex Phill.

Perennial; tufted; 450-850
mm tall. Leaf blades 300-500
mm long; setaceous or to 2 mm
wide. Spikelets 5-6 mm long
(sessile and pedicellate). Ligule
5-12 mm long, papery, pointed;
lower glume of sessile spikelets
winged, flattish.

Flowering November to April.

Open veld and rocky hillsides. Infrequent. Biome: Savanna
and Grassland. Southern Africa. In habit similar to C.
plurinodis, which has short ligules, broader blades and the
lower glume of the sessile spikelet is wingless and deeply
grooved.

Description: Chippindall 1955 (508). Voucher:

Dieterlen 390B. PRECIS code 9900720-00100.

Cymbopogon excavatus (Hochst.) Stapf ex Burtt Davy

Fig. 54.

Common turpentine grass,
lemoengras, buchugras.

Perennial; tussocky; to 1500
mm tall. Leaf blades 50-300 mm
long; to 14 mm wide. Spikelets
(sessile) 3. 5-5.0 mm long (pedi-
cellate slightly shorter). Leaf
blades rounded at base; lowest
culm nodes exposed; lower glume of sessile spikelets with
a deep narrow groove.

Flowering mostly November to May. Open veld and
hillsides. Very common. Biome: Savanna and Grassland.
Southern Africa. Domestic use (thatching). Our species is
here retained as separate from the closely-related tropical
C. caesius, pending a generic revision. C. giganteus ,
another tropical species which is similar to C. excavatus but
larger in all parts, may possibly occur in the extreme north.
Diheteropogon amplectens has similar leaf blades but it is
not aromatic and lacks swollen raceme bases.

Description: Chippindall 1955 (506), Clayton et al.
1970-1982 (761). Voucher: De Winter & Codd 467.
PRECIS code 9900720-00200.

Cymbopogon marginatus (Steud.) Stapf ex Burtt Davy

PI. 49.

Motwortelterpentyngras,
muskusgras.

Perennial; densely tufted;

300-800 mm tall. Leaf blades
150-350 mm long; 2-5 mm wide.

Spikelets 5-6.5 mm long (sessile
and pedicellate). Racemes con-
spicuously hairy, lower glume of
sessile spikelets winged, flattish.

Flowering July to June. Rocky hillsides. Common.
Biome: Fynbos and Nama-Karoo. Endemic. This is the the
‘winter rainfall’ species. It can be distinguished from wide-
bladed forms of C. plurinodis by the concave and wingless
lower glume of the sessile spikelet in that species. However,
specimens from several localities in the southwestern Cape
appear to be intermediate.

Description: Chippindall 1955 (506). Voucher: Taylor
3197. PRECIS code 9900720-00300.

Cymbopogon plurinodis (Stapf) Stapf ex Burtt Davy

Bitter turpentine grass.

Perennial; tufted; 300-1000
mm tall. Leaf blades 150-300
mm long; 2-4 mm wide (often
folded and appearing setaceous).
Spikelets (sessile) 5-6 mm long
(pedicellate slightly shorter).
Lower glume of sessile spikelets

deeply concave, usually wingless or with a narrow wing.

Flowering October to May. Grassveld. Very common.
Biome: Fynbos, Savanna, Grassland, Nama-Karoo, and
Succulent Karoo. Southern and eastern Africa. Our species
is retained as separate from the tropical C. pospischilii
which has longer racemes.

Description: Chippindall 1955 (508), Clayton et al.
1970-1982 (765). Illustration: Chippindall 1955 (fig. 407).
Voucher: Smith 4091. PRECIS code 9900720-00400.

Cymbopogon prolixus (Stapf) Phill.

Tamboekiegras.

Perennial; tufted; 900-1200
mm tall. Leaf blades 300-500
mm long; 3-6 mm wide. Spike-
lets (sessile) 5-6 mm long (pedi-
cellate equalling it or slightly
smaller). Racemes nearly
glabrous, lower glume of sessile
spikelets winged, flattish.

Flowering October to April. Rocky hillsides. Common.
Biome: Savanna and Grassland. Endemic. In its habit
this species appears intermediate between C. plurinodis ,
which is distinguished by the concave and wingless lower
glume of the sessile spikelet, and C. validus, which is larger
and very robust.

Description: Chippindall 1955 (507). Voucher: Smith
1315. PRECIS code 9900720-00500.

Cymbopogon validus (Stapf) Stapf ex Burtt Davy

(=C. afronardus Stapf) 2.

Reuse terpentyngras,
giant turpentine grass.

Robust perennial; tufted;

1200-2400 mm tall. Leaf blades
500-700 mm long; 5-10 mm
wide. Spikelets (sessile) 4. 5-6.0
mm long (pedicellate slightly shorter). Racemes nearly
glabrous; lower glume of sessile spikelets winged, flattish.

Flowering July to June (but usually in autumn). Rocky
hillsides and scrub vegetation, often in damp places. Com-
mon. Biome: Savanna and Grassland. Southern Africa.
Probably to be included with the tropical C. nardus , but
kept separate pending a generic revision.

Description: Chippindall 1955 (507), Clayton et al.
1970-1982 (764). Illustration: Chippindall 1955 (fig. 406).
Voucher: Moll 1665. PRECIS code 9900720-00600.

Cynodon Rich.

Capriola Adans., Dactilon Vill., Fibichia Koel.

Perennial; long-rhizomatous and long-stoloniferous
(often sward-forming). Culms 40-600(-1000) mm high;
herbaceous. Leaf blades linear; flat, or folded. Ligule a
fringed membrane (very short), or a fringe of hairs.

Inflorescence of spike-like main branches ; digitate or
subdigitate (sometimes in two or more closely spaced
whorls); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; 1 .7-3 mm long ; compressed
laterally; disarticulating above the glumes (or between
them). FI airy callus absent. Glumes two; more or less equal;
awnless; similar (narrow, lanceolate ). All florets normally
female-fertile, or distal incomplete florets also present;
proximal incomplete florets absent.

Female-fertile florets 1. Lemmas 1-4 nerved;
membranous; entire; awnless. Palea present; relatively
long. Lodicules fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small; ellipsoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;

96

NAD-ME (2 species); XyMS+. PCR sheath outlines even.
PCR sheath extensions absent. PCR cell chloroplasts elon-
gated; with well developed grana; centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 10. Chloridoideae; Chlorideae sensu lato.
10 species. Tropical and subtropical. Mesophytic, or xero-
phytic; in open habitats; maritime-arenicolous, halophytic,
and glycophytic. Namibia, Botswana, Transvaal, Orange
Free State, Swaziland, Natal, Lesotho, and Cape Province.
Indigenous species (6), naturalized species (2).

Intergeneric hybrids with Chloris (X Cynochloris
Clifford & Everist: several species involved).

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1974. FTEA.

Species treatment by M. Koekemoer.

1(0). Keel of lemma winged; rachilla not produced; glumes
usually shorter than half the spikelet length ... 2
Keel of lemma not winged; rachilla usually produced;
glumes usually longer than half the spikelet length

3

2(1). Leaf blades densely hairy; spikelets 2.0-2. 5 mm long,

broadly ovate C. hirsutus

Leaf blades glabrous or very sparsely hairy; spikelets

2.5-3. 0 mm long, narrowly ovate

C. incompletus

3( 1 ). Spikes 2 or 3 (very rarely 1 or 4); plants stoloniferous,
mat-forming and seldom taller than 150 mm ... 4
Spikes 4-20; plants rhizomatous and/or stoloniferous,
sometimes mat-forming, usually 100-1000 mm tall

6

4(3). Leaves sparsely or densely hairy; spikes most often

3, erect or spreading C. bradleyi

Leaves glabrous; spikes most often 2, ascending when

young and often reflexed at maturity 5

5(4). Rachilla not produced; culms firm; leaves rigid, more
than 1.5 mm wide; spikes not reflexed at maturity

C. polevansii

Rachilla produced and often longer than the lower
glume; culms delicate; leaves very fine, less than

1.5 mm wide; spikes reflexed at maturity

C. transvaalensis

6(3). Plants rhizomatous and stoloniferous, up to 400 mm
tall; spikes usually 4 or 5 (occasionally 3 or up to

6), in a single whorl C. dactylon

Plants stoloniferous, 300-1000 mm tall; spikes 5-20

in 1-5 whorls 7

7(6). Keel of lemma glabrous or with a few solitary hairs;
spikes usually stiff and tardily spreading; plants
robust, often woody, coarse, 400-1000 mm tall

C. aethiopicus

Keel of lemma very densely pubescent; spikes usually
slender to flexuous and spreading; plants fairly
slender to robust, not woody, usually soft, 300-600
mm tall C. nlemfuensis

Cynodon aethiopicus Clayton & Harlan

(experimental plantings on roadsides). Similar to C.
nlemfuensis , which is smaller and less robust and has the
lemma keel very densely pubescent.

Description: Clayton et al. 1970-1982 (319).

Illustration: Clayton et al. 1970-1982 (fig. 89). Voucher:
Smook 4140. PRECIS code 9902960-00100.

Star grass, reuse kweekgras.

Robust perennial; stolonifer-
ous (often woody and coarse);

350-900 mm tall. Leaf blades
30-250 mm long; 3-7 mm wide.

Spikelets 2. 5-3.0 mm long.

Racemes stiff, purple-pigmented,
in multiple whorls; keel of lemma
not winged, glabrous or with a few single hairs.

Flowering January to June. Rich soils, particularly old
cattle kraals and abandoned cultivation, also at moist
streamsides. Infrequent. Naturalized from tropical Africa.
Biome: Savanna. Tropical Africa. Weed, or erosion control

Cynodon bradleyi Stent

Bradley grass.

Perennial; stoloniferous; 50-
100(-300) mm tall. Leaf blades
10-35 mm long; to 2.5 mm wide.

Spikelets 2-3 mm long. Leaves
densely or sparsely hairy; spikes
usually three; rachilla sometimes
produced; lemma keel not
winged.

Flowering December to March. Fertile, well-drained
soils. Infrequent. Biome: Grassland. Endemic. Ornamental

97

(useful lawn grass). Very similar to C. hirsutus , which has
a wing on the lemma keel.

Description: Chippindall 1955 (202). Voucher: De
Winter 382. PRECIS code 9902960-00200.

Cynodon dactylon (L.) Pers.

Fig. 3. Fig. 8. Fig. 55. PI. 50.

Couch grass, kweekgras.

Sward-forming perennial;
rhizomatous and stoloniferous;

50-350 mm tall. Leaf blades
10-120 mm long; 2^1 mm wide.

Spikelets 2. 0-2. 5 mm long.

Racemes (3— )4— 5(— 6), in a single
whorl; lemma keel wingless;
upper glume 1/2-3/4 the spikelet length; rachilla produced.

Flowering September to May. In most soils along
roadsides and overgrazed, trampled areas. Locally domin-
ant. Biome: Fynbos, Savanna, Grassland, Nama-Karoo, and
Desert. Worldwide in warm and temperate regions. Food
and drink (leaves rich in Vitamin C), or pasture (certain
strains), or erosion control (hardy pioneer), or ornamental
(planted as lawn in gardens or sports fields), or traditional
medicine (for heartburn, wounds, indigestion or as a blood
purifier), or chemicals (cynodin and triticin), or weed (in
cultivated lands, and a host for many fungi and viruses).
This species has been reported as a weed in more than 80
countries and because of the rhizome that can be up to 1000
mm deep it is difficult to eradicate. It is, however, also one
of our most valuable grasses because it protects the soil and
provides some grazing in areas that suffer from
overstocking.

Description: Holm et al. 1977 The Worlds Worst Weeds
(25), Stapf 1898-1900 (634), Hitchcock & Chase 1950
(483), Chippindall 1955 (200), Clayton et al. 1970-1982
(318). Illustration: Chippindall 1955 (fig. 175), Hitchcock
& Chase 1950 (fig. 1031). Voucher: De Winter & Hardy
8110, Brueckner 38, Smook 4751. PRECIS code 9902960-
00300.

Cynodon hirsutus Stent

Red quick grass, Transvaal-
kweek.

Perennial; stoloniferous; 50-
250 mm tall. Leaf blades 15-30
mm long; 2^4 mm wide. Spike-
lets 2. 0-2. 5 mm long. Leaf blades
hairy; spikelets broadly ovate;
glumes more than 1/2 the
spikelet length; lemma keel winged; rachilla not produced.

Flowering October to April. Well-drained loam soils.
Common. Biome: Savanna and Grassland. Endemic. Ero-
sion control, or traditional medicine (for indigestion and as
blood purifier). Very similar to C. bradleyi, which lacks a
wing on the lemma keel, andC. incompletus , which has less
hairy leaves and shorter glumes.

Description: Chippindall 1955 (202). Illustration: Chip-
pindall 1955 (fig. 177). Voucher: Potts 3720, Smook &
Gibbs Russell 2467. PRECIS code 9902960-00400.

Cynodon incompletus Nees

Karroo quick grass, soet-
kweek.

Perennial; stoloniferous; 50-
300 mm tall. Leaf blades 30-60
mm long; 2-3 mm wide. Spike-
lets 2. 5-3. 0 mm long. Leaf blades
glabrous or sparsely hairy; spike-
lets narrowly ovate; glumes

less than 1/2 the spikelet length; lemma keel winged;
rachilla not produced.

Flowering November to May. Sandy loam to turf soils.
Common. Biome: Fynbos, Savanna, Grassland, and Nama-
Karoo. Australia and Argentina (probably a cultivated
pasture). Close to C. hirsutus, which has more hairy leaves
and longer glumes.

Description: Stapf 1898-1900 (635), Chippindall 1955
(203). Illustration: Chippindall 1955 (fig. 178). Voucher:
Acocks 8479. PRECIS code 9902960-00500.

Cynodon nlemfuensis Vanderyst

* Star grass, reuse kweekgras.

Perennial; stoloniferous (sto-
lons stout and woody, plants
otherwise fairly soft); 200-600
mm tall. Leaf blades 50-160 mm
long; 2-6 mm wide. Spikelets 2-3
mm long. Spikes usually slender,
flexuous and spreading; lemma
very densely, silky pubescent, keel not winged.

Flowering January to March. In disturbed areas such as
old lands, cattle paddocks and road verges, also moist
streamsides and weedy places. Infrequent. Naturalized from
Kenya. Biome: Savanna. Tropical Africa. Very similar to
C. dactylon which has rhizomes, and C. aethiopicus, which
is larger and more robust and has the lemma keel glabrous
or with a few single hairs. Some specimens have previously
been wrongly identified as C. plectostachyus (K. Schum.)

Pilg-

Description: Clayton et al. 1970-1982 (319). Voucher:
Scheepers 148. PRECIS code 9902960-00550.

Cynodon polevansii Stent

Compact perennial; rhizomat-
ous; 50-120 mm tall. Leaf blades
10-20 mm long; about 2 mm
wide. Spikelets 2. 7-3. 5 mm long.

Leaves rigid; spikes 2, not
reflexed; rachilla not produced.

Flowering December. Moist
areas. Rare. Biome: Grassland.

Endemic. The status of this
species is very uncertain. Apart from the type specimen
there are no other records at PRE. De Wet (1971) J1 S. Afr.
Bot. 37,1 (53) regards this as a variety of C. dactylon.

Description: Chippindall 1955 (202). Voucher: Pole
Evans 334 (type). PRECIS code 9902960-00650.

Cynodon transvaalensis Burtt Davy

Transvaal quick grass.

Perennial; rhizomatous; 50-
300 mm tall. Leaf blades 1.0- 1.5
mm wide. Spikelets 2.0- 2.5 mm
long. Leaf blades involute and
filiform; spikes usually 2, reflex-
ed at maturity; rachilla produced
and often longer than the lower
glume.

Flowering November to May. Roadsides and weedy
places. Infrequent. Biome: Fynbos and Grassland. Northern
Africa, cultivated in Zimbabwe. Weed and ornamental
(lawns). Delicate fine leaves, a long rachilla and spikes that
are reflexed at maturity distinguish it from other Cynodon
species.

Description: Chippindall 1955 (202), Clayton et al.
1970-1982 (317). Voucher: Smook 4747, Muller 1340.
PRECIS code 9902960-00700.

98

Cynosurus L.

Falonia Adans.

Annual, or perennial; caespitose. Culms 100-900 mm
high; herbaceous; unbranched above. Sheath margins free.
Leaf blades linear; flat .Ligule an unfringed membrane. The
spikelets of sexually distinct forms on the same plant;
overtly heteromorphic (fertile spikelets mixed with and
more or less concealed by sterile ones consisting of rigid,
lanceolate, awned glumes and lemmas).

Inflorescence paniculate; contracted; espatheate.
Spikelet-bearing axes persistent.

Female-fertile spikelets 2.8-10 mm long; compressed
laterally; disarticulating above the glumes. Glumes two;
more or less equal; decidedly shorter than the adjacent
lemmas, or long relative to the adjacent lemmas; awnless;
similar (narrow, thin). All florets female-fertile, or distal in-
complete florets also present; proximal incomplete florets
absent.

Female-fertile florets (I-)2-5. Lemmas similar in
texture to the glumes to decidedly firmer than the glumes;
5 nerved; entire, or incised; awned. Awns I ; median; from
the sinus, or apical; non-geniculate; much shorter than the
body of the lemma, to much longer than the body of the

lemma. Palea present; relatively long. Lodicules 2;
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short, or long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. 8 species. Europe,
western Asia, North and South Africa. Mesophytic, or xero-
phytic; in open habitats (meadows, disturbed ground).
Orange Free State and Cape Province. 1 naturalized species
(possibly 1 indigenous species).

References. 1. Chippindall. 1955. Gr. & Past. 2. Linder.
1986. Bothalia 16: 61. 3. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Glumes and lemmas of sterile spikelets produced into
awns that are purple at the base and pale above;
awns 15-20 mm long; anthers 0.4—0. 6 mm long;
plants to 200 mm tall; lemmas 3.5^L0 mm long;

fertile spikelets 1 -flowered C. coloratus

Glumes and lemmas of sterile spikelets produced into
pale awns; awns to 15 mm long; anthers 3-4 mm
long; plants to 600 mm tall; lemmas 5-7 mm long;
fertile spikelets 2-3-flowered C. echinatus

Fig. 56. Cynosurus echinatus

Cynosurus coloratus Lehm. ex Nees

PI. 51.

Annual; loosely tufted; 50-
200 mm tall. Leaf blades 10-50
mm long; 2-3 mm wide. Spike-
lets 10-25 mm long. Female-
fertile spikelets 1 -flowered;
glumes and lemmas of sterile
spikelets produced into awns
which are purple at the base and
pale above, 15-20 mm long;
glumes 4-7 mm long; lemmas 3. 5-4.0 mm long; anthers
0.4—0. 6 mm long.

Flowering March to April. In rocky areas, usually
calcareous soils. Rare. Biome: Fynbos. Mediterranean
region. Although the type specimen was collected in South
Africa, this species has a very doubtful and unsure status.
A deeper investigation of more material is needed before
it can be decided that both the Cynosurus species are
represented in South Africa.

Description: Bor 1985 (1725), Linder (31). Voucher:
Leistner 275. PRECIS code 9903730-00050.

Cynosurus echinatus L.

Fig. 56.

Dogstail.

Annual; tufted (culms erect or
decumbent at the base); 200-600
mm tall. Leaf blades 50-150 mm
long; 3-10 mm wide. Spikelets
7-20 mm long. Female-fertile
spikelets 2-3-flowered; glumes
and lemmas of sterile spikelets
produced into pale awns up to 15 mm long; glumes 7-12
mm long; lemmas 5-7 mm long; anthers 3^1 mm long.

Flowering July to January. On rocky, well-drained soils
and disturbed places such as roadsides, often in the shade.
Infrequent. Naturalized from Europe. Biome: Fynbos and
Nama-Karoo. Atlantic islands and Mediterranean region
eastwards to India. Weed.

Description: Bor 1985 (1726), Chippindall & Crook
1976 (204), Linder (30), Stapf 1898-1900 (690), Chippin-
dall 1955 (61). Illustration: Chippindall 1955 (fig. 33).
Voucher: Crook 2318. PRECIS code 9903730-00100.

99

Dactylis L.

Amaxitis Adans., Trachypoa Bub.

Perennial; caespitose (with short, oblique rhizomes and/
or stolons). Culms 150-2000 mm high; herbaceous; un-
branched above. Sheath margins joined (to halfway, at least
in the upper leaves). Leaf blades linear to linear-lanceolate;
flat, or rolled (involute). Ligule an unfringed membrane .

Inflorescence paniculate', open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets secund (in dense, one-sided clusters
terminating the panicle branches); 4-8 mm long; com-
pressed laterally; disarticulating above the glumes. Glumes
two; relatively large; very unequal, or more or less equal;
markedly shorter than the spikelets to about equalling the
spikelets; awned to awnless; carinate ; similar

(membranous, somewhat curved). Incomplete florets distal
to the female-fertile florets; proximal incomplete florets
absent.

Female-fertile florets 2-5. Lemmas decidedly firmer
than the glumes', 5 nerved; entire; awned. Awns 1; median;
dorsal, or apical; non-geniculate; much shorter than the
body of the lemma. Palea present; relatively long. Lodicules
2; membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 7. Pooideae; Poodae; Poeae. 1 species (or up
to 5, by recognition of minor segregates). Temperate
Eurasia. Mesophytic; in shade and in open habitats
(meadows, woodlands and disturbed ground, in moist to dry
places). Transvaal, Orange Free State, Natal, and Cape
Province. 1 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Linder.
Unpubl. ms, FSA.

Dactylis glomerata L.

Fig. 57. PI. 52.

Cocksfoot, orchard grass.

Perennial; densely and coarse-
ly tufted, or rhizomatous (rhi-
zome oblique); 150-800(-1400)
mm tall. Leaf blades 100-450
mm long; 2-14 mm wide. Spike-
lets 5-9 mm long. Leaf blades
folded at first, pilose on upper
surface, glabrous, shining and smooth below; ligule
membranous, 2-10 mm long; panicle 50-300 mm long,
racemes closely spaced, lowest usually solitary, remote and
bare at the base; spikelets 3-5(-7)-flowered, laterally
compressed; glumes and lemmas strongly keeled, keels
finely or coarsely ciliate, tips acuminate, mucronate or
awned, awn to 1.5 mm long.

Flowering July, August and November to February.
Mostly in cultivation but also in other disturbed places like
roadsides. Infrequent. Naturalized from Europe. Biome:
Fynbos and Grassland. Introduced to most temperate
countries. Used to a limited extent as winter pasture.

Description: Linder(55), Stapf 1898-1900(696), Flitch-
cock & Chase 1950 (184), Chippindall 1955 (49), Clayton
et al. 1970-1982 (43). Illustration: Chippindall 1955 (fig.
18), Clayton et al. 1970-1982 (fig. 16), Hitchcock & Chase
1950 (fig. 366). Voucher: Du Toit 2543. PRECIS code
9903980-00100.

Dactyloctenium Willd.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent (mostly low,
sometimes sward-forming). Culms 50— 1 000( — 1 600 ) mm
high; herbaceous. Leaf blades linear to linear-lanceolate;
flat, or rolled. Ligule a fringed membrane (narrow), or a
fringe of hairs. The spikelets all alike in sexuality (or
terminal spikelets sterile).

Inflorescence of spike -like main branches ( spikelets
almost at right-angles to the rachides)', digitate or subdigi-
tate; axes not ending in spikelets (produced into a flattened
point)', espatheate. Spikelet-bearing axes disarticulating',
falling entire.

Female-fertile spikelets biseriate; 2.3-8 mm long; com-
pressed laterally; disarticulating above the glumes; not dis-
articulating between the florets (rachilla tough). Glumes
two (persistent, membranous, laterally compressed); more
or less equal; markedly shorter than the spikelets; awned,
or awnless (the lower muticate, the upper awned or
mucronate). Incomplete florets distal to the female-fertile
florets; proximal incomplete florets absent.

Female-fertile florets 3-6. Lemmas 1-3 nerved; entire,
or incised; awnless, or mucronate, or awned. Awns when
present 1; apical; much shorter than the body of the lemma.
Palea present; relatively long. Lodicules 2; fleshy. Stamens
3. Ovary glabrous. Fruit small (0.7-1. 1 mm long); ellipsoid
to subglobose; hilum short; pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions present. Maximum number of extension cells 1.
PCR cell chloroplasts with well developed grana;
centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10 and 12. Chloridoideae; Chlorideae sensu
lato. 13 species. In warm regions. Mesophytic to xero-
phytic; in open habitats; maritime-arenicolous, halophytic,
and glycophytic (sometimes in saline habitats or dunes,
mostly in dry sandy soils). Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, and Cape Province.
4 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Species treatment by M. Koekemoer.

100

1(0). Plants annual, tufted; caryopsis triangular with a

truncate to concave apex; spikes 3-9 2

Plants perennial, stoloniferous; caryopsis with a

Fig. 58. Dactyloctenium giganteum

rounded to convex apex; spikes 1-3 (rarely to 6)

3

2(1). Anthers 0.3-0. 8 mm long; plants not robust, culms
geniculate and ascending; glume awn shorter than

twice the glume length D. aegyptium

Anthers longer than 0.8 mm; plants robust, culms
erect; glume awn longer than twice the glume

length D. giganteum

3(1). Lemma keel smooth, tip acute .... D. geminatum
Lemma keel scabrid, tip awned (awn 0.5-0. 7 mm
long) D. australe

Dactyloctenium aegyptium (L.) Willd.

Crowfoot, duck grass.

Mat-forming annual; tufted
(culms geniculately ascending
and rooting at the nodes); 70-750
mm tall. Leaf blades 30-250 mm
long; 3-8 mm wide. Spikelets
3.5 — 4.5 mm long. Spikes 4-8,

15-65 mm long; lemma keel
scabrid above the middle, ending in a mucro to 1 mm long;
anthers 0. 3-0.8 mm long; grains broadly triangular, apex
truncate to concave.

Flowering January to April. Disturbed areas near water.
Common. Biome: Savanna, Grassland, Nama-Karoo, and
Desert. Tropical and warm temperate regions worldwide.
Food and drink (seed used as food in times of famine), or
pasture, or poisonous (bruised young seed used as a fish
poison), or traditional medicine (extracted to treat kidney
ailment and coughing), or weed (in ricefields and waste
ground, host for viruses).

Description: Chippindall & Crook 1976 (5), Fisher &
Schweickerdt 1941 Ann. Nat. Mus. 10(1), Stapf 1898-1900
(646), Hitchcock & Chase 1950 (481), Chippindall 1955
(132), Clayton et al. 1970-1982 (252). Illustration: Chip-
pindall 1955 (fig. 104), Hitchcock & Chase 1950 (fig.
1029). Voucher: G.J. du Toil 229. PRECIS code 9903320-
GO 100.

Dactyloctenium australe Steud.

Durban grass. Natal crowfoot.

Perennial; stoloniferous;

130-810 mm tall. Leaf blades
50-270 mm long; 2.0 — 4.5 mm
wide. Spikelets 3^4- mm long.

Spikes 2-3, 30-50 mm long;
lemma keel scabrid above middle,
awn 0.5-0. 7 mm long; anthers
1.3-1. 7 mm long; grains obovate, apex rounded to convex.

Flowering January to May. Sandy soils on seashores,
dunes and along forest roads, often in light shade. Common.
Biome: Savanna. Tropical east Africa. Pasture (grazed by
stock), or erosion control (good sandbinder), or ornamental
(lawns near coast, also grows in shade).

Description: Chippindall 1955 (131), Clayton et al.
1970-1982 (256). Voucher: Smook 5532. PRECIS code
9903320-00200.

Dactyloctenium geminatum Hack.

Mat-forming perennial; rhizo-
matous and stoloniferous;

350-870 mm tall. Leaf blades
40-250 mm long; 3-6 mm wide.

Spikelets 3. 0-5. 3 mm long.

Spikes usually 2(— 3), 25-70 mm
long; lemma tips acute; anthers
1.1-1. 7 mm long; grains obovate,
apex rounded to convex.

101

Flowering December to March. On sandy soil of alkaline
pans and on sand dunes, in swamp forest undergrowth,
coastal sandflats and open grassveld. Locally common.
Biome: Savanna. Tropical east Africa to Somalia. Erosion
control (sand binder).

Description: Chippindall 1955 (131), Clayton et al.
1970-1982 (255). Illustration: Clayton et al. 1970-1982
(fig. 70). Voucher: De Winter & Codd 545. PRECIS code
9903320-00300.

Dactyloctenium giganteum Fisher & Schweick.

Sterretjiegras, gaint crowfoot. Fig- 58. PI. 53.

Robust annual; tufted (erect);

480-1140 mm tall. Leaf blades
110-450 mm long; 5-12 mm
wide. Spikelets 4.0-6. 2 mm long.

Spikes 3-9, 35-110 mm long;
lemma keels scabrid; awns 0.7-
2.0 mm long; grains triangular,
apex truncate to concave.

Flowering November to May. Open veld or disturbed
areas on river banks or near water, often in shade. Common.
Biome: Savanna and Grassland. Tropical east Africa. Weed.
The voucher specimen is one of seven duplicates at PRE,
which are progeny of the type specimen and were cultivated
at the Natal Herbarium.

Description: Chippindall & Crook 1976 (5), Chippindall
1955 (132), Clayton et al. 1970-1982 (251). Voucher:
Schweickerdt 1451. PRECIS code 9903320-00400.

Danthoniopsis Stapf

Gazachloa Phipps, Jacquesfelixia Phipps, Petrina
Phipps, Pleioneura (C. E. Hubb.) Phipps, Rattraya Phipps,
Xerodanthia Phipps.

Annual (rarely), or perennial; caespitose (sometimes
densely so). Culms 250-2000 mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear
to lanceolate; flat, or rolled (but only slightly so). Ligule a
fringe of hairs. Plants with hermaphrodite florets.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets in triplets (rarely), or in pairs; not secund ; 5-20
mm long; compressed laterally, disarticulating above the
glumes. Glumes two; very unequal; awnless, or awned (G2
sometimes aristate); very dissimilar (G1 acute to
acuminate, G2 with the tip extended). Proximal incomplete
florets 7; paleate, palea fully developed (membranous
between the two narrowly winged keels); male.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes (to slightly firmer); hairy (the hairs in tufts, or not
in tufts; in transverse rows, or not in transverse rows (in
Sect. Pleioneura)); having the margins tucked in onto the
palea; with a clear germination flap (just above the callus,
often hidden by hairs); 7-9 nerved; incised; deeply cleft',
awned. Awns 1, or 3; median, or median and lateral. The
median awn different in form from the laterals (when
laterals present); from the sinus (flat, basally twisted); gen-
iculate; much longer than the body of the lemma. Palea
present; relatively long (about equalling the lemma). Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Hilum
long-linear; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional, or unconventional. Organization
of PCR tissue when unconventional Arundinella type.
XyMS-. PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x - 9, or 12 (?). Panicoideae; Panicodae; Arun-
dinelleae. About 20 species. Africa, Arabia. Mesophytic to
xerophytic; in open habitats (savanna woodland and desert

fringes); glycophytic. Namibia, Botswana, Transvaal,
Natal, and Cape Province. 6 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1967. Kew Bull. 21: 123. 3. Launert. 1970. FSWA. 4.
Phipps. 1972. Bolm. Soc. Broteriana 46: 423. 5. PRE
Herbarium practice, following Smook & Gibbs Russell.

Species treatment by H.M. Anderson.

102

1(0). Plants annual; spikelets 14-20 mm long, with striking
colouration, light green variegated with dark purple

D. dinteri

Plants perennial; spikelets less than 14 mm long,

without striking colouration, mainly straw-coloured

and sometimes tinged with purple 2

2(1). Spikelets in lax triads . . * D. scopulorum

Spikelets in pairs or borne singly 3

3(2). Culms profusely branched and many-jointed

D. ramosa

Culms sparingly branched or simple 4

4(3). Culms thick-walled and woody; plants growing near

water D. lignosa

Culms brittle or delicate; plants growing amongst

rocks 5

5(4). Spikelets straw-coloured, 4 mm long; culms delicate

D. parva

Spikelets light sienna brown, 5-9 mm long; culms
brittle D. pruinosa

Danthoniopsis dinteri (Pilg.) C.E. Hubb.

Annual; tufted; to 2000 mm
tall. Leaf blades 300-600 mm
long; 8-15 mm wide. Spikelets
14-20 mm long. Culms robust,
unbranched; inflorescence light
green variegated with dark pur-
ple; spikelets in lax triads, pedi-
cels unequal, usually the 2 lower
spikelets are reduced; lower lem-
ma 7-nerved; female-fertile (upper) lemma loosely hairy,
lobes acute and 4 mm long, central awn 10-22 mm long.

Flowering February to June. Among rocks and in rock
crevices on mountains. Infrequent. Biome: Savanna and
Grassland. Zimbabwe, Angola. Natural pasture (grazed
when young).

Description: Muller 1984 (118), Chippindall 1955 (286).
Illustration: Chippindall 1955 (255). Voucher: Smook
5137. PRECIS code 9901770-00200.

Danthoniopsis lignosa C.E. Hubb.

Robust perennial; tufted; to
2000 mm tall. Leaf blades to 400
mm long; 8 mm wide. Spikelets
4-6 mm long. Culms very thick-
walled and woody; inflorescence
dense; spikelets in pairs, straw
coloured or tinged with light pur-
ple; female-fertile (upper) lemma
1 1 -nerved, awns 5 mm long.

Flowering July. River edges in flowing water. Infre-
quent. Biome: Savanna. Angola.

Description: Hubbard 1949 Kew Bull. 3: (351).
Voucher: Leistner, Oliver, Steenkamp & Vorster 313.
PRECIS code 9901770-00250.

Danthoniopsis parva (J.B. Phipps) Clayton

Perennial; tufted; 300-600
mm tall. Leaf blades to 50 mm
long; 3 mm wide. Spikelets about
4 mm long. Culms delicate; in-
florescence straw coloured;
spikelets in pairs; upper glume 5-
nerved; lower lemma 3-nerved,
awns 7-10 mm long.

Flowering January to May.

Rock crevices on cliffs. Infrequent. Biome: Savanna. This
species is close to D. pruinosa , which has larger spikelets
and a 3-nerved upper glume.

Description: Phipps 1964 Kirkia 4 (118). Voucher:
Smook 5401. PRECIS code 9901770-00300.

Danthoniopsis pruinosa C.E. Hubb.

Rock powder grass.

Perennial; tufted; 600-1800
mm tall. Leaf blades 100-250
mm long; 5-9 mm wide. Spike-
lets 5-9 mm long. Rhizome swol-
len, woody; culms brittle, often
branched, with a waxy bloom
below the node, nodes occasion-
ally hairy; spikelets light or sienna brown; upper glume and
lower lemma 3-nerved; lower lemma with three tufts of
white hairs on either side above the middle, lobes acute and

I. 0-1. 5 mm long, awns 7-12 mm long.

Flowering December to June. Among rock and rock
crevices on mountains; granite outcrops and other rock
types. Locally common. Biome; Savanna. Zambia,
Zimbabwe and southern Malawi.

Description: Chippindall & Crook 1976 (100), Stapf
1898-1900 (286). Illustration: Chippindall 1955 (fig. 256).
Voucher: Raal 467. PRECIS code 9901770-00500.

Danthoniopsis ramosa (Stapf) Clayton

( =Trichopteryx ramosa
Stapf) 5; (=Loudetia anomala
C.E. Hubb. & Schweick.) 3;

(=Loudetia ramosa (Stapf) C.E.

Hubb.) 2.

Shrub or dwarf shrub; tufted;

450- 600 mm tall. Leaf blades to
300 mm long; 3 mm wide. Spike-
lets about 10 mm long. Culms profusely branched with
many joints; spikelets borne singly or in pairs, pale green
to straw coloured, usually tinged with purple; lower lemma
5-7-nerved and glabrous; female-fertile (upper) lemma
loosely hairy, lobes acute, 2 mm long, awns 8-12 mm long.

Flowering December to June. Among rocks on hills and
in ravines. Locally common. Biome: Nama-Karoo. Pasture.

Description: Chippindall 1955 (284). Illustration:
Muller 1984 (fig. 59). Voucher: De Winter 2776. PRECIS
code 9901770-00700.

Danthoniopsis scopulorum (J.B. Phipps) J.B. Phipps

(=Gazochloa scopulorum

J. B. Phipps) 4.

Perennial; tufted; 300-400
mm tall. Leaf blades to 200 mm
long; filiform or to 2 mm wide.

Spikelets about 9 mm long.

Spikelets arranged in lax triads,
straw coloured; lower lemma 7-
nerved, awns 12 mm long.

Flowering June. Growing on rock faces. Infrequent.
Biome: Savanna. Similar to the tropical species D.
chimanimaniensis , which has purple spikelets and lower
lemma 5-nerved.

Description: Phipps 1965 Kirkia 5 (229). Voucher: Codd
4314 (the type specimen). PRECIS code 9901770-00750.

103

Deschampsia P. Beauv.

Airidium Steud., Aristavena Albers & Butzin, Avenella
Pari., Campella Link, Czerniaevia Ledeb., Erioblastus
Honda, Homoiachne Pilger, Lerchenfeldia Schur,
Podinapus Dulac.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent (but usually
caespitose). Culms 80-2000 mm high; herbaceous; un-
branched above. Leaf blades linear; flat, or folded, or rolled
(convolute). Ligule an unfringed membrane.

Inflorescence paniculate ; espatheate. Spikelet-bearing
axes persistent.

Spikelets not secund\ 3-9 mm long\ compressed
laterally; disarticulating above the glumes. Rachilla hairy.
Glumes two; very unequal (rarely), or more or less equal;
markedly shorter than the spikelets to about equalling the
spikelets; long relative to the adjacent lemmas ; awnless;
similar (subscarious to membranous, with thin margins).
All florets female-fertile, or distal incomplete florets also
present; proximal incomplete florets absent.

Female-fertile florets 2-3 (usually 2, rarely only one).
Lemmas similar in texture to the glumes to decidedly firmer
than the glumes; non-carinate\ 4-7 nerved; entire, or
incised (2-lobed, toothed or truncate); awned. Awns 1;

Fig. 60. Deschampsia cespitosa

median; dorsal; non-geniculate, or geniculate; much shorter
than the body of the lemma, to much longer than the body
of the lemma. Palea present; relatively long. Lodicules 2;
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; embryo large (rarely), or small.

Cytology, classification, distribution. Chromosome base
number, x = 7 and 13. Pooideae; Poodae; Aveneae. 40
species. North and South temperate, high altitude tropics.
Helophytic, or mesophytic; in shade and in open habitats
(meadows, upland grasslands and woods). Orange Free
State, Lesotho, and Cape Province. 2 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA. 3. Clarke. 1980. FI. Europ.

Species treatment by T.M. Sokutu.

1(0). Ligule deeply lobed, 1-2 mm long; spikelets 5-6 mm
long; awn geniculate, well exserted beyond the

glumes; lemma apices notched D. flexuosa

Ligule entire, 5.0-11.5 mm long; spikelets 3-4 mm
long; awn almost straight, hardly exserted beyond
the glumes; lemma apices jagged . . D. cespitosa

Deschampsia cespitosa (L.) Beauv.

Fig. 60. PI. 55.

Perennial; tufted (to densely
so); 250-850 mm tall. Leaf blades
70-200 mm long; 2-4 mm wide.

Spikelets 3-A mm long. Leaf
blades expanded, scabrid, pale
green to light brown when dry;
ligule entire, 5.0-11.5 mm long;
lemma apices deeply notched,
truncate, awn almost straight,
hardly exserted from the glumes.

Flowering January to March. Typical grassland habitat,
usually damp or black nutrient-rich soil. Infrequent. Nat-
uralized from Europe. Biome: Grassland. Northwards
through Africa and the Mediterranean to Europe. Easily
distinguished from D . flexuosa by its jagged lemma apices
and leaves that are never filiform.

Description: Hubbard 1954 Grasses (227), Clarke 1980
(5:225), Chippindall 1955 (85), Clayton et al. 1970-1982
(92). Voucher: Hoener 1769, Du Toit 2233. PRECIS code
9901890-00100.

Deschampsia flexuosa (L.) Trin.

Perennial; tufted; 50-500 mm
tall. Leaf blades 200-1200 mm
long; 0.5-1. 5 mm wide. Spikelets
5-6 mm long. Leaf blades
convolute, glabrous, dark green;
ligule deeply lobed, 1-2 mm
long; lemma apices shallowly
notched, awns geniculate. Well
exserted beyond the glumes.

Flowering January to March. Uplands, dry to wet sandy
loam soil, 2200-2300 m. Infrequent. Naturalized from
Europe. Throughout Africa, America, Europe. It is not
always clear whether the awn originates basally or from the
lower 1/3 of the lemma. Only a few specimens of this
species were available. Based on the descriptions and
comments on herbarium sheets, our material represents var.
afromontana C.E. Hubb.

Description: Hubbard 1954 Grasses (225), Clarke 1980
(5:226), Chippindall 1955 (85), Clayton et al. 1970-1982
(94). Illustration: Chippindall 1955 (fig. 56). Voucher:
Esterhuysen 28262. PRECIS code 9901890-00200.

104

Diandrochloa DeWinter

Sometimes included in Eragrostis Wolf.

Annual, or perennial; caespitose. Culms 100-1500 mm
high; herbaceous (soft, geniculate or erect); branched
above, or unbranched above. Leaf blades linear to linear-
lanceolate; flat. Ligule an unfringed membrane.

Inflorescence paniculate ; open, or contracted; non-
digitate (the branches in pseudo-whorls on a central axis);
espatheate. Spikelet-bearing axes persistent.

Fig. 61. Diandrochloa namaquensis

Spikelets solitary; 1-3.5 mm long\ compressed laterally;
disarticulating above the glumes; disarticulating between
the florets. Hairy callus absent (‘ callus swollen, truncate,
glabrous' ). Glumes two; very unequal to more or less equal;
markedly shorter than the spikelets; awnless; similar
(membranous or sub-hyaline, ovate to lanceolate, often
green). Upper glume 1 nerved. All florets female-fertile, or
distal incomplete florets also present, these merely underde-
veloped; proximal incomplete florets absent.

Female-fertile florets 2-14. Lemmas similar in texture
to the glumes to decidedly firmer than the glumes
(translucent or thinly leathery); 3 nerved; entire, or incised;
awnless. Palea present; relatively long. Lodicules 2, fleshy,
glabrous. Stamens 2. Ovary glabrous. Hilum short; pericarp
fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 7
species. Americas, Australia, Asia, Africa. In shade, or in
open habitats; glycophytic. Namibia, Botswana, Transvaal,
Natal, and Cape Province. 2 indigenous species.

References. 1. De Winter. 1960. Bothalia 7: 387.

Species treatment by M. Koekemoer.

1(0). Spikelets 1.0-1. 5 mm long, 2-4-flowered; lemmas
0.5-0. 6 mm long; inflorescence branches spreading

D. pusilla

Spikelets 2-3 mm long, 4— 8-flowered; lemmas about
1 mm long; inflorescence branches somewhat
contracted D. namaquensis

Diandrochloa namaquensis (Nees) De Winter

Fig. 61. PI. 56.

(-Eragrostis namaquensis
Nees ex Schrad.) 1.

Annual; tufted; 250-1500 mm
tall. Leaf blades 60-300 mm
long; 2-6 mm wide. Spikelets 2-3
mm long. Inflorescence branches
somewhat contracted, young
branches often wavy; spikelets
4-8-flowered, very delicate.

Flowering throughout the year (with peak from March
to May). Always near water on sandy or clayey soil. Locally
common (moist areas). Biome: Savanna, Grassland, Nama-
Karoo, and Desert. Tropical Africa. Distinguished from
Eragrostis by a membranous ligule and only two stamens
per floret.

Description: De Winter 1960 (387), Chippindall 1955
(182), Clayton et al. 1970-1982 (209). Illustration: Chip-
pindall 1955 (fig. 156). Voucher: Smith 1480. PRECIS
code 9902852-00100.

Diandrochloa pusilla (Hack.) De Winter

(-Eragrostis pusilla
Hack.) 1.

Annual; tufted; 1 00 — 420 mm
tall. Leaf blades 50-150 mm
long; 2-5 mm wide. Spikelets
1.0-1. 5 mm long. Inflorescence
branches spreading when fully
developed, spikelets 2^1-
flowered, very delicate.

Flowering March to May. Always near water in sandy
or clayey soil, often in the shade. Conservation status not
known. Infrequent. Biome: Savanna. Zimbabwe and
Angola. Distinguished from Eragrostis by a membranous
ligule and only two stamens per floret.

Description: De Winter 1960 (387), Chippindall 1955
(182). Illustration: Chippindall 1955 (fig. 157). Voucher:
De Winter & Codd 313. PRECIS code 9902852-00200.

105

Dichanthium Willem.

Diplasanthum Desv., Lepeocercis Trin.

Annual (rarely), or perennial; long-rhizomatous, or
long-stoloniferous, or caespitose, or decumbent. Culms
150-2000 mm high; herbaceous; branched above, or un-
branched above. Leaf blades flat. Ligule an unfringed
membrane to a fringed membrane . Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant ; overtly heteromorphic (the pedicellate
spikelets smaller, awnless), or homomorphic; in both ho-
mogamous and heterogamous combinations (the lowest
pair being imperfect and homogamous).

Inflorescence of spike-like main branches (many-jointed
‘racemes’ ), or paniculate ( rarely : the lower ‘racemes’
sometimes branched again at the base)-, digitate or subdigi-
tate (the racemes often almost palmate, towards the culm
tips)-, spatheate, orespatheate; not comprising ‘partial inflo-
rescences’ and foliar organs. Spikelet-bearing axes
‘racemes’ (with many — more than 8 — sessile spikelets);
solitary, or paired, or clustered; with very slender rachides\
disarticulating at the joints. The pedicels and internodes of
the rachis without a longitudinal, translucent furrow.
‘Articles’ without a basal callus-knob.

Spikelets in pairs (with a terminal triplet); consistently
in Tong-and-short' combinations; these pedicellate/sessile.
Pedicels free of the rachis. The sessile spikelets hermaphro-
dite (save at the raceme base, where the spikelet pairs are
homogamous). The pedicellate spikelets male-only, or
sterile; awnless. Female-fertile spikelets compressed dorsi-
ventrally; falling with the glumes. Glumes two; relatively
large; more or less equal; awnless; very dissimilar (lower

Fig. 62. Dichanthium annulatum var. papillosum

bicarinate, upper narrower and naviculate). Proximal in-
complete florets 7; epaleate; sterile.

Female-fertile florets 1 . Lemmas less firm than the
glumes (reduced to a hyaline stipe); entire; awned. Awns
1; median; apical; geniculate; much longer than the body
of the lemma. Palea present, or absent; when present very
reduced. Lodicules 2; fleshy; glabrous. Stamens 1-3. Ovary
glabrous. Fruit small; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. About 16 species. Old World
Tropics. Helophytic to xerophytic; in open habitats (from
marshes to subdesert and disturbed ground); glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, Cape Province. Indigenous
species (1), naturalized species (1).

Intergeneric hybrids with Bothriochloa.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Culm glabrous below inflorescence, nodes with a ring
of spreading hairs; lower glume of sessile spikelet

to 1.5 mm across D. annulatum

Culm velvety below inflorescence, nodes glabrous to
short-woolly; lower glume of sessile spikelet to 2.5
mm across D. aristatum

Dichanthium annulatum (Forssk.) Stapf var. papillosum

(A. Rich.) De Wet & Harlan

(-D. nodosum sensu Acocks,
non Willemet) 1; (=D.
papillosum (Hochst.)

Stapf) 1.

Blue grama, vlei finger grass.

Perennial; densely tufted; to
1000 mm tall. Leaf blades 30-300
mm long; to 7 mm wide. Spikelets (sessile) 2. 5-5.0 mm
long; 1.0-1. 5 mm wide. Nodes with a ring of spreading
hairs; culms glabrous below inflorescence; lower glume of
sessile spikelets to 1.5 mm wide.

Flowering July to June (mostly in late summer).
Riverbanks and wet places. Common. Biome; Savanna and
Nama-Karoo. North to Ethiopia.

Description: Chippindall 1955 (480), Clayton et al.
1970-1982 (725). Illustration: Chippindall 1955 (fig. 394).
Voucher: Edwards 3057. PRECIS code 9900640-00050.

Dichanthium aristatum (Poir.) C.E. Hubb.

Perennial; tufted; to 1100 mm
tall. Leaf blades 30-250 mm
long; 2-7 mm wide. Spikelets
(sessile) 3. 0^1.5 mm long; to 2.5
mm wide. Nodes glabrous or
short-woolly; culms velvety
below inflorescence; lower glume
of sessile spikelets to 2.5 mm
wide.

Flowering October to June.

Disturbed and moist places. Infrequent. Naturalized from
tropical Asia. Biome: Savanna. Introduced to most tropical
areas.

Description: Chippindall 1955 (481), Clayton et al.
1970-1982 (723). Voucher: Mogg 13709. PRECIS code
9900640-00100.

Fig. 62. PI. 57.

106

Digitaria Haller

Acicarpa Raddi, Digitariella De Winter,
Elytroblepharum Steud., Elytroblepharum (Steud.)
Schlecht., Eriachne Phil., Gramerium Desv., Sanguinaria
Bub ..Sanguinella Gleichen ,Syntherisma Walt ,,Trichachne
Nees, Valota Adans.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent (sometimes sward
forming). Culms (60-) 150-3000 mm high (or more?); her-
baceous; branched above, or unbranched above. Leaf blades
linear to lanceolate; flat, or folded, or rolled. Ligule an
unfringed membrane (usually).

Inflorescence of spike-like main branches', open, or
contracted; digitate or subdigitate, or non-digitate; espathe-
ate. Spikelet-bearing axes persistent.

Fig. 63. Digitaria eriantha

Spikelets solitary, or in pairs, or in triplets; consistently
in ‘long-and-short’ combinations, or not in distinct ‘long-
and-short’ combinations; abaxial ; compressed dorsiven-
trally; falling with the glumes. Glumes one per spikelet, or
two; very unequal (lower tiny or suppressed); awnless; very
dissimilar. Proximal incomplete florets /; paleate, or
epaleate, palea when present reduced; sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes; smooth to
striate; hairless (no more than minutely striate-papillate);
having the margins lying flat and exposed on the pa lea ;
with a clear germination flap; 1-3 nerved (obscured);
entire; awnless (but often apiculate). Palea present; rela-
tively long (about equalling the lemma). Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit small.
Hilum short; embryo large.

Photosynthetic pathway. C4; NADP-ME (D.
sanguinalis)', XyMS-. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9, 15, and 17. Panicoideae; Panicodae; Pani-
ceae. 220 species. Mainly in warm regions. Mesophytic, or
xerophytic; mostly in open habitats (diverse habitats,
including weedy ground and sandy beaches). Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. Indigenous species (35),
naturalized species (1).

References. 1 . De Winter. 1961 . Bothalia 7: 467. 2. Kok.
1981. Bothalia 13: 435. 3. Clayton & Renvoize. 1982.
FTEA. 4. Kok. 1984. SA. J. Bot. 3: 185. 6. Kok. 1978. DSc,
Univ. Pretoria. 6. Kok et al. 1989. SA. J. Bot. 55: 141. 7.
Veldkamp. 1973. Blumea 21: 1. 8. Webster. 1987. Sida 12:
209.

Species treatment by P.D.F. Kok and H.M. Anderson.

1(0). Pedicel tips with stiff white hairs, some overtopping

the spikelet D. diagonalis var. diagonalis

Pedicel tips with or without stiff hairs, but if present

not overtopping the spikelet 2

2(1). Internode present between the glumes, or between the

glumes and rest of spikelet 3

Intemode absent between the glumes, or between the

glumes and rest of spikelet 5

3(2). Internode present between glumes and rest of spikelet

D. gymnostachys

Intemode present between lower and upper glume .

4

4(3). Upper glume longer than 6.0 mm . D. remotigluma

Upper glume shorter than 4.5 mm D. debilis

5(2). All or the lower racemes arranged in 6 or more whorls

D. perrottetii

Racemes arranged digitately, single or irregularly, or
if in whorls then these 3 or fewer 6

6(5). Racemes without spikelets for the lower third of their
length; central axis longer than the racemes ....

D. flaccida

Racemes bearing spikelets along their whole length;
central axis shorter than the racemes (except in

some specimens of D. velutina) 7

7(6). Inflorescence a single raceme; female-fertile (upper)
lemma not black at maturity . . . D. monodactyla
Inflorescence of two or more racemes (except in some
specimens ofD. eylesii', but then the female-fertile

(upper) lemma black at maturity) 8

8(7). Spikelet hairs silvery or silky and extending more
than 1 mm beyond apex of spikelets into a brush-
like point 9

Spikelet hairs if present not silvery or silky, and
extending less than 0.5 mm beyond apex of

spikelets, not forming a brush-like point 11

9(8). Rhizome horizontal to oblique . D. tricholaenoides
Rhizome, if present, not horizontal to oblique . . 10

10(9). Plants annual, lower leaf sheaths glabrous or
sparsely hairy D. gayana

107

Plants perennial, lower leaf sheaths densely hairy

D. brazzae

11(8). Spikelets in threes, if some are paired then the
female-fertile (upper) lemma dark brown or
black, or paired spikelets alternate with solitary

spikelets 12

Spikelets paired or solitary; female-fertile (upper)

lemma pale brown, greyish or purple 20

12(11). Upper glume as long as the female-fertile (upper)
lemma, concealing the lemma which is pallid to

dark brown 13

Upper glume shorter and narrower than the female-
fertile (upper) lemma, exposing the lemma which

is dark brown or black 15

13(12). Rachis triquetrous; spikelets longer than 2.0 mm

D. angolensis

Rachis winged; spikelets shorter than 1.8 mm . 14
14(13). Upper lemma grey; racemes typically paired ....

D. longiflora

Upper lemma dark brown; racemes typically three

or more D. violascens

15(12). Plants perennial 16

Plants annual 18

16(15). Spikelets longer than 3.0 mm D. setifolia

Spikelets shorter than 2.8 mm 17

17(16). Branched rhizome present; racemes 1-3, longer

than 100 mm D. eylesii

Branched rhizome absent; racemes 3-6, shorter

than 90 mm D. maitlandii

18(15). Nerves of the lower lemma thickened

D. comifera

Nerves of the lower lemma not thickened .... 19
19(18). Spikelets longer than 2.2 mm; pedicel with a corona

of hairs D. ternata

Spikelets shorter than 2.0 mm; pedicel without a

corona of hairs D. thouaresiana

20(11). Annuals, not mat-forming; stolons absent but culms
sometimes rooting from the lower nodes ... 21
Perennials or mat-forming annuals; stolons present

or absent 25

21(20). Nerves of the lower lemma scaberulous 22

Nerves of the lower lemma smooth 23

22(21). Lower lemma 2.3-3. 1 mm long, shorter to slightly
longer than the female-fertile (upper) lemma .

D. sanguinalis

Lower lemma 0.2-0. 5 mm long, longer than the

female-fertile (upper) lemma

D. acuminatissima

23(21). Racemes scattered on a central axis which is longer
thari the shortest racemes, but the longer racemes
are longer than the axis; racemes delicate and

spikelets loosely imbricate D. velutina

Racemes digitate, central axis if present not
exceeding the length of the shortest racemes;
racemes not delicate and spikelets not loosely

imbricate 24

24(23). Lower glume absent or a minute rim and obscure;

spikelet usually less than 2.3 mm long

D. nuda

Lower glume a distinct triangular scale; spikelet
usually more than 2.3 mm long .... D. ciliaris
25(20). Lower glume completely clasping the spikelet . .

D. maniculata

Lower glume not clasping the spikelet 26

26(25). Spikelets glabrous; upper glume as long as the
lower lemma, plants mat-forming

D. abyssinica

Spikelets hairy, or if glabrous then the upper glume
shorter than the lower lemma; plants usually

tufted, rarely mat-forming 27

27(26). Upper glume as long as the lower lemma,
concealing the female-fertile (upper) lemma . .

D. gazensis

Upper glume shorter than the lower lemma, partly
exposing the female-fertile (upper) lemma . 28

28(27). Inflorescence panicle-like, central axis longer than

the racemes D. rukwae

Inflorescence digitate, central axis not longer than

the racemes 29

29(28). Narrowest interspaces on lower lemma adjacent to
the central nerve; racemes often adhering to each

other by the hairs D. argyrograpta

Broadest interspaces on the lower lemma adjacent
to the central nerve, or interspaces of similar
width; racemes not adhering to each other by the

hairs 30

30(29). Lamina of the lower leaves reduced; culms mainly
branched at the middle nodes . . . D. polyphylla
Lamina of the lower leaves not reduced; culms
unbranched, or branched mainly at the lower

nodes 31

31(30). Rhizomes with innovations covered by hairy

cataphylls D. seriata

Rhizomes without innovations covered by hairy

cataphylls 32

32(31). Ligule longer than 4.0 mm; lower leaf sheaths rusty

brown D. natalensis

Ligule shorter than 3.5 mm; lower leaf sheaths not

rusty brown 33

33(32). Nerves of lower lemma scaberulous

D. milanjiana

Nerves of lower lemma smooth 34

34(33). Plants robust and always erect, densely tufted,
usually more than 350 mm tall; culms at base 2.5

mm or more across D. eriantha

Plants delicate and often decumbent, loosely tufted,
usually less than 350 mm tall; culms at base 2.0

mm or less across 35

35(34.) Spikelets longer than 3.0 mm . . D. diversinervis
Spikelets shorter than 2.8 mm D. didactyla

Digitaria abyssinica (A. Rich.) Stapf

( =D . scalarum (Schweinf.)

Chiov.) 4; ( =D . vestita Fig. &

De Not. var. scalarum
(Schweinf.) Henr.) 3.

Abyssinian finger grass.

Perennial; rhizomatous (mat-
forming); 200-350 mm tall. Leaf
blades 40-70 mm long; 3-6 mm wide. Spikelets 1.8-2. 2
mm long; 1 mm wide. Long wiry rhizomes; culms branched
at nodes; racemes 3-1 1, subdigitate, 20-80 mm long; spike-
lets paired, rarely solitary; upper glume and lower lemma
as long as spikelet, glabrous and often tinged with purple;
female-fertile (upper) floret light brown, grey and purple.

Flowering November to June. Mainly disturbed ground.
Locally common. Biome: Fynbos, Savanna, and Grassland.
Tropical Africa. Cultivated pasture and ornamental (lawns),
or weed.

Description: Clayton et al. 1970-1982 (641).

Illustration: Clayton et al. 1970-1982 (fig. 147). Voucher:
Smook 3573. PRECIS code 9900890-00100.

Digitaria acuminatissima Stapf

( -D . acuminatissima Stapf
subsp. inermis Goetghebeur) 4.

Annual; loosely tufted; 600-
1 200 mm tall. Leaf blades 30-250
mm long; 3-10 mm wide. Spike-
lets 2. 5-4.0 mm long; 1 mm wide.

Culms rooting at lower nodes; ra-
cemes 4—20, digitate to sub-
digitate, 70-250 mm long; lower lemma conspicuously

108

longer than spikelet, 0.2-0. 5 mm long, acuminate, scaberu-
lous along nerves.

Flowering February. Riversides and near damp rocks.
Infrequent. Biome: Savanna. Tropical Africa.

Description: Clayton et al. 1970-1982 (650). Voucher:
De Winter and Giess 7005. PRECIS code 9900890-00150.

Digitaria angolensis Rendle

Annual; loosely tufted; 150-
400 mm tall. Leaf blades 40-90
mm long; 4-6 mm wide. Spike-
lets 2. 0-2. 5 mm long; 0.8 mm
wide. Culms often decumbent; ra-
cemes 2-5, digitate or subdigi-
tate, 120-180 mm long; spikelets
in clusters of 3; rachis triquet-
rous; upper glume as long
as spikelet; upper glume and lower lemma covered with
silver hairs 1 mm long.

Flowering August. Sandy soil. Infrequent. Biome: Sa-
vanna. Tropical Africa southwards.

Description: Clayton et al. 1970-1982 (633).

Illustration: Kok 1978 (fig. 8.9). Voucher: Smook 1324.
PRECIS code 9900890-00250.

Digitaria argyrograpta (Nees) Stapf

Silver finger grass.

Perennial; rhizomatous and
tufted; 200-600 mm tall. Leaf
blades 40-200 mm long; 1-3 mm
wide. Spikelets up to 3 mm long;

0.8 mm wide. Rhizomes knotty,
culms profusely branched from
lower nodes; racemes usually in
pairs 40-100 mm long, often adhering to each other because
the hairs on the spikelets become entangled; lower glume
2/3-4/5 as long as spikelet; lower lemma 7-nerved with 3
nerves close together in the middle, thus with narrow inter-
spaces adjacent to central nerve.

Flowering November to March. Wide range of habitats.
Common. Biome: Savanna, Grassland, and Nama-Karoo.
Mozambique. Natural pasture.

Description: Chippindall 1955 (414). Illustration: Chip-
pindall 1955 (fig. 346). Voucher: Smook 993. PRECIS code
9900890-00400.

Digitaria brazzae (Franch.) Stapf

Brown finger grass.

Perennial; densely tufted;

500-1100 mm tall. Leaf blades
60-100 mm long; 2^1 mm wide.

Spikelets 2. 8-3. 2 mm long; 1 mm
wide. Cataphylls and basal
sheaths densely covered with
silky hairs; racemes 2^1, digitate,

150-200 mm long; spikelets in clusters of 2— 4(— 5); upper
glume 1/2-3/4 as long as spikelet; lower lemma depressed
and hyaline beside the midnerve, with hairs 2-4 mm long
forming 2 tufts at the base and fringing the upper half of
the margin.

Flowering September to April. Grows in grassland,
usually sandy soil, often on stony hillsides. Locally com-
mon. Biome: Savanna and Grassland. Tropical and sub-
tropical Africa.

Description: Clayton et al. 1970-1982 (627).

Illustration: Kok 1978 (fig. 8.16). Voucher: Du Toit 1236.
PRECIS code 9900890-00600.

Digitaria ciliaris (Retz.) Koeler

(=D. adscendens Henr.) 2;

( -D . marginata Link) 4.

Tropical finger grass.

Annual; tufted; 200-550 mm
tall. Leaf blades 30-160 mm
long; 3-10 mm wide. Spikelets
2. 3-3.4 mm long; 1 mm wide.

Racemes 3-7, digitate, 40-100 mm long; lower glume a tri-
angular scale up to 0.3 mm long; lower lemma not
scaberulous.

Flowering January to April. Mainly disturbed sandy soil.

109

Locally common. Biome: Savanna, Grassland, and Nama-
Karoo. Pan-tropical. Weed.

Description: Chippindall 1955 (399). Illustration: Kok
1978 (fig. 8.23). Voucher: Smook 1982. PRECIS code
9900890-00700.

Digitaria comifera Pilg.

Annual; tufted; 300-800 mm
tall. Leaf blades 40-100 mm
long; 2-5 mm wide. Spikelets
2. 2-2. 7 mm long; 1 mm wide.

Racemes 2-7, subdigitate, 30-
150 mm long; spikelets in clusters
of 3-5 on a winged rachis with
sharply angular midrib; upper
glume 3/4 as long as spikelet,
covered with rows of white clavate hairs; lower lemma gla-
brous or hairy, midrib prominent because hyaline
interspaces allow the dark brown colour of the lower floret
to show through.

Flowering February. Open sandy places by roadsides.
Infrequent. Biome: Savanna. East Africa.

Description: Clayton et al. 1970-1982 (630). Voucher:
De Winter 9125. PRECIS code 9900890-00750.

Digitaria debilis (Desf.) Willd.

Finger grass.

Annual; tufted; 100-500 mm
tall. Leaf blades 20-25 mm long;

2^1 mm wide. Spikelets 2. 1-3.1
mm long; 1 mm wide. Culms de-
cumbent at base and rooting from
the lower nodes; racemes 60-160
mm long, 4—12, subdigitate, on a
central axis up to 60 mm long; spikelets solitary or paired;
lower and upper glume separated by an intemode 0. 1-0.3
mm long; upper glume longer than spikelet but less than 4.5
mm long and tip acuminate.

Flowering November to June. Mainly damp places and
sandy soil. Locally dominant. Biome: Fynbos and Savanna.
Central and north Africa and southern Europe.

Description: Chippindall 1955 (396), Clayton et al.
1970-1982 (637). Illustration: Kok 1978 (fig. 8.4).
Voucher: Smook 4183. PRECIS code 9900890-00800.

Digitaria diagonalis (Nees) Stapf var. diagonalis

Fig. 65.

(=D. trichopodia Stent) 4;

(=D. uniglumis (A. Rich.)

Stapf) 2.

Brown seed finger grass.

Perennial; tufted; 400-1500
mm tall. Leaf blades 90-170 mm
long; 2-3 mm wide. Spikelets
1. 4-2.4 mm long; 1 mm wide.

Culms swollen and bulbous at
base; basal leaf sheaths silky-
hairy, breaking up into fibres; inflorescence a panicle;
spikelets in groups of 3(-6), on unequal pedicels with
10-15 white setae as long as or longer than spikelets;
female-fertile (upper) floret bright dark brown or black.

Flowering January to April. Grows in open, usually
sourveld grassland, often hillsides and in damp places. Lo-
cally dominant. Indigenous. Biome: Savanna and Grass-
land. Two additional varieties occur in eastern and western
Africa. May be confused with D. eylesii, which lacks long
setae and has racemes in pairs or threes.

Fig. 64.

Fig. 65. Digitaria diagonalis var. diagonalis

Description: Chippindall 1955 (419), Clayton et al.
1970-1982 (625). Illustration: Chippindall 1955 (fig. 349),
Clayton et al. 1970-1982 (fig. 145). Voucher: Davidse
6759. PRECIS code 9900890-01000.

110

Digitaria didactyla Willd.

( -D . swazilandensis Stent) 4.

Blue couch, Swaziland finger
grass.

Perennial; weakly tufted, or sto-
loniferous (and mat-forming);

150-300 mm tall. Leaf blades
25-50 mm long; 1.0-1. 8 mm
wide. Spikelets 2. 5-2. 8 mm long;

0.8 mm wide. Rhizomes knotty; culms may be branched and
rooted at lower nodes; racemes 2-4, digitate, 30-65 mm
long.

Disturbed sandy soil. Locally common. Madagascar.
Biome: Savanna and Grassland. Cultivated worldwide. Or-
namental (lawns). Spikelet similar to some forms of D.
eriantha where the spikelet is 2. 2-4.0 mm long.

Description: Veldkamp 1973(44). Voucher: Smook
1986. PRECIS code 9900890-01050.

Digitaria diversinervis (Nees) Stapf

(=D. albomarginata Stent) 4;

(=D. diversinervis (Nees) Stapf
var. woodiana Henr.) 2.

Richmond and Wynberg fin-
ger grass.

Perennial; rhizomatous and sto-
loniferous (and mat-forming);

200-350 mm tall. Leaf blades 20-90 mm long; 3-8 mm
wide. Spikelets 3-4 mm long; 1 mm wide. Rhizomes
knotty, much branched; racemes 2-5, digitate, 30-77 mm
long; lower glume a well developed scale up to 1 mm long;
upper glume 1/2-2/3 the length of the spikelet; upper glume
and lower lemma glabrous or hairy along margins.

Flowering November to June. Mainly sandy coastal
areas. Locally common. Biome: Fynbos, Savanna, and
Grassland. Endemic. Ornamental (lawn grass, grows well
in shade).

Description: Chippindall 1955 (398). Illustration: Chip-
pindall 1955 (335). Voucher: Smook 1855. PRECIS code
9900890-01200.

Digitaria eriantha Steud.

Fig. 63. PI. 58.

(=D. bechuanica (Stent)

Henr.) 2; (=D. decumbens Stent)

2; (=D. dinteri Henr.) 4; (= D .
eriantha subsp. pentzii (Stent)

Kok) 4; (=D. eriantha subsp.
stolonifera (Stapf) Kok) 4; (=D.
eriantha subsp. transvaalensis
Kok) 4; (-D. eriantha var.
stolonifera Stapf) 2; ( -D .
geniculata Stent) 2; (=D. glauca sensu Stent, non A.
Camus) 2; (=D. pentzii Stent) 2; ( =D . pentzii Stent var.
stolonifera (Stapf) Henr.) 2, 4; (=D. setivalva Stent) 4;

(=D. smutsii Stent) 2; (=D. stentiana Henr.) 2; (=D. valida
Stent) 2; (=D. valida Stent var. glauca Stent) 2.

Common finger grass.

Perennial; sometimes stoloniferous, or tufted and rhizo-
matous; 350-1400 mm tall. Leaf blades 50-400 mm long;
2-14 mm wide. Spikelets 2. 2^1.0 mm long; 1 mm wide.
Rhizomes knotty and unbranched; culms simple or
branched, basal leaf sheaths silky and hairy, racemes 3-15,
digitate, 50-200 mm long; lower glume a membranous
scale, ovate to acute; upper glume 1/3— 2/3 as long as spike-

let; lower lemma as long as spikelet; upper glume and lower
lemma covered with hairs 1 mm long, purple and silvery.

Flowering January to April. Occurs in a wide range
habitats. Dominant. Biome: Fynbos, Savanna, Grassland,
and Nama-Karoo. Zimbabwe. Pasture (cultivated and
natural). A very variable grass with many forms.

Description: Chippindall 1955 (403). Illustration: Chip-
pindall 1955 (fig. 338). Voucher: Smook and Gibbs Russell
2166. PRECIS code 9900890-01400.

Digitaria eylesii C.E. Hubb.

Perennial; rhizomatous, or
tufted; 400-65 0(- 1000) mm tall.

Leaf blades 50-130 mm long; 2-4
mm wide. Spikelets 2. 3-3.0 mm
long; 1 mm wide. Rhizomes
creeping, branched, with cata-
phy 11s; racemes usually in pairs or
threes, or rarely solitary, 100-180
(-240) mm long; female-fertile
(upper) floret purplish brown or
black.

Flowering January to April. Grows in wet places. Local-
ly common. Biome: Savanna and Grassland. Zambia and
Zimbabwe. See comment under D. diagonalis var.
diagonalis.

Description: Chippindall 1955 (419). Illustration: Kok
1987 (fig. 8.14). Voucher: De Winter 739. PRECIS code
9900890-01500.

Digitaria flaccida Stapf

Flaccid finger grass.

Perennial; densely tufted and
rhizomatous; 250-400(-600) mm
tall. Leaf blades 40-100 mm
long; 1 .5-3.0 mm wide. Spikelets

1- 3 mm long; 1 mm wide. Knotty
rhizomes with hairy cataphylls;
inflorescence panicle-like with
6-17 racemes, 30-55 mm long; spikelets paired; upper
glume and lower lemma covered with silky purplish hairs
up to 2 mm long.

Flowering November to January. Mainly on rocky
ground, mountain sourveld. Locally common. Biome:
Grassland. Central Africa southwards. The silky hairy pani-
cle resembles those in Melinis and Brachiaria species.

Description: Chippindall 1955 (400). Illustration: Kok
1978 (fig. 8.17). Voucher: Smook 1399. PRECIS code
9900890-01600.

Digitaria gayana (Kunth) Stapf

Annual; loosely tufted; 1 30—

300 mm tall. Leaf blades 20-70
mm long; 3. 5-6.0 mm wide.

Spikelets 2. 4-2. 8 mm long; 1 mm
wide. Lower leaf sheaths glab-
rous or sparsely hairy; racemes

2- 3, digitate, 40-100 mm long;
spikelets in clusters of 3( — 4);
upper glume 4/5 length of spike-
let, covered with silvery hairs 1 mm long; lower lemma with
membranous interspaces, depressed on either side of
midnerve, margin with copious purplish or silvery hairs 2-4
mm long.

Flowering February. A weedy species of disturbed areas,
mainly sandy soils. Infrequent. Biome: Savanna. Tropical
and sub-tropical Africa. Weed.

Description: Clayton et al. 1970-1982 (627).

Illustration: Kok 1987 (fig. 8.17). Voucher: De Winter
9203. PRECIS code 9900890-01700.

Ill

Digitaria gazensis Rendle

Perennial; usually rhizomat-
ous and stoloniferous (rarely), or
tufted (densely); 350-700 mm
tall. Leaf blades 50-130 mm
long; 3. 0^1. 5 mm wide. Spikelets
1. 8-2.4 mm long; 0.8 mm wide.

Rhizome short and knotty; basal
sheaths hairy;racemes 5-10, sub-
digitate, 30-70 mm long; spike-
lets paired; upper glume as long as or slightly shorter than
spikelet; upper glume and lower lemma covered with 2 mm
long hairs, often purplish; female-fertile (upper) floret
yellowish to dark purple-grey.

Flowering January to March. Mainly sandy soil but not
in flood plain. Infrequent. Biome: Savanna. Tropical and
sub-tropical Africa and Madagascar. Close to D. rukwae
which has a brown upper floret and grows in flood plains.

Description: Clayton et al. 1970-1982 (643). Voucher:
De Winter & Wiss 4450. PRECIS code 9900890-01800.

Digitaria gymnostachys Pilg.

Perennial; loosely tufted; 600-
1000 mm tall. Leaf blades 60-
300 mm long; 7-15 mm wide.

Spikelets h-A mm long; 1 mm
wide. Racemes 6-13, 100-300
mm long, subdigitate on central
axis up to 60 mm long; spikelets
paired; lower and upper glumes
are small scales 0.2-0. 5 mm long,
separated from rest of spikelet by an intemode 0.2-0. 5 mm
long.

Flowering February to April. Mainly sandy soil. Infre-
quent. Biome: Savanna. Tropical Africa.

Description: Clayton et al. 1970-1982 (636). Voucher:
Ward 3852. PRECIS code 9900890-02000.

Digitaria longiflora (Retz.) Pers.

False couch finger grass.

Perennial (short lived), or an-
nual; mat-forming and stolonifer-
ous, or tufted (loosely); 100-350
mm tall. Leaf blades 18-70 mm
long; 1.6-4. 5 mm wide. Spikelets
1.2-1 .6 mm long; 0.6 mm wide.

Culms rooting from lower nodes;
racemes 2(-3), digitate, 30-70 mm long; spikelets in pairs
or threes and tend to bend outwards; rachis winged with low
rounded midrib; upper glume and lower lemma as long as
spikelet and covered with hairs 0.2 mm long; female-fertile
(upper) lemma grey to purplish.

Flowering October to June. Disturbed areas, mainly
sandy soil. Common. Biome: Savanna. Pan-tropical. Orna-
mental (lawns), or weed.

Description: Clayton et al. 1970-1982 (635).

Illustration: Kok 1987 (fig. 8.10). Voucher: Smook 3157.
PRECIS code 9900890-02100.

Digitaria maitlandii Stapf & C.E. Hubb.

(=D. apiculata Stent) 3.

Perennial; tufted; 220^110
mm tall. Leaf blades 40-130 mm
long; 2^1 mm wide. Spikelets 1-2
mm long; 0. 8-1.0 mm wide. No
rhizome; racemes 3-6, 50-80 mm
long, subdigitate on a short cen-
tral axis; spikelets in groups of
3, glabrous or with a few hairs; female-fertile (upper) floret

purplish brown or black.

Flowering November to May. Mountain grassland. In-
frequent. Biome: Grassland. Mozambique and up to
Zambia.

Description: Chippindall 1955 (418). Illustration: Kok
1987 (fig. 8.15). Voucher: Scheepers 831. PRECIS code
9900890-02200.

Digitaria maniculata Stapf

Annual; partially mat-form-
ing; 80-150 mm tall. Leaf blades
10-30 mm long; 1.5-3 mm wide.

Spikelets 2. 5-2.7 mm long; 0.8
mm wide. Culms rooting from
lower nodes; racemes 2-3, digi-
tate, 40-80 mm long; spikelets
paired; lower glume clasping the
spikelet and hairy; upper glume
and lower lemma as long as spikelet and covered with hairs
0.2 mm long.

Flowering December to March. Sandy flats near rivers.
Infrequent. Biome: Savanna. Zaire southwards.

Description: Kok 1987 (176). Illustration: Kok 1978
(fig. 8.8). Voucher: De Winter and Marais 4638. PRECIS
code 9900890-02400.

Digitaria milanjiana (Rendle) Stapf

Milanje finger grass.

Perennial; rhizomatous and
stoloniferous, or tufted; 500-
1300 mm tall. Leaf blades 60-
300 mm long; 2-8 mm wide.

Spikelets 2. 5-3. 2 mm long; 0.8
mm wide. Rhizomes branched,
slender and elongate; culms usu-
ally straight and erect; racemes 3-12, digitate, 80-250 mm
long; spikelet similar toD. eriantha but hairs yellowish or
brown and nerves of lower lemma scaberulous.

Flowering January and February. Occurs in a wide range
of habitats, often in disturbed areas. Locally common.
Biome: Savanna. Tropical and sub-tropical Africa. Natural
pasture. A variable tropical species, according to Clayton
et al. 1982 (648) which can be separated from its southern
allies D. eriantha and D. didactyla by the scabrid nerves
on the lower lemma.

Description: Chippindall 1955 (407), Clayton et al.
1970-1982 (647). Illustration: Kok 1978 (fig. 8.31).
Voucher: Voster 2779. PRECIS code 9900890-02500.

Digitaria monodactyla (Nees) Stapf

PI. 59.

( -D . monodactyla Stapf var.
explicata Stapf) 2.

One finger grass.

Perennial; densely tufted;

200-550 mm tall. Leaf blades
30-60 mm long; 1 .8-2.2 mm
wide. Spikelets 2. 8-3. 2 mm long;

1 mm wide. Culm with hairy cataphylls; leaf blades often
rolled; raceme solitary, 50-180 mm long; spikelets in pairs;
upper glume and lower lemma covered with yellow hairs
1 mm long; female-fertile (upper) floret light brown.

Flowering November to February. Open grassland,
usually highland sourveld. Locally dominant. Biome: Sa-
vanna and Grassland. Tropical Africa southwards. May be
confused with Elionurus muticus, which has a sessile spike-
let and a 2-lobed lower glume.

Description: Chippindall 1955 (415). Illustration: Chip-
pindall 1955 (fig. 347). Voucher: Smook 4789. PRECIS
code 9900890-02700.

112

Digitaria natalensis Stent

Digitaria polyphylla Henr.

( -D . lirtoralis sensu Stent,
non Salisb.) 2; (=D. littoralis
sensu Stent, non Salisb. var.
prostrata Stent) 4; ( =D .
macroglossa Henr.) 2; ( =D .
macroglossa Henr. var.
prostrata (Stent) Henr.) 2; ( =D .
natalensis Stent subsp. stentiana
Henr.) 2; ( -D . rigida Stent) 2.

Coast finger grass.

Perennial; loosely to densely tufted and rhizomatous;
600-1500 mm tall. Leaf blades 80-250 mm long; 2-6 mm
wide. Spikelets 2. 8-3. 5 mm long; 0.8 mm wide. Rhizomes
knotty; culm lower nodes straight and covered by bright,
reddish-brown sheaths; ligule membranous, 3.5-12.0 mm
long; racemes 4-12, digitate, 100-200 mm long; spikelet
characters similar to D. eriantha.

Flowering December to June. Mainly sandy ground. Lo-
cally dominant. Biome: Fynbos, Savanna, and Grassland.
Mozambique.

Description: Chippindall 1955 (407). Illustration: Kok
1987 (fig. 8.32). Voucher: Smook 5759. PRECIS code
9900890-02750.

Digitaria nuda Schumach.

( -D . borbonica Desv.) 3.

Annual; tufted; 200-500 mm
tall. Leaf blades 20-135 mm
long; 2-10 mm wide. Spikelets
1.6-2. 3 mm long; 0.8 mm wide.

Culms decumbent at base, geni-
culate and ascending, rooting
from lower nodes; racemes 3-8,
digitate or subdigitate, 40-100 mm long; lower glume
absent, or if present then a minute rim, obscure.

Flowering November to April. Open disturbed areas.
Locally common. Biome: Savanna. Mainly tropical Africa,
also Brazil and Indonesia. Weed.

Description: Clayton et al. 1970-1982 (654). Voucher:
Du Toil PRE 58243. PRECIS code 9900890-02900.

Digitaria perrottetii (Kunth) Stapf

Whorled finger grass.

Annual; loosely tufted; 400-
800 mm tall. Leaf blades 30-90
mm long; 3-15 mm wide. Spike-
lets 1 .5-2.0 mm long; 1 mm wide.

Culms decumbent and rooting
from lower nodes; panicle
80-200 mm long, racemes ar-
ranged in 8-12 whorls on a central axis, each whorl consists
of 4-8 racemes 20-50 mm long; spikelets solitary or in
pairs, widely spaced on axis; upper glume as long as spike-
let.

Flowering January to June. Damp, shady, sandy areas.
Locally common. Biome: Savanna. Central Africa and
Madagascar.

Description: Chippindall 1955 (400). Illustration: Kok
1987 (fig. 8.20). Voucher: Smook 1985. PRECIS code
9900890-03100.

Perennial; rhizomatous and
stoloniferous, or tufted (densely);

300-500 mm tall. Leaf blades
50-130 mm long; 2. 5-5.0 mm
wide. Spikelets 2. 3-3.0 mm long;

1 mm wide. Rhizome knotty,
culms with lower nodes unbran-
ched, while middle and upper
nodes are profusely branched
and form leafy tufts; racemes 2-6, digitate, 30-80 mm
long; spikelet characters similar to D. eriantha.

Flowering February to April. Sandy and stony ground,
low rainfall areas. Locally common. Biome: Savanna.
Endemic.

Description: Chippindall 1955 (414). Illustration: Chip-
pindall 1955 (fig. 345). Voucher: Acocks 2078. PRECIS
code 9900890-03300.

Digitaria remotigluma (De Winter) Clayton

(-Digitariella remotigluma
De Winter) 3.

Annual; tufted; 100—400 mm
tall. Leaf blades 15-80 mm long;

1-3 mm wide. Spikelets 7-10 mm
long; 1 mm wide. Culms decum-
bent at base and rooting from
lower nodes; racemes 2-5, digi-
tate, 25-120 mm long; spikelets in pairs; intemode 0. 7-2.0
mm long between lower and upper glume; upper glume
acuminate and 7-10 mm long; lower lemma finely acute,
13-16 mm long.

Flowering November to February. Damp, sandy soils.
Infrequent. Biome: Savanna. East Africa and southwards.
The awnlike tip to the upper glume and lower lemma are
characteristic for this species.

Description: Clayton et al. 1970-1982 (638).

Illustration: Kok 1978 (fig. 8.5). Voucher: Smith 2659.
PRECIS code 9900890-03350.

Digitaria rukwae Clayton

Perennial; rhizomatous and
tufted (loosely); 250-1200 mm
tall. Leaf blades 50-300 mm
long; 2-7 mm wide. Spikelets
2. 0-2. 6 mm long; 0.8 mm wide.

Rhizomes scaly;basal sheaths gla-
brous or silky pubescent; racemes
6-25, 40-100 mm long, borne
irregularly or in untidy whorls
upon an axis 40-200 mm long; upper glume 3/4 to as long
as to equalling spikelet; lower lemma as long as the spikelet
with fine brown hairs mainly along the margins; female-
fertile (upper) floret ellipsoid, brown.

Flowering March. Flood plains and alluvial soils. Infre-
quent. Biome: Savanna. Northwards to Tanzania. Close to
D. gazensis, which has the upper floret pale grey or
yellowish to dark purple-grey and does not grow in
floodplains.

Description: Clayton et al. 1970-1982 (646). Voucher:
Ellis 4517. PRECIS code 9900890-03360.

1 13

Digitaria sanguinalis (L.) Scop.

Fig. 66.

Crab finger grass.

Annual; tufted; 200-600 mm
tall. Leaf blades 27-170 mm
long; 3-10 mm wide. Spikelets
2.3-3. 1 mm long; 1 mm wide.

Culms branched, decumbent and
rooting from lower nodes; ra-
cemes 3-12, digitate or subdigi-
tate, 40-105 mm long; lower lemma equalling or shorter
than spikelet, scaberulous along nerves, very rarely with a
ciliate frill.

Flowering November to May. Disturbed areas. Locally
dominant. Naturalized from Europe. Biome: Fynbos, Sa-
vanna, Grassland, Nama-Karoo, and Succulent Karoo. Pan-
temperate. Weed.

Description: Chippindall 1955 (399). Illustration: Kok
1987 (fig. 8.22). Voucher: Smook 4674. PRECIS code
9900890-03370.

Digitaria seriata Stapf
( =D . polevansii Stent) 4.

Kuruman finger grass.

Perennial; loosely or densely
tufted, stoloniferous, and rhizo-
matous; 500—1200 mm tall. Leaf
blades 100-350 mm long; 3-9
mm wide. Spikelets 2.5-3.2 mm long; 1 mm wide. Rhi-

zomes knotty, branched and woody; base of culm bulbous
and covered with densely hairy scales; racemes 3-12, digi-
tate to subdigitate, 100-230 mm long; spikelets like those
of D. e riant ha.

Flowering January to April. Mainly sandy soil. Locally
common. Biome: Savanna and Grassland. Zimbabwe.

Description: Chippindall 1955 (414). Illustration: Chip-
pindall 1955 (fig. 344). Voucher: De Winter 7377. PRECIS
code 9900890-03450.

Digitaria setifolia Stapf

Fine-leaved finger grass.

Perennial; densely tufted;

200-500 mm tall. Leaf blades
50-140 mm long; 1. 5-3.0 mm
wide. Spikelets 3-4 mm long; 1
mm wide. Old leaf sheaths break
up into brown fibres at base; leaf
blades usually rolled; racemes
40-120 mm long, (2— )3 — 4(— 5), subdigitate on a short
central axis; spikelets in pairs or threes; upper glume and
lower lemma with conspicuous rows of bright brown,
clavate hairs; female-fertile (upper) floret dark brown.

Flowering September to January. Grows in mountain
sourveld areas, usually in damp places or vleis. Locally
common. Biome: Grassland. From the Congo south,
excluding east Africa and Angola.

Description: Chippindall 1955 (416). Illustration: Kok
1987 (fig. 8.13). Voucher: Smook 2570. PRECIS code
9900890-03550.

Digitaria ternata (A. Rich.) Stapf

Blackseed finger grass.

Annual; loosely tufted; 200-
600 mm tall. Leaf blades 20-200
mm long; 1-7 mm wide. Spike-
lets 1.5-2. 3 mm long; 0.8 mm
wide. Racemes 40-200 mm long,

2-6, in pairs or subdigitate on
short central axis, midrib wing-
ed; spikelets in clusters of (2— )3; pedicels with a corona of
short hairs at apex; upper glume 3/4, as long as spikelet;
upper glume and lower lemma covered with white clavate
hairs; female-fertile (upper) floret dark brown or purplish
black.

Flowering November to May. Damp and disturbed areas.
Locally dominant (roadsides). Biome: Savanna and Grass-
land. Africa to Far East.

Description: Chippindall 1955 (418). Illustration: Chip-
pindall 1955 (fig. 348). Voucher: Smook 4712. PRECIS
code 9900890-04100.

Digitaria thouaresiana (Fluegge) A. Camus

( -D . melanochila Stapf) 3;

(-D. tricostulata (Hack.)

Henr.) 3.

Annual; tufted; 20-100 mm
tall. Leaf blades 3-20 mm
long; 2-8 mm wide. Spikelets
1.0-1. 7 mm long; 0.8 mm wide.

Racemes 2-14, subdigitate, 20-
120 mm long; spikelets in clusters of 3-4 on winged rachis;
pedicels without corona of hairs, only a few hairs at apex;
upper glume and lower lemma rarely glabrous, usually
covered with white clavate hairs; female-fertile (upper)
floret dark brown to black.

Flowering February to May. Marshy places and disturb-
ed sites. Locally common. Biome: Savanna and Grassland.
Ecist Africa

Description: Clayton et al. 1970—1982 (631). Voucher:
Smook 3071. PRECIS code 9900890-04300.

114

Digitaria tricholaenoides Stapf

Purple finger grass.

Perennial; rhizomatous and
tufted; 200-550 mm tall. Leaf
blades 40-200 mm long; 2-7 mm
wide. Spikelets 3-5 mm long; 1
mm wide. Rhizome oblique,up to
100 mm long, covered with hairy,
persistent, overlapping bases of

Fig. 67.

old leaf sheaths; racemes 2— 3(— 7), subdigitate, 30-130 mm
long; spikelets in clusters of 2-5; upper glume 1/2-2/3
length of spikelet, covered with silvery or purplish hairs
1-2 mm long; lower lemma covered (except for central
interspace) with silvery or purplish hairs 2-4 mm long.

Flowering November to March. In open, sourveld
grassland mainly on stony soil. Locally dominant. Biome;
Savanna and Grassland. Endemic.

Description: Chippindall 1955 (402). Illustration: Chip-
pindall 1955 (fig. 337). Voucher: Smook 4932. PRECIS
code 9900890-04400.

Digitaria velutina (Forssk.) Beauv.

( -D . zeyheri (Nees) Henr.) 3.

Long-plumed finger grass.

Annual; very loosely tufted;

150-800 mm tall. Leaf blades
40-150 mm long; 3-10 mm wide.

Spikelets 1. 5-2.0 mm long; 0.5
mm wide. Culms decumbent at
base, rooting from lower nodes; inflorescence panicle-like,
racemes 35-100 mm long, 5-15, subdigitate on a relatively
long central axis 25-50 mm long; lower glume obscure or
an ovate scale up to 0.2 mm long; upper glume and lower
lemma almost glabrous.

Flowering December to May. Open disturbed areas. Lo-
cally dominant. Biome: Savanna and Grassland.

Northwards to Egypt. Weed.

Description: Chippindall 1955 (400). Illustration: Chip-
pindall 1955 (fig. 336). Voucher: Smook 2655. PRECIS
code 9900890-04900.

Digitaria violascens Link.

Annual; tufted; to 900 mm
tall. Leaf blades 40-220 mm
long; 3-6 mm wide. Spikelets
1.35-1.7 mm long; 0.8 mm wide.

Culms erect or decumbent and
rooting from lower nodes; ra-
cemes 3-6, digitate to subdigi-
tate, 30-50 mm long; upper
glume and lower lemma as long
as spikelet, glabrous or with minute hairs, green veins very
distinctive; female-fertile (upper) lemma dark brown.

Flowering March. Disturbed areas and woodland
margins. Infrequent. Naturalized, area of origin unknown.
Biome: Grassland. Tropics and sub-tropics of world. Weed.

Description: (Webster 1983 (21 1)). Voucher: Ellis 4416.
PRECIS code 9900890-04970.

Diheteropogon Stapf

Annual, or perennial (slender); caespitose. Culms
150-2300 mm high; herbaceous (to woody at base); un-
branched above (mainly). Leaf blades linear or linear-
lanceolate; flat. Ligule an unfringed membrane. Plants
bisexual, with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant (all male or sterile at bases
of ‘racemes’ , heterogamous above)', overtly heteromorphic
(pedicellate spikelets awnless or aristulate, larger, callus
glabrous).

Inflorescence of spike-like main branches, or paniculate
(of paired ‘racemes’ , terminal or in a scanty false panicle);
spatheate; a complex of ‘partial inflorescences’ and inter-
vening foliar organs. Spikelet-bearing axes ‘ racemes’ ;
paired (not deflexed); with substantial rachides; disarticu-
lating at the joints. ‘Articles’ non-linear (thickened and
hollowed at the summit).

Spikelets in pairs; consistently in ‘long-and-short’ com-

115

binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite (save at the base
of the raceme). The pedicellate spikelets male-only, or
sterile (?). Female-fertile spikelets 5-9 mm long; com-
pressed dorsiventrally; falling with the glumes (or with a
slight tendency to disarticulate above them). Callus long.
Glumes two; more or less equal; awnless; very dissimilar
(somewhat leathery; G1 bicarinate, grooved between the
keels; G2 not bicarinate). Proximal incomplete florets 1\
epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); incised; awned. Awns 1; median; from
the sinus; geniculate (twice-geniculate); much longer than
the body of the lemma (and sturdy). Palea present; conspic-
uous but relatively short (small). Lodicules 2. Stamens 3.
Ovary glabrous.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 5 species. Tropical Africa. Helo-
phytic; in open habitats (savanna); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 2 indigenous species.

References. 1. Clayton. 1966. Kew Bull. 20: 75. 2.
Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

1(0). Leaf blades filiform, usually narrower than 3 mm if
unrolled, base not rounded; lower glume of sessile
spikelet acutely bent on either side of the deep

median longitudinal groove D. filifolius

Leaf blades expanded, usually wider than 3 mm, base
rounded or amplexicaul; lower glume of sessile
spikelet rounded on either side of the deep median
longitudinal groove D, amplectens

Diheteropogon amplectens (Nees) Clayton

Fig. 68. PI. 60.

{■^Andropogon amplectens
Nees) 1.

Broad-leaved bluestem, bree-
blaarandropogon.

Perennial; rhizomatous; 300-
2000 mm tall. Leaf blades 1 50—

300 mm long; to 20 mm wide.

Spikelets (sessile) 7-9 mm long (pedicellate longer). Leaf
blade bases rounded, clasping the stem; young growth
waxy.

Flowering November to April. Poor shallow soils on
stony slopes and in woodland. Common. Biome: Savanna
and Grassland. Tropical Africa. Cymbopogon excavatus has
similar leaf blades, but it is aromatic and often has swollen
raceme bases.

Description: Chippindall 1955 (498), Clayton et al.
1970-1982 (784). Illustration: Chippindall 1955 (pi. 21),
Clayton et al. 1970-1982 (fig. 182), Flower. PI. Afr. (24:
922). Voucher: Moll 617. PRECIS code 9900810-00100.

Diheteropogon filifolius (Nees) Clayton

(=Andropogon filifolius
(Nees) Steud.) 1.

Draadbloustam, thread-leaved
andropogon.

Perennial; rhizomatous and
tufted; 150-600 mm tall. Leaf
blades 100-500 mm long;
filiform or to 3.5 mm wide. Spikelets (sessile) 6-8 mm long
(pedicellate much longer). Leaf blades thread-like, bluish-
green.

Flowering October to April. Sour open grassveld on
hillsides. Common. Biome: Savanna and Grassland.
Southern tropical Africa. Andropogon schirensis is closely
related, but has expanded leaf blades and a shorter callus.

Description: Chippindall 1955 (497). Illustration: Chip-
pindall 1955 (fig. 401). Voucher: Pole Evans 1009. PRECIS
code 9900810-00200.

1 16

Dinebra Jacq.

Annual ; caespitose to decumbent. Culms 150-1200 mm
high; herbaceous; unbranched above. Leaf blades usually
flat. Ligule a fringed membrane (very narrow).

Inflorescence of spike-like main branches (a raceme of
numerous small spikes which become deflexed at maturity,
the lower spikelets of each spike often replaced by small
deciduous branchlets)', espatheate. Spikelet-bearing axes
disarticulating, or persistent; falling entire (the smaller
laterals deciduous).

Spikelets solitary; biseriate; all sessile ; 3.5-10 mm long;
compressed laterally; disarticulating above the glumes; dis-
articulating between the florets (when two or more florets).
Glumes present; two; more or less equal; much exceeding
the spikelets; awned (acuminate-aristate); very dissimilar,
or similar (leathery or membranous, the lower often very
asymmetrical). Incomplete florets distal to the female-
fertile florets; proximal incomplete florets absent.

Female-fertile florets 1-2. Lemmas less firm than the
glumes to similar in texture to the glumes (thinly
membranous); 3 nerved; awnless to mucronate. Palea
present. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small; ellipsoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions absent. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 3
species. Tropical Africa, Asia. Helophytic to mesophytic
(in seasonally wet places); in shade, or in open habitats
(savanna); glycophytic. Namibia, Botswana, Transvaal,
Swaziland, Natal, and Cape Province. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Dinebra retroflexa (Vahl) Panz. var. condensata S.M.

Phillips

Fig. 69. PI. 61.

Kattestertgras, catstail grass.

Annual; loosely tufted;

130-820 mm tall. Leaf blades
45-280 mm long; 4—8 mm wide.

Spikelets 5. 7-9.0 mm long.

Spikes up to 50 mm long; glumes
6. 0-8. 2 mm long, with spreading
aristate tips.

Flowering December to May. Usually on disturbed soil
in moist weedy places, often in the shade and on black turf
or waterlogged soils. Common. Biome: Savanna. Tropical
Africa through Egypt and Iraq to India. Weed (in ricefields
mostly).

Description: Phillips 1974 Kew Bull. 28(3),Chippindall
1955 (185), Clayton et al. 1970-1982 (273). Illustration:
Chippindall 1955 (fig. 160), Clayton et al. 1970-1982 (fig.
75). Voucher: Acocks 16804. PRECIS code

9903300-00100.

Diplachne P. Beauv.

Sometimes included in Leptochloa.

Perennial ; long-stoloniferous, or caespitose (some tall).
Culms 300-2700 mm high; herbaceous; unbranched above.
Leaf blades linear; flat, or rolled (often involute). Ligule an
unfringed membrane, or a fringed membrane ( sometimes
much reduced).

Inflorescence of spike-like main branches (a contracted
panicle of spikelike racemes ), or a single raceme (rarely),
or paniculate-, open ; digitate or subdigitate, or non-digitate;
espatheate. Spikelet-bearing axes persistent.

Spikelets not secund (or scarcely so)’, biseriate; 6—15
mm long (narrow); not noticeably compressed to com-
pressed dorsiventrally (more or less terete ); disarticulating
above the glumes; disarticulating between the florets.
Glumes two; relatively large; very unequal, or more or less
equal; markedly shorter than the spikelets; awnless; similar
(membranous). All florets female-fertile, or distal incom-
plete florets also present, these awnless; proximal incom-
plete florets absent.

Female-fertile florets 5-20. Lemmas 3 nerved; entire, or
incised (bidentate); awnless, or mucronate (from the sinus),
or awned. Awns when present 1; from the sinus, or dorsal;
non-geniculate; much shorter than the body of the lemma.
Palea present. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small; hilum short; pericarp free, or
loosely adherent, or fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 18
species. Tropical and subtropical. Helophytic, or meso-
phytic, or xerophytic; in shade and in open habitats
(woodland, savanna, dry and swampy soils); halophytic and

117

glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 4 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA. 3. Launert. 1974. Bol. Soc. Brot. Ser. 2. 47:
349. 4. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

1(0). Ligule a fringe of hairs, to 1.5 mm long; lemma tips
awnless, obtuse to notched; inflorescence of 2-8
distant spikes; spikelets densely packed and

overlapping D. eleusine

Ligule membranous, exceeding 2 mm; lemma tips
acute or shortly mucronate; inflorescence an open
panicle; spikelets less dense and usually not

overlapping 2

2(1). Plants reed-like, exceeding 1500 mm in height;
panicle 230-400 mm long; spikelets overlapping;
primary branches slender, 120-200 mm long;
lemma awned; awn to 1 .2 mm long; from Botswana

and Namibia D. gigantea

Plants not reed-like, to 1600 mm tall; panicle usually
90-220 mm long; spikelets not overlapping;
primary branches firm, to 60 mm long; lemma

awnless; also growing outside Botswana 3

3(2). Glumes shorter than 2 mm; lemmas shorter than 3
mm; spikelets to 10 mm long; plants to 700 mm tall;
leaf blades not exceeding 150 mm; racemes

spreading horizontally; from Namibia

D. cuspidata

Glumes 2. 5-3. 2 mm long; lemmas 4. 0-5. 2 mm long;
spikelets 5-15 mm long; plants to 1600 mm tall;
leaf blades to 300 mm long; racemes not spreading

more than 70 degrees; widely distributed

D. fusca

Diplachne cuspidata Launert

Perennial; tufted (culms geni-
culate); 320-650 mm tall. Leaf
blades 20-150 mm long; 2-A mm
wide. Spikelets 7-10 mm long.

Ligule membranous, to 3 mm
long; lemmas 2. 8-3.0 mm long,
tips rounded and minutely awned.

Flowering March to April.

Clayey soils, in water or in
mopaneveld. Infrequent. Biome: Savanna. Possibly also in
Angola. Closely related to D. fusca, which is larger in all
dimensions and has a wider distribution.

Description: Launert 1970 (160:68). Voucher: Smook
5111. PRECIS code 9903450-00200.

Diplachne eleusine Nees

Perennial; rhizomatous and
tufted (culms geniculately
ascending); 520-1270 mm tall.

Leaf blades 120-270 mm long;

2-4 mm wide. Spikelets 4-8 mm
long. Ligule a hairy membrane,

1.5 mm long; spikes 2-8, distant;
spikelets overlapping, 5-10-
flowered; lemma tip obtuse to
minutely notched.

Flowering November to April. Sandy soils, rocky slopes
or in the shade of trees, occasionally on turf soils. Common.
Biome: Savanna and Grassland. ?Endemic. Other

Diplachne species have membranous ligules and acute or
mucronate lemma tips.

Description: Stapf 1898-1900 (591), Chippindall 1955
(121). Illustration: Chippindall 1955 (fig. 91). Voucher:
Smook 4249. PRECIS code 9903450-00300.

Diplachne fusca (L.) Beauv. ex Roem. & Schult.

(=D. malaharica sensu
Adamson, non (L.) Merr.) 4.

Kuilgras, swamp grass.

Perennial; hydrophyte, rhizo-
matous, stoloniferous, and tufted;

220-1550 mm tall. Leaf blades
250-550 mm long; 3-5 mm wide.

Spikelets 6-14 mm long. Ligule a conspicuous membrane,
to 5 mm long; racemes numerous; spikelets not
overlapping, 5-1 2-flowered; lemma tips acute.

Flowering October to May. Sandy soil, almost always
near or in fresh or brackish water to 500 mm deep. Com-
mon. Biome: Fynbos, Savanna, Grassland, Nama-Karoo,
and Succulent Karoo. Old world tropics & subtropics and
in Australia. Pasture (in vleis and brackish soil). This grass
covers nearly all possible variations in habit posible in a
grass. Closely related toD. cuspidata from Namibia, which
is smaller with shorter leaf blades, glumes and lemmas.

Description: Chippindall & Crook 1976 (205), Launert

Fig. 70. PI. 62.

Fig. 70. Diplachne fusca

118

1970 (160:68), Stapf 1898-1900 (591), Chippindall 1955
(119), Clayton et al. 1970-1982 (281). Illustration: Chip-
pindall 1955 (fig. 90), Clayton et al. 1970-1982 (fig. 77).
Voucher: Scheepers 1495, Goldblatt 2820. PRECIS code
9903450-00400.

Diplachne gigantea Launert

Robust perennial; hydrophyte
and rhizomatous; 1500-2700 mm
tall. Leaf blades 300-650 mm
long; 4-5 mm wide. Spikelets
10-14 mm long. Ligule 4-6 mm
long; lemma tip ending in a short
mucro, 0.25-0.50 mm long.

Flowering February to May.

Amongst reeds and waterlilies, on
sandbanks and along rivers. Rare. Biome: Savanna.
Tropical Africa. Distinguished from D. cuspidata and D.
fusca, which are not reed-like and have smaller panicles in
which the spikelets do not overlap.

Description: Clayton et al. 1970-1982 (282). Voucher:
Smith 1387. PRECIS code 9903450-00500.

Photosynthetic pathway. C3 (obviously so in D. pumila,
but but the anatomy of D. calvinensis is equivocal, to say
the least: most mesophyll cells are no more than one cell
distant, and the only seeming exceptions are at the tops of
the adaxial ribs. A candidate for intermediacy); XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 2 species. South and south west Africa.
Xerophytic (D. pumila in blown sand over rocks); in open
habitats; maritime-arenicolous (sometimes), or glycophytic
(usually). Namibia and Cape Province. 2 indigenous
species.

References. 1. Conert. 1966. Senckenb. Biol. 47; 338.

Species treatment by N.P. Barker.

1 (0). Body of lemma densely pubescent below tufts of hairs
across back; leaf blade apices rounded, terminating

in a minute spine D. pumila

Body of lemma glabrous above and below tufts of
hairs across back; leaf apices pungent but not
terminating in a spine D. calviniensis

Dregeochloa Conert

Sometimes included in Rytidosperma , Datithonia sensu
lato.

Perennial; long-stoloniferous (sometimes), orcaespitose
(with short often branched creeping rhizomes). Culms
40-250 mm high; herbaceous; unbranched above (but
usually considerably branched just below the soil surface ).
Leaf blades linear, or ovate-lanceolate to ovate ; to 3 mm
wide ; usually folded; not disarticulating. Ligule a fringe of
hairs (minute).

Inflorescence a single raceme, or paniculate (of 4-12
spikelets, rarely a reduced, contracted panicle)', contracted;
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; not two-ranked; not in distinct Tong-
and-short’ combinations; 10-13 mm long; somewhat com-
pressed laterally; disarticulating above the glumes. Callus
long. Glumes two; more or less equal; about equalling the
spikelets to much exceeding the spikelets; awnless; similar
(lanceolate; scarious, or herbaceous below. G1 narrower).
Incomplete florets distal to the female-fertile florets, merely
underdeveloped, awned; proximal incomplete florets
absent.

Female-fertile florets 3-8. Lemmas similar in texture to
the glumes (membranous); hairy (hairs in tufts, in trans-
verse rows, lobes minutely hairy, with a row of tufts at their
base, and larger marginal tufts beneath); 7-9 nerved;
incised; 2 lobed (the lobes acute or bristle-tipped); awned.
Awns 7; median; from the sinus; geniculate; about as long
as the body of the lemma. Palea present; relatively long;
2-nerved. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
sparsely hairy. Fruit small; hilum short (punctiform); peri-
carp free.

Fig. 7 1 . Dregeochloa pumila

Dregeochloa calviniensis Conert

Perennial; shortly rhizomat-
ous; 150-250 mm tall. Leaf
blades 10-120 mm long; 1.6-2. 3
mm wide. Spikelets 12-15 mm
long. Plant bases somewhat
swollen, covered by old leaf
sheaths; sheath mouth with short,
inconspicuous hairs; leaf blades
glabrous or inconspicuously
pubescent; panicle contracted, 30-50 mm long; spikelets
4—5-flowered, uppermost reduced; glumes 9-15 mm long,
3-5-nerved; lemmas 4. 5-6.0 mm long, including lobes,
with 3 tufts of white hairs on either side of central nerve,
glabrous above and below these tufts; lemma lobes
acuminate, apically produced into a short bristle; central
awn 8-10 mm long, geniculate.

Flowering October. Limestone outcrops. Locally com-
mon. Biome: Nama-Karoo. Endemic. A relatively unknown
species from a poorly collected area.

Description: Conert 1966 (335). Illustration: Conert
1966 (335). Voucher: Acocks 19040. PRECIS code
9902045-00100.

Dregeochloa pumila (Nees) Conert

Fig. 71. PI. 63.

(= Danthonia pumila Nees) 1.

Perennial; shortly rhizomat-
ous; 40-70 mm tall. Leaf blades
10-25 mm long; 1.6-3. 5 mm
wide. Plant base covered in broad
scales; leaf sheath mouth
pubesent; leaf blades succulent in
appearance, minutely but densely
pubescent, apex rounded, spiny-apiculate; inflorescence a
raceme (occasionally a panicle); spikelets 6-10-flowered;
glumes 9-13 mm long, 5(-7)-nerved; lemma 3. 0-3. 5 mm
long, including small, truncate lobes, with a row of hairs
across the back below the awn, with a large tuft of hairs on
each margin, projecting at an angle away from the margin
and with two smaller tufts on each side of central nerve;
central awn 4-7 mm long.

Flowering August to January (but also later). Rocky
areas, in crevices or loose sand. Infrequent. Biome: Succu-
lent Karoo and Desert. Endemic. Restricted to coastal belt,
to 15 km inland, in areas subject to sea mists.

Description: Conert 1966 (335-343), Chippindall 1955
(245). Illustration: Conert 1966 (335-343), Chippindall
1955 (fig. 217). Voucher: Ellis 5076. PRECIS code
9902045-00200.

119

Echinochloa P. Beauv.

Ornithospermum Dumoulin, Tema Adans.

Annual, or perennial; caespitose to decumbent (or
floating). Culms 400-3600 mm high; herbaceous; branched
above, or unbranched above. Leaf blades broad ; flat. Ligule
when present a fringe of hairs.

Inflorescence of spike -like main branches (spike lets
often hispid)\ with axes ending in spike lets', espatheate.
Spikelet-bearing axes persistent.

Spikelets paired or clustered; 2.3-7 mm long; probably
best interpreted as adaxial — i.e., in relation to the reduced,
spikelet-bearing branch; compressed dorsiventrally; falling
with the glumes. Glumes two; very unequal; awned, or
awnless; very dissimilar ( G1 usually much shorter, ovate,
often mucronate. G2 strongly concave, acute, cuspidate or
awned). Proximal incomplete florets 1\ sterile lemma
awned or acuminate', paleate, palea fully developed, or
reduced (e.g. E. kimberleyensis)', male (rarely), or sterile.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes', smooth; becoming indurated, or not becoming
indurated (subcoriaceous to crustaceous); hairless (shiny);
having the margins tucked in onto the palea; with a clear
germination flap; 5 nerved; entire; awnless (obtuse to
apiculate). Palea present (the tip reflexed); relatively long.
Lodicules fleshy; glabrous. Stamens 3. Ovary glabrous.
Fruit small; hilum short; embryo large.

Photosynthetic pathway. C4; NADP-ME (3 species);
XyMS- PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. 30-40
species. In warm regions. Hydrophytic, helophytic, and
mesophytic; mostly in open habitats (in water and moist or
marshy places, also in disturbed ground and weedy places);

Fig. 72. Echinochloa crus-galli

glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 10 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

1(0). Ligule absent in lower and upper leaves 2

Ligule a fringe of hairs, at least in the lower leaves

5

2(1). Racemes distinctly compound with short secondary
branchlets, the inflorescence untidily ovate;

spikelets often with a short curved awn

E. crus-pavonis

Racemes not or inconspicuously compound,
inflorescence elongate; spikelets rarely with a short

curved awn 3

3(2). Plants perennial, rhizomatous E. haploclada

Plants annual, not rhizomatous 4

4(3). Racemes untidily 2 to several rowed, 20-100 mm long
and usually with secondary branchlets at the base;
spikelets 3-4(-7) mm long, awnless or rarely with

awns up to 5 mm long E. crus-galli

Racemes neatly 4-rowed, usually 10-25 mm long,
with no secondary branchlets at the base, awnless

E. colona

5(1). Plants always annual, racemes neatly 4-rowed ....

E. ugandensis

Plants mainly perennial, racemes not neatly 4-rowed

6

6(5). Spikelets seldom over 2.5 mm long, commonly round
to elliptic, often awned, the awns 5(— 1 5) mm long

E. haploclada

Spikelets seldom under 3 mm long, commonly elliptic
to elongate, awns absent or when present 5-25 mm

long 7

7(6). Spikelets awnless (rarely with a subulate point up to

3 mm long) 8

Spikelets with awns usually longer than 5 mm ... 9
8(7). Culms 500-900 mm tall; leaves 180-220 mm long;

inflorescence (80—) 1 20(— 1 80) mm long

E. holubii

Culms 1000-4000 mm tall; leaves 200-600 mm long;

inflorescence ( 1 50— )200( — 400) mm long

E. pyramidalis

9(7). Inflorescence open, the branches clearly secund;

spikelets narrowly ovate, 4-6 mm long

E. stagnina

Inflorescence dense, the branches not clearly secund;
spikelets narrowly elliptic, 3. 0-3. 5(-4. 0) mm long
E. jubata

Echinochloa colona (L.) Link

Jungle rice.

Annual; hydrophyte, stolon-
iferous, and tufted; 100- 1000 mm
tall. Leaf blades 50-300 mm
long; 2-8 mm wide. Spikelets
1.5-3 mm long; 1.0-1. 5 mm
wide. Ligule absent; inflores-
cence 10-150 mm long; racemes
neatly 4-rowed, 10-25 mm long; spikelet pubescent; lower
floret male or sterile; lower lemma not awned (tip may be
up to 1 mm long).

Flowering January to April. Muddy or swampy places.
Common. Biome: Savanna, Grassland, and Nama-Karoo.
Worldwide tropics and subtropics. Food and drink (cereal).
This species may hybridize with E. crus-galli and E.
haploclada.

Description: Chippindall 1955 (361), Clayton et al.

120

1970-1982 (557). Illustration: Clayton et al. 1970-1982
(fig. 134). Voucher: Smook 4398. PRECIS code

9901120-00100.

Echinochloa crus-galli (L.) Beauv.

( =E . subverticillata Pilg.) 2.

Barnyard millet.

Annual; hydrophyte, stolon-
iferous, and tufted; 250- 1 000 mm
tall. Leaf blades 70-350 mm
long; 4-20 mm wide. Spikelets
3 — 4(— 7) mm long; 1-2 mm wide. Ligule absent; inflores-
cence 60-220 mm long; racemes untidily 2 to several-
rowed, 20-100 mm long and usually with secondary
branchlets at base; lower floret sterile; lower lemma usual-
ly awnless, rarely with an awn 5-10 mm long.

Flowering January to April. Swampy areas, wet places
of cultivation. Locally common. Naturalized from
temperate Eurasia. Biome: Fynbos, Savanna, Grassland,
and Nama-Karoo. Worldwide temperate and subtropical
regions. Food and drink (cereal), or weed (polymorphic
especially in rice fields). A very variable species, may be
confused with E. colona, which has neat 4-rowed racemes
and E. crus-pavonis, which has a larger inflorescence and
racemes with many secondary branches.

Description: Chippindall 1955 (362), Clayton et al.
1970-1982 (557). Illustration: Chippindall 1955 (fig. 312).
Voucher: Smook 5871. PRECIS code 9901 120-00200.

Echinochloa crus-pavonis (Kunth) Schult.

Gulf barnyard grass.

Annual, or perennial (rarely);
hydrophyte, stoloniferous, and
tufted; 500-2000 mm tall. Leaf
blades 150-600 mm long; 5-25
mm wide. Spikelets 2.0-3. 5 mm
long; 1.0-1. 5 mm wide. Ligule
absent; inflorescence 100-300
mm long, untidily ovate; racemes 30-150 mm long,
distinctly compound with short secondary branchlets;
spikelets in dense clusters; lower floret male or sterile; low-
er lemma acute or with short curved awn 1— 3(— 7) mm long.

Flowering February to March. Along stream banks and
swamps. Locally common. Biome: Fynbos and Grassland.
Tropics of Africa and America. See note under E. crus-
galli.

Description: Chippindall 1955 (362), Clayton et al.
1970-1982 (556). Illustration: Haefliger and Scholz 1980
Grass Weeds I, Documenta, Ciba-Geigy (p.56). Voucher:
Pole-Evans PRE 34612. PRECIS code 9901120-00300.

Echinochloa haploclada (Stapf) Stapf

Perennial; hydrophyte, rhizo-
matous, and tufted; 300-3000
mm tall. Leaf blades 50-100 mm
long; 3— 1 0(— 20) mm wide. Spike-
lets 1 .5— 2.5(— 3.0) mm long; 1.0-
1.5 mm wide. Ligule absent or a
fringe of hairs; inflorescence 70-
250 mm long; racemes 10-50 mm
long, densely crowded with ap-
pressed spikelets; lower floret male; lower lemma acute or
with awns 5-15 mm long.

Flowering March to April. Stream banks, dry river beds.
Biome: Savanna and Grassland. Northwards to Sudan and
Ethiopia. This species can hybridize with E. colona.

Description: Clayton et al. 1970-1982 (560). Voucher:
Davidse and Ellis 5869. PRECIS code 9901 120-00500.

Echinochloa holubii (Stapf) Stapf

Kalahari water grass.

Perennial; hydrophyte, rhizo-
matous, and tufted; 500-900 mm
tall. Leaf blades 180-220 mm
long; (2— )4(— 8) mm wide. Spike-
lets 2. 5-3. 5 mm long; 1.0-1. 5
mm wide. Ligule a fringe of hairs,
may be absent in upper leaves; in-
florescence (80 — ) 1 20(— 180) mm long, racemes distant,
15-40 mm long; lower floret male or sterile; lower lemma
awnless, tip acute to acuminate, 3 mm long.

Flowering December to April. Swampy areas, pans and
vleis. Locally common. Biome: Savanna, Grassland, Nama-
Karoo and Succulent Karoo. To Zimbabwe. Clayton 1982
(562) regards this species as a synomym of E. pyramidalis,
which extends through tropical Africa. This has not been
adopted for the FSA region, where the two species are dist-
inguishable on size characters.

Description: Chippindall 1955 (361). Illustration: Clay-
ton et al. 1970-1982 (fig. 309). Voucher: Smook 4415.
PRECIS code 9901120-00600.

Echinochloa jubata Stapf

Perennial; hydrophyte, rhizo-
matous, and stoloniferous; 500-
2000 mm tall. Leaf blades 1 00 —

250 mm long; 3-15 mm wide.

Spikelets 3.0-3.5(-4.0) mm long;

1 mm wide. Ligule a fringe of
hairs, may be absent from upper
leaves; inflorescence dense,

80-200 mm long; racemes 20^40
mm long; spikelets narrowly elliptic and closely packed;
lower floret male or sterile; lower lemma with awns 3-25
mm long.

Flowering November to May. Growing in water and
stream sides, often floating in water. Locally common.
Biome: Savanna and Grassland. Northwards to the tropics.
Clayton 1982 (564) records ih&iE. jubata may be a southern
variant of E. stagnina.

Description: Clayton et al. 1970-1982 (563).

Illustration: Chippindall 1955 (fig. 311). Voucher:

Zambatis 1376. PRECIS code 9901120-00650.

Echinochloa pyramidalis (Lam.) Hitchc. & Chase

Limpopo grass.

Perennial; hydrophyte, rhizo-
matous, stoloniferous, and tufted;

1000-4000 mm tall. Leaf blades
80-600 mm long; 5-20 mm wide.

Spikelets 2. 5^4.0 mm long; 1.0-
1.8 mm wide. Culms robust; lig-
ule a fringe of hairs, may be ab-
sent in upper leaves; inflorescence ( 150 — )200( — 400) mm
long; racemes simple or compound, 25-35 mm long; lower
floret male; lower lemma awnless, tip acute to acuminate,
3 mm long.

Flowering December to May. Swamps and riversides,
usually standing in water and may be floating. Locally com-
mon. Biome: Fynbos, Savanna, and Grassland. To tropical
Africa and Madagascar. Domestic use (cereal), or pasture
(natural and cultivated).

Description: Chippindall & Crook 1976 (133), Chippin-
dall 1955 (361), Clayton et al. 1970-1982 (561).
Illustration: Chippindall 1955 (fig. 310). Voucher: Smook
1882. PRECIS code 9901120-00700.

121

Echinochloa stagnina (Retz.) Beauv.

Long-awned water grass, wat-
er grass.

Perennial and annual (rarely);
hydrophyte, rhizomatous, and
stoloniferous; 800-1500 mm tall.

Leaf blades 100-450 mm long;

4-15 mm wide. Spikelets 4—6 mm
long; 1.0-1. 8 mm wide. Ligule a
fringe of hairs, often absent in upper leaves; inflorescence
open, 80-250 mm long; racemes 20-80 mm long, branches
clearly secund; spikelets narrowly ovate, with rigid hairs on
nerves; lower floret male or sterile; lower lemma with
awns ( 1 — )3— 20(— 50) mm long.

Flowering December to May. Growing in water,
streamsides and often floating in water. Locally common.
Biome: Savanna and Grassland. Tropical Africa,

Madagascar, Assan to Indo-China. Pasture.

Description: Chippindall & Crook 1976 (132),Chippin-
dall 1955 (360), Clayton et al. 1970-1982 (562).
Illustration: Chippindall 1955 (fig. 311). Voucher: Jacobsen
2978. PRECIS code 9901 120-00800.

Echinochloa ugandensis Snowden & C.E. Hubb.

Annual; hydrophyte, stolon-
iferous, and tufted; 250-800 mm
tall. Leaf blades 70-200 mm
long; 3-6 mm wide. Spikelets
2.3-3.0 mm long; 1.5 mm wide.

Ligule a fringe of hairs; inflores-
cence 50-200 mm long, linear
with racemes neatly 4-rowed and
up to 30 mm long; spikelets
pubescent; lower floret sterile; lower lemma acute or with
awn up to 6 mm long.

Flowering January. Swampy areas, shallow pools. Lo-
cally common. Biome: Savanna. Up to tropical east Africa.

Description: Clayton et al. 1970-1982 (561). Voucher:
Smook 5337. PRECIS code 9901 120-00900.

Ehrharta Thunb.

Diplax Bennett, Trochera L. Rich.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 60-1500 mm
high; woody and persistent, or herbaceous; branched above,
or unbranched above. Leaf blades linear to linear-
lanceolate; flat, or folded, or rolled; disarticulating from the
sheaths, or not disarticulating. Ligule an unfringed
membrane, or a fringed membrane, or a fringe of hairs.
Plants with hermaphrodite florets.

Inflorescence a single raceme, or paniculate (then
narrow, with slender branches)', espatheate (though in two
species the mature inflorescence base is enclosed in the
uppermost leaf sheath). Spikelet-bearing axes persistent.

Spikelets solitary; not in distinct ‘long-and-short’ com-
binations; 2-17 mm long; compressed laterally, or not
noticeably compressed; disarticulating above the glumes.
Hairy callus absent. Glumes two; very unequal, or more or
less equal; decidedly shorter than the adjacent lemmas, or
long relative to the adjacent lemmas; awnless; similar
(membranous). Lower glume 5 nerved. Proximal incom-
plete florets florets 2 (very variable in form and structure);
epaleate; sterile; lemmas awned (abruptly from the back, or
the lemma tapering into the awn), or awnless; less firm than
the female-fertile lemmas, or similar in texture to the
female-fertile lemmas.

Female-fertile florets I. Lemmas entire; usually
awnless, or mucronate (occasionally); 5-7 nerved. Palea
present (keeled); relatively long (narrow); thinner than the

lemma ; 1 -nerved to with several nerves, or nerveless
(rarely). Lodicules 2; membranous; ciliate, or glabrous.
Stamens 3, or 4, or 6. Ovary glabrous. Fruit small; hilum
long-linear; embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells (sometimes, in southern African species), or without
arm cells; without fusoids. Midrib with one bundle only.

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Oryzodae; Ehrharteae. 27

122

species. Southern and tropical Africa, Mascarene Is., New
Zealand. Helophytic (most annuals), or mesophytic; in
shade (£. erecta), or in open habitats; maritime-arenicolous
(. E . villosa ), or glycophytic. Namibia, Transvaal, Orange
Free State, Natal, Lesotho, and Cape Province. 23 indige-
nous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Gibbs
Russell. 1984. Bothalia 15: 145 & 149. 3. Gibbs Russell &
Ellis. 1987. Bothalia 17: 5 1-65. .4. Gibbs Russell. 1987a.
Bothalia 17:73-67. 5. Gibbs Russell. 1987b. Bothalia
17:191-194. 6. Gibbs Russell & Ellis. 1988. Bothalia
18:165-171. 7. Gibbs Russell & Ellis. 1989. Bothalia 19:
189-207.

Species treatment by G.E. Gibbs Russell.

1(0). First sterile lemma thin, triangular, with raised nerves,
less than half the length of the second sterile
lemma; second sterile lemma and female-fertile

lemma similar, with canoe-shaped tips 2

First sterile lemma similar in texture to second sterile
lemma, with similar nervation, half as long as to
equalling the second sterile lemma; female-fertile
lemma differing from second sterile lemma ... 8
2(1). Glumes about half the length of the longest lemma

3

Glumes longer than half the length of the longest

lemma 5

3(2). Plants delicate, herbaceous, less than 250 mm tall;
inflorescences of 1-4 spikelets, barely overtopping

the leaves; spikelets 4. 5-5.0 mm long

E. rupestris subsp. dodii

Plants not delicate, herbaceous to suffrutescent,
200-450 mm tall; inflorescences of 4—9 spikelets,
considerably overtopping leaves; spikelets 4. 5-6. 3

mm long 4

4(3). Leaf blades rolled and appearing setaceous, held
erect, or flat and held nearly spreading, tips not
hooded; spikelets to 2 mm across, outline oblong

to linear E. rupestris subsp. tricostata

Leaf blades folded, somewhat thickened, held
ascending, tips hooded; spikelets to 2.5 mm across,

outline oblong to nearly square

E. rupestris subsp. rupestris

5(2). Plants erect, 250-400 mm tall; inflorescences of 5-15
spikelets; glumes appressed to florets at maturity

6

Plants sprawling, or if erect then less than 250 mm
tall; inflorescences of \-b spikelets; glumes gaping

widely at maturity 7

6(5). Leaf blades tightly rolled, appearing setaceous, rigid,
erect or curved slightly outward from the middle,

texture smooth; spikelets 5. 5-6. 5 mm long

E. setacea subsp. setacea

Leaf blades flat, to 6 mm across at base, rolled near
tip, held ascending, texture scabrous; spikelets
(6.5-)7.0-8.0 mm long . E. setacea subsp. scabra
7(5). Plants sprawling or trailing; culms herbaceous, lowest
nodes bearing leaves with blades; spikelets 4. 5-6. 5
mm long; glumes usually a little longer than

lemmas E. setacea subsp. uniflora

Plants erect; culms suffrutescent below, lowest nodes
usually leafless; spikelets 4-5 mm long; glumes

slightly shorter than lemmas

E. setacea subsp. disticha

8(1). Culms with the lowest node swollen and hard

(‘bulblike’) 9

Culms not swollen and hard (‘bulblike’) at the lowest

node 13

9(8). Sterile lemmas with a fringe of long hairs on keel;
‘bulbs’ fusiform, ivory coloured; basal sheaths

dark purple E. eburnea

Sterile lemmas not fringed with long hairs; ‘bulbs’
spherical or cylindrical, white or orange; basal
sheaths not dark purple 10

10(9). First sterile lemma broadest at middle, margins
inrolled at basal third; glumes often less than half
the length of the lemmas; first sterile lemma with

strong ribs, at least on the basal half 11

First sterile lemma with margins straight from base
to tip; glumes 1/2-2/3 length of the lemmas; first
sterile lemma longitudinally nerved or weakly

corrugated 12

11(10). Leaf blades flat, marginal vein pale, thickened,
usually undulate; ‘bulbs’ taller than wide, light
orange, polished, crowded; spikelets 8-12 mm

long E. capensis

Leaf blades rolled or sometimes flat, marginal vein
usually not prominent or undulate; ‘bulbs’
chalky white, not polished, well separated on a

thin rhizome; spikelets 7-10 mm long

E. bulbosa

12(10). ‘Bulbs’ taller than wide, dark orange, polished,
crowded; spikelets 8-10 mm long .. E. ottonis
‘Bulbs’ spherical, pale orange, somewhat shining,
obscurely punctate; spikelets 10.0-1 1.5 mm long

E. longifolia

13(8). Sterile lemmas lacking long hairs on sides, keel or
margins, glabrous to strongly scabrous (but

sometimes bearded at base) 14

Sterile lemmas with long hairs on sides, keel or

margins 28

14(13). Sterile lemmas with tip drawn out into an awn at
least 1/3 as long as body of lemma, usually
equalling or longer than body of lemma ... 15
Sterile lemmas not awned (sometimes mucronate,
but if mucro is as much as 1/3 as long as lemma,
then lemma with long hairs on sides or keel) . .

18

15(14). Plants annual; basal sheaths thin and loose, not
flabellate; first sterile lemma more than 2/3
length of the second; fertile floret shorter than

both sterile lemmas 16

Plants tufted rhizomatous perennials; basal sheaths
hard, flabellate; first sterile lemma 1/2-2/3 length
of the second; fertile floret longer than body of

first sterile lemma 17

16(15). Spikelets (7— )10— 25 mm long (including awns);

sterile lemmas bearded at base; sides smooth or
nerved, or with 6-12 small transverse

corrugations; stamens 6 E. longiflora

Spikelets 6—1 1 (—14) mm long (including awns);
sterile lemmas not bearded at base, sides with
4-8 strong transverse corrugations; stamens 3 .

E. triandra

17(15). Leaf blades expanded, 4—10 mm across, lanceolate;

basal sheaths persistent, hard, reddish brown;

awns 2-16 mm long E. dura

Leaf blades reduced, setaceous, to 1 mm across;
basal sheaths eventually deciduous,
membranous, light brown or whitish; awns 1 3-25

mm long E. microlaena

18(14). Sterile lemmas shorter than 5.5 mm (rarely to 7 mm,
but then with glumes less than 3/4 the length of

the lemmas) 19

Sterile lemmas longer than 5.5 mm, glumes longer

than 3/4 the length of the lemmas 24

19(18). Glumes 1/2-3/4 the length of the lemmas; sterile
lemmas with longitudinal nerves or transverse

corrugations 20

Glumes nearly as long as to longer than the lemmas;

sterile lemmas smooth, shining 23

20(19). Spikelets 2-3 mm long; glumes 2/3— 3/4 the length
of the lemma; second sterile lemma with an

earlike appendage at base E. delicatula

Spikelets 3-6(-7) mm long; glumes 1/2-2/3 the
length of the lemmas; second sterile lemma not

appendaged at the base ,. . 21

21(20). Spikelets 5-7 mm long; sterile lemmas gradually

tapering to an acute tip

E. erecta var. abysinnica

123

Spikelets less than 5 mm long; sterile lemma tips

tapering or abruptly rounded 22

22(21). Spikelets (4.0-)4.2-5.0 mm long; sterile lemmas
tapering gradually to a subacute tip; second
sterile lemma often bearded at base, sides
longitudinally nerved or only shallowly

transversely corrugated

E. erecta var. natalensis

Spikelets 3. 0^4. 2 mm long; sterile lemmas abruptly
rounded at tip; second sterile lemma not bearded
at base, sides usually deeply transversely

corrugated, especially on the distal half

E. erecta var. erecta

23(19). Glumes 0.5-2. 0 mm longer than lemmas;

mountainsides in the Drakensberg

E. longigluma

Glumes nearly as long as to 0.5 mm longer than
lemmas; dry places in the SW and NW Cape . .

E. melicoides

24(18). Plants robust, strongly suffrutescent; leaf blades
absent or reduced; sterile lemmas usually

mucronate 25

Plants not robust, usually herbaceous; leaf blades
present, expanded; sterile lemmas usually

muticous 26

25(24). Plants very robust, culms to 5 mm across; glumes
usually slightly shorter than sterile lemmas;
inflorescence usually contracted, pedicels erect

to ascending; leaves usually bladeless

E. ramosa subsp. ramosa

Plants robust, culms to 2.5 mm across; glumes
slightly to considerably longer than sterile
lemmas; inflorescence usually open, pedicels
spreading to reflexed; leaves rarely with small

blades E. ramosa subsp. aphylla

26(24). Inflorescence contracted, pedicels and spikelets

erect; glumes subcoriaceous

E. rehmannii subsp. subspicata

Inflorescence open, pedicels spreading to reflexed
and spikelets spreading to nodding; glumes

membranous 27

27(26). Inflorescence of fewer than 20 spikelets; leaf blades
narrower than 4 mm; sterile lemmas glabrous on

sides E. rehmannii subsp. filiformis

Inflorescence of more than 20 spikelets; leaf blades
to 6 mm across; sterile lemmas sometimes shortly
hairy on sides or tips and/or strongly scabrous on

keels E. rehmannii subsp. rehmannii

28(13). Second sterile lemma with ear-like appendage at
base; spikelets usually less than 8.5 mm long

(rarely to 1 1 mm) 29

Second sterile lemma not appendaged at base;

spikelets longer than 8.5 mm 32

29(28). First sterile lemma more than 2/3 the length of the

second 30

First sterile lemma about 1/2 the length of the

second 31

30(29). Sterile lemmas with tips truncate or with mucro
arising abruptly from central nerve; plants
perennial (very rarely annual) .... E. calycina
Sterile lemmas with tips running out gradually to
mucros 1-2 mm long; plants annual . E. pusilla
31(29). Sterile lemmas with tips rounded; second sterile
lemma inflated; spikelets more or less terete,
2. 7-3. 5 mm long . E. brevifolia var. brevifolia
Sterile lemmas with tips aristate, not inflated;
spikelets laterally compressed, 3. 5-4. 5 mm long

E. brevifolia var. cuspidata

32(28). Sterile lemmas with hairs only on keels and
margins; leaf blades hairy, flat; nodes often with
a ring of retrorse hairs; rhizomes short, woody,

knotted E. barbinodis

Sterile lemmas with profuse hairs on sides; leaf
blades glabrous, usually rolled; nodes lacking
retrorse hairs; rhizomes long, no thicker than
culms 33

33(32). Glumes 1/2-3/4 as long as the spikelet, 5-nerved,
upper glume to 8 mm long; spikelets 8-10 mm
long; rhizomes densely covered with hairy

cataphylls, internodes often sub-bulbous

E. thunbergii

Glumes 3/4 as long to about equalling the spikelet,
5-9 nerved, upper glume 8-13 mm long;
spikelets (10-)1 1-18 mm long; rhizomes naked,

slender, not sub-bulbous 34

34(33). Inflorescence exserted from uppermost leaf sheath,
the sheath usually not inflated; upper glume 9-13

mm long; culms to 3 mm across

E. villosa var. villosa

Inflorescence closely subtended or enveloped by
inflated uppermost leaf sheath; upper glume
( 1 0—) 1 3— 1 8 mm long; culms to 5 mm across . .
E. villosa var. maxima

Ehrharta barbinodis Nees ex Trin.

Shrub or dwarf shrub; tufted;

300-900 mm tall. Leaf blades
10-100 mm long; to 4 mm wide.

Spikelets 10-13 mm long; about
3 mm wide. Culms several,
branched, woody, nodes retrorse-
ly hairy; leaf blades short; sterile
lemmas similar, smooth, with
hairs on keel and margins.

Flowering July to October. Rocky hillsides, often
growing through bushes. Infrequent to locally common.
Biome: Fynbos and Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (679), Chippindall 1955
(44). Illustration: Chippindall 1955 (fig. 4(11) & 14).
Voucher: Acocks 16439. PRECIS code 9901600-00100.

Ehrharta brevifolia Schrad. var. brevifolia

Erect annual; 180-300 mm
tall. Leaf blades 25-90 mm long;

3-5 mm wide (flat or folded).

Spikelets 2. 7-3. 3 mm long.

Sterile lemmas similar in texture,
sides long-hairy, 1st about half
the length of 2nd, the 2nd with a
pair of ear-like appendages at
base, tip rounded, sides inflated,
the spikelets therefore nearly terete.

Flowering August to October. Sandy soil of coastal
Fynbos and Strandveld. Infrequent. Biome: Fynbos and
Succulent Karoo. Endemic. A few specimens have sterile
lemmas with apiculate tips and somewhat flattened sides,
and are apparently intermediate between the two varieties.
In both varieties the mature sterile lemmas sometimes have
dark blotches.

Description: Stapf 1898-1900 (673), Chippindall 1955
(42). Illustration: Chippindall 1955 (fig. 4(6)). Voucher:
Smith 3031. PRECIS code 9901600-00200.

Ehrharta brevifolia Schrad. var. cuspidata Nees

Erect annual; 200-500 mm
tall. Leaf blades 25-100 mm
long; 3-5 mm wide (flat or
folded). Spikelets (3.2-)3.5-4.5
mm long (including arista).

Sterile lemmas similar in texture,
sides long-hairy, 1st about half
the length of 2nd, the 2nd with a
pair of ear-like appendages at
base, tip aristate, sides laterally compressed, the spikelets
therefore flattened.

Flowering August to November (only rarely in
November). Sandy soil, hillsides and Strandveld. Infre-

124

quent. Biome: Fynbos and Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (674), Chippindall 1955
(42). Voucher: Goldblatt 2279. PRECIS code 9901600-
00300.

Ehrharta bulbosa J.E. Sm.

Bulb or corm; long-rhizomat-
ous and tufted; to 700 mm tall.

Leaf blades 60-350 mm long; to
8 mm wide (flat or rolled,
ascending). Spikelets 7-10 mm
long. Lowest culm node bulbous,
spherical, whitish; sterile lemmas
similar in texture, transversely
corrugated, the 2nd broadest
at middle, inrolled below.

Flowering October to November. Hillsides and flats, alt.
50-250 m. Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (666), Chippindall 1955
(38). Illustration: Chippindall 1955 (fig. 4(24)). Voucher:
Manson 205. PRECIS code 9901600-00400.

Ehrharta calycina J.E. Sm.

Fig. 74. PI. 65.

Very variable perennial, or an-
nual (possibly); often rhizomat-
ous; 300-700(-1800) mm tall.

Leaf blades filiform or to 7 mm
wide (flat or rolled). Spikelets
4. 0-8. 5 mm long. Sterile lemmas
similar in texture, sides long-
hairy, 1st more than 2/3 length of
2nd, the 2nd with tip acute,
truncate or commonly with a mucro arising abruptly from
the central nerve, and a pair of ear-like appendages at base.

Flowering July to June (but usually in spring). Many
habitats and soil types. Common. Biome: Fynbos, Savanna,
and Succulent Karoo. Endemic, but introduced to Australia
as a pasture grass and naturalized there, also in California.
Pasture (local strains have been tested for forage value; this
species is one of the few winter-rainfall grasses even
potentially valuable for grazing). This widespread and
variable entity is a species complex showing polyploidy and
probably aneuploidy. Many ecotypes and regional variants
can be recognized. Some have been formally described, e.g.
var. angustifolia and var. versicolor, but their status
requires a full biosystematic study and they are therefore
not treated here.

Description: Stapf 1898-1900 (674), Chippindall 1955
(42). Illustration: Chippindall 1955 (fig. 4(14) &12).
Voucher: Anderson 46. PRECIS code 9901600-00600.

Ehrharta capensis Thunb.

Fig. 73. PI. 66.

Bulb or corm; long- rhizomat-
ous and tufted; 400-1000 mm
tall. Leaf blades 50-220 mm
long; to 10 mm wide (flat,
spreading, with thickened undu-
late marginal vein). Spikelets
8-12 mm long. Lowest culm node
bulbous, cylindrical, orange,
shining; sterile lemmas similar in
texture, transversely corrugated, the 2nd broadest at middle,
inrolled below.

Flowering September to November. Mountains and
hillsides, on a variety of soils. Infrequent to locally com-
mon. Biome: Fynbos and Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (667), Chippindall 1955
(38). Illustration: Chippindall 1955 (fig. 4(25) & 6).
Voucher: Adamson 3040. PRECIS code 9901600—00700.

Fig. 74. Ehrharta calycina

125

Ehrharta delicatula (Nees) Stapf

Leafy annual; 45-250 mm tall.

Leaf blades 20-160 mm long;

1-1 1 mm wide (flat, thin). Spike-
lets 2-3 mm long. Glumes
1/2-3/4 length of lemmas; sterile
lemmas similar, sides not long-
hairy, with 2-3 corrugations, the
2nd sterile lemma with a pair of
ear-like appendages at base.

Flowering July to October. In mesic microhabitats in
arid areas: between rocks on outcrops, in shade of shrubs
and in streambeds. Locally common. Biome: Fynbos,
Nama-Karoo, and Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (672), Chippindall 1955
(40). Illustration: Chippindall 1955 (fig. 4(9) & 10).
Voucher: Goldblatt 2463. PRECIS code 9901600-00800.

Ehrharta dura Nees ex Trin.

Perennial; rhizomatous and
tufted (erect, rarely long-
rhizomatous); to 800 mm tall.

Leaf blades 90-320 mm long;

4-10 mm wide (flat). Spikelets
9-16 mm long (excluding awns);

3 mm wide. Basal sheaths
flattened, reddish brown; sterile
lemmas similar, subglabrous,
with awns 13-25 mm long.

Flowering September to December. Mountain Fynbos in
seasonally moist open habitats, on sandstone or granite-
derived soils, alt. 430-1300 m. Infrequent. Biome: Fynbos.
Endemic.

Description: Stapf 1898-1900 (665), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig. 4(17)). Voucher:
Taylor 4211. PRECIS code 9901600-01000.

Ehrharta eburnea Gibbs Russell

Bulborcorm; tufted; 200-500
mm tall. Leaf blades 40-150 mm
long; to 5 mm wide. Spikelets
9-13 mm long; 3^4 mm wide.

Lowest 1 or 2 culm nodes
‘bulbous’, fusiform, whitish,
smooth; basal sheaths purple;
sterile lemmas similar, smooth,
with hairs on keels and margins.

Flowering September to November. Mountainsides,
often in Rhenosterbosveld, alt. 1000-1400 m. Rare. Biome:
Fynbos. Endemic.

Description: Gibbs Russell (1984) 145. Illustration:
Gibbs Russell (1984) Fig. 9. Voucher: Acocks 15129.
PRECIS code 9901600-01050.

to India, and is also naturalized in Europe, Australia and
North America. Weed. E. erecta is the most widespread of
all ehrhartas and its spikelet size and basal hairiness
gradually increase northwards from the southwestern Cape.
This clinal variability has traditionally been been treated as
three varieties.

Description: Stapf 1898-1900 (671), Chippindall 1955
(40). Illustration: Chippindall 1955 (fig. 4(19) & 9).
Voucher: Adamson 760. PRECIS code 9901600-01100.

Ehrharta erecta Lam. var. natalensis Stapf

Perennial; loosely tufted (or
rambling); to 900 mm tall. Leaf
blades to 270 mm long; to 16 mm
wide. Spikelets 4. 2-5.0 mm long.

Glumes 1/2-3/4 length of
lemmas; sterile lemmas similar in
texture, sides not long-hairy,
longitudinally nerved or only
shallowly corrugated, tapering
gradually to subacute tips, the upper sterile lemma often
bearded at base.

Flowering October to April (occasionally to June).
Shady moist places, especially forest margins. Locally com-
mon. Biome: Savanna and Forest. Endemic. The third
variety, E. erecta var. abyssinica (Hochst.) Pilg., is only
doubtfully present in the Transvaal, but is the only variety
that occurs in tropical Africa. It is distinguished by much
larger spikelets, 5-7 mm long. Variety natalensis is
intermediate between the other two in spikelet size and
bearding on the upper sterile lemma.

Description: Stapf 1898-1900 (671), Chippindall 1955
(40). Voucher: Cleghorn 3124. PRECIS code

9901600-01200.

Ehrharta longiflora J.E. Sm.

PI. 67.

Leafy annual; 150-600 mm
tall. Leaf blades to 200 mm long;

5- 15 mm wide (flat, collar often
dark). Spikelets 10-25 mm long
(including awns). Sterile lemmas
similar, sides subglabrous, with

6- 12 small corrugations, tips
long-awned, bases bearded.

Flowering July to November.

Hillslopes, in the shade of rocks and shrubs, wet places, and
often in disturbed areas such as roadsides, gardens. Locally
common. Biome: Fynbos and Succulent Karoo. Endemic.
Some specimens from Namaqualand have sterile lemmas
with more deeply corrugated sides, and tend toward E.
triandra .

Description: Stapf 1898-1900 (664), Chippindall 1955
(38). Illustration: Chippindall 1955 (fig. 4(10)). Voucher:
Crook 1028. PRECIS code 9901600-01400.

Ehrharta erecta Lam. var. erecta

Perennial; loosely tufted;

200-600 mm tall. Leaf blades
30-150 mm long; 3-12 mm wide
(soft). Spikelets 3. 0^1. 2 mm
long. Glumes 1/2-3/4 length of
lemmas; sterile lemmas similar in
texture, sides not long-hairy,
usually deeply corrugated
especially on upper half, tips
abruptly rounded, bases not bearded.

Flowering throughout the year (commonly October to
January). Shady moist places, often at forest margins. Lo-
cally common. Biome: Fynbos, Savanna, and Forest. This
variety is endemic; the species extends through east Africa

Ehrharta longifolia Schrad.

Bulb or corm; rhizomatous
and tufted; to 1200 mm tall. Leaf
blades 70-200 mm long; to 5 mm
wide (rolled, erect). Spikelets
10-12 mm long. Lowest culm
node bulbous, spherical, whitish,
dull; sterile lemmas similar,
longitudinally veined, margins
straight from base to tip.

Flowering November and December. Mountainsides,
alt. 100-1800 m. Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (667), Chippindall 1955
(38). Illustration: Chippindall 1955 (fig. 4(16)). Voucher:
Liebenberg 4047. PRECIS code 9901600-01500.

126

Ehrharta longigluma C.E. Hubb.

Perennial; rhizomatous (rhi-
zomes long and slender), or tufted
(erect); 300-600 mm tall. Leaf
blades 25-120 mm long; 2-4 mm
wide (flat). Spikelets 4.0-7. 5 mm
long. Glumes 0. 5-2.0 mm longer
than lemmas; sterile lemmas
similar, glabrous, unawned, the
2nd with a pair of ear-like
appendages at base.

Flowering November to March. Mountain grassland,
2300-3300 m, often in peaty soil. Infrequent. Biome: Afro-
montane. Endemic. This is our only species of Ehrharta
whose range does not include the southwestern or
northwestern Cape.

Description: Chippindall 1955 (41). Illustration: Chip-
pindall 1955 (fig. 4(1) & 11). Voucher: Killick 1478.
PRECIS code 9901600-01600.

Ehrharta melicoides Thunb.

Haasgras.

Perennial; rhizomatous (rhi-
zomes stout), or tufted (densely);

300-700 mm tall. Leaf blades
50-250 mm long; to 4 mm wide
(tightly rolled or sometimes flat).

Spikelets 3. 5^L0 mm long. Culm
bases tightly clad by thick old
leaf sheaths and sometimes appearing bulbous, but the
culms themselves not swollen; glumes about as long as the
lemmas; sterile lemmas similar, glabrous, unawned, the 2nd
with a pair of ear-like appendages at base.

Flowering August to November. Mountainsides, in
Rhenosterveld and in overgrazed grassland, often in rocky
places among dolerite or shale. Infrequent. Biome: Fynbos
and Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (673), Chippindall 1955
(41). Illustration: Chippindall 1955 (fig. 4(15)). Voucher:
Acocks 17303. PRECIS code 9901600-01700.

Ehrharta microlaena Nees ex Trin.

Perennial; tufted (erect); to
1 100 mm tall. Leaf blades 70-150
mm long; to 1 mm wide
(setaceous). Spikelets 13-15 mm
long (excluding awns); to 2.5 mm
wide. Basal sheaths pale; sterile
lemmas similar, subglabrous with
awns 2-14 mm long.

Flowering December to Feb-
ruary. Mountain Fynbos at streamsides and in damp peaty
places, alt. 400-1330 m. Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (665), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig. 4(18)). Voucher:
Esterhuysen 28427. PRECIS code 9901600- 01800.

Ehrharta ottonis Kunth ex Nees

Bulb or corm; rhizomatous
and tufted; to 1200 mm tall. Leaf
blades to 600 mm long; to 5 mm
wide (rolled, erect). Spikelets
8-10 mm long. Lowest culm node
bulbous, cylindrical, orange,
shining; sterile lemmas similar,
longitudinally veined, margins
straight from base to tip.

Flowering September to November. Hills and
mountains, and on flats in disturbed places, alt. 50-800 m.

Infrequent to locally common. Biome: Fynbos and Succu-
lent Karoo. Endemic.

Description: Stapf 1898-1900 (677), Chippindall 1955
(39). Illustration: Chippindall 1955 (fig. 4(23)). Voucher:
Lamb 111. PRECIS code 990 1 600-0 1 900.

Ehrharta pusilla Nees ex Trin.

Sprawling annual; 50-350 mm
tall. Leaf blades 15-110 mm
long; 2-7 mm wide (flat or
folded, sheaths often inflated).

Spikelets (5.6-)6.5-8.5 mm long.

Sterile lemmas similar, sides
long-hairy, tips gradually running
out into an arista 1-2 mm long,
the 1st sterile lemma more than
2/3 length of 2nd, the 2nd with a pair of ear-like appendages
at base.

Flowering July to October. Sandy soil, usually in dry
streambeds. Locally common. Biome: Nama-Karoo and
Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (674), Chippindall 1955
(43). Illustration: Chippindall 1955 (fig. 4(8) & 13).
Voucher: Goldblatt 5678. PRECIS code 9901600-02000.

Ehrharta ramosa (Thunb.) Thunb. subsp. aphvlla

(Schrad.) Gibbs Russell

(=£. aphylla Schrad.) 1.

Shrub or dwarf shrub, or per-
ennial (culms woody); rhizomat-
ous (rhizomes woody, branched);

300-800 mm tall. Leaf blades to
30 mm long; to 1 mm wide (often
absent). Spikelets (5.5— )6.0— 7.5
(-9.0) mm long. Culms to 2.5 mm across; leaves usually
bladeless, but reduced blades sometimes present; panicle
usually open, pedicels spreading to reflexed; glumes
usually somewhat longer than lemmas; sterile lemmas
similar, sides hairless, tips usually mucronate, the 2nd
sterile lemma with a pair of ear-like appendages at base.

Flowering September to January. Mountain Fynbos on
TMS-derived soils, between rocks, often in dry micro-
habitats. Locally common. Biome: Fynbos. Endemic.
Intermediates exist to subsp. ramosa and to subspecies of
E. rehmannii.

Description: Stapf 1898-1900 (678), Chippindall 1955
(39). Voucher: Esterhuysen 28110. PRECIS code

9901600-02100.

Ehrharta ramosa (Thunb.) Thunb. subsp. ramosa

Shrub or dwarf shrub, per-
ennial (culms woody); rhizomat-
ous (rhizomes woody, branched);

300-1000 mm tall. Leaf blades
absent. Spikelets (5.5-)6.0-7.5
(-9.0) mm long; 2-3 mm wide.

Culms to 5 mm across; leaves
bladeless; panicle usually con-
tracted, pedicels erect to as-
cending; glumes usually slightly shorter than lemmas;
sterile lemmas similar, sides hairless, tips usually
mucronate, the 2nd sterile lemma with a pair of ear-like
appendages at base.

Flowering October to January. Mountain or grassy
Fynbos, on sandy or stony TMS or lateritic soils, often in
rocky places. Locally common (at high altitudes). Biome:
Fynbos. Endemic. The robust leafless culms are similar to
E. thunbergii and E. villosa , but these species have
profusely hairy sterile lemmas.

Description: Stapf 1898-1900 (677), Chippindall 1955
(39). Illustration: Chippindall 1955 (fig. 8). Voucher:
Taylor 4235. PRECIS code 9901600-02200.

127

Ehrharta rehmannii Stapf subsp. filiformis (Stapf)
Gibbs Russell

Ehrharta rupestris Nees ex Trin. subsp. dodii (Stapf)
Gibbs Russell

( -E . rehmannii Stapf var.
filiformis Stapf) 7.

Perennial (sometimes deli-
cate); tufted (erect or straggling,
often growing in dense masses);

120-800 mm tall. Leaf blades
15-100 mm long; to 4 mm wide
(usually soft and thin). Spike-
lets 4.0-6.5(-8.0) mm long. Inflorescence an open raceme
with 1— 1 5(— 24) spikelets, pedicels spreading to reflexed;
glumes about as long as lemmas, membranous; sterile
lemmas similar, sides hairless, tips usually muticous, the
2nd sterile lemma with a pair of ear-like appendages at
base.

Flowering October to February. Sandy (TMS) and
humic soils, at streamsides, moist places and in shade of
rocks. Infrequent. Biome: Fynbos. Endemic. Intermediates
link this subspecies to the other two.

Description: Stapf 1898-1900 (677), Chippindall 1955
(39). Voucher: Acocks 22484. PRECIS code 9901600-
02300.

Ehrharta rehmannii Stapf subsp. rehmannii

Perennial; tufted (erect);

300-1000 mm tall. Leaf blades
60-300 mm long; to 6 mm wide.

Spikelets (5— )6— 8 mm long; about
2 mm wide. Inflorescence an
open raceme or verticillate
panicle, with more than 20
spikelets; glumes about as long as
lemmas, membranous; sterile
lemmas similar, scabrous to shortly hairy, tips usually
muticous, the 2nd sterile lemma with a pair of ear-like
appendages at base.

Flowering August to December. Mountain slopes, on
streambanks and rocky places, sometimes under trees. In-
frequent. Endemic. A particularly tall, long-leaved form
with thick but soft culms and numerous short spikelets
(5. 5-6.0 mm long) occurs in forests and on rocky ground
at George and Knysna.

Description: Stapf 1898-1900 (677), Chippindall 1955
(39). Illustration: Chippindall 1955 (fig. 4(21)). Voucher:
Compton 23076. PRECIS code 9901600-02400.

Ehrharta rehmannii Stapf subsp. subspicata (Stapf)

Gibbs Russell

( -E . subspicata Stapf) 7.

Perennial; tufted (erect);

300-600 mm tall. Leaf blades
30— 1 20(— 1 70) mm long; 4.0-8. 5
mm wide (erect). Spikelets
6. 0-8. 5 mm long. Inflorescence a
narrow raceme of 12-36 erect
appressed spikelets; glumes about
as long as lemmas, subcoriaceous; sterile lemmas similar,
sides hairless, tips usually muticous, the 2nd sterile lemma
with a pair of ear-like appendages at base.

Flowering October to December. Sandy or gravelly soil
in moist places, near sea level. Rare. Endemic.

Description: Stapf 1898-1900 (676), Chippindall 1955
(39). Illustration: Chippindall 1955 (fig. 4(22) & 7).
Voucher: Esterhuysen 33720. PRECIS code 9901600-
02440.

(=£. dodii Stapf) 2.

Delicate perennial; rhizomat-
ous (trailing, rarely erect); less
than 250 mm tall. Leaf blades
rolled, erect. Spikelets 4. 5-5.0
mm long; to 2 mm wide.

Inflorescence a raceme of 1^4
spikelets; glumes 1/3 as long as
lemmas; 1st sterile lemma short, glumelike, 2nd with a
canoe-shaped tip.

Flowering November to January. Wet places on
mountainsides, among rocks and at cliff bases, alt.
660-1660 m. Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (670), Chippindall 1955
(35). Illustration: Chippindall 1955 (fig. 4(2)). Voucher:
Esterhuysen 33084. PRECIS code 9901600-02490.

Ehrharta rupestris Nees ex Trin. subsp. rupestris

Suffrutescent perennial; long
rhizomatous; to 300 mm tall. Leaf
blades 20-30 mm long; 2-4 mm
wide (folded, distichous, tips
hooded). Spikelets 4. 5-6.0 mm
long; to 2.5 mm wide.

Inflorescence a raceme of 4-8
spikelets; spikelets nearly square;
glumes 1/3 as long as lemmas; 1st
sterile lemma short, glumelike, 2nd with a canoe-shaped tip.

Flowering October to January. Mountain slopes among
rocks, alt. 910-1970 m. Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (668), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig. 4(3)). Voucher:
Esterhuysen 21044. PRECIS code 9901600-02500.

Ehrharta rupestris Nees ex Trin. subsp. tricostata

(Stapf) Gibbs Russell

(=E. tricostata Stapf) 2.

Suffrutescent perennial; rhizo-
matous; 200—450 mm tall. Leaf
blades to 100 mm long; to 2.5 mm
wide (usually rolled, setaceous
(rarely flat)). Spikelets 4. 6-6. 3
mm long; to 2 mm wide. Lowest
leaf sheaths blade-bearing;
inflorescence a raceme of 4-9 spikelets; glumes 1/3 as long
as lemmas; 1st sterile lemma short, glumelike, 2nd with a
canoe-shaped tip.

Flowering October to February. Wet places on mountain
slopes and at base of cliffs, alt. 300-2030 m. Infrequent.
Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (669), Chippindall 1955
(35). Illustration: Chippindall 1955 (fig. 4(4a)). Voucher:
Fourcade 3132. PRECIS code 9901600-02540.

Ehrharta setacea Nees subsp. disticha Gibbs Russell

Delicate but suffrutescent per-
ennial; rhizomatous (cushion-
forming); to 250 mm tall. Leaf
blades to 30 mm long; distichous,
hard, rolled. Spikelets 4-5 mm
long. Inflorescence a raceme of
1-2 spikelets; glumes slightly
shorter than lemmas; 1st sterile
lemma short, glumelike, 2nd with
a canoe-shaped tip.

128

Flowering October to November. Dry rocky places on
mountain slopes, alt. 580-1225 m. Rare. Biome: Fynbos.
Endemic.

Description: Gibbs Russell (1984) Bothalia 15: 151.
Voucher: Esterhuysen 31735. PRECIS code 9901600-
02560.

Ehrharta setacea Nees subsp. scabra (Stapf) Gibbs

Russell

(=£. setacea Nees var.
scabra Stapf) 2.

Suffrutescent perennial; long-
rhizomatous and stoloniferous;

250-600 mm tall. Leaf blades
30-1 10 mm long; to 6 mm wide
(scabrous, flat at base, rolled near
tip). Spikelets (6.5-)7.0-8.0 mm long. Inflorescence a
raceme of 5-15 spikelets; glumes 2/3 — 3/4 as long as
lemmas; 1st sterile lemma short, glumelike, 2nd with a
canoe-shaped tip.

Flowering October to January (sporadically to March).
Mountainsides, among rocks, in seepage areas and in
disturbed places, alt. 350-1212 m. Rare. Biome: Fynbos.
Endemic.

Description: Stapf 1898-1900 (669). Voucher: Haynes
868. PRECIS code 9901600-02580.

Ehrharta setacea Nees subsp. setacea

Suffrutescent perennial; long-
rhizomatous; 250-400 mm tall.

Leaf blades 50— 80(— 110) mm
long; setaceous, hard, smooth.

Spikelets 5. 5-6. 8 mm long.

Lowest leaf sheaths bladeless;
inflorescence a raceme of 5-15
spikelets; glumes 2/3 or more as
long as lemmas; 1st sterile lemma
short, glumelike, 2nd with a canoe-shaped tip.

Flowering September to December (sporadically to
April). Infrequent. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (668), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig. 4(4) & 5).
Voucher: Esterhuysen 28669. PRECIS code 9901600-
02600.

Ehrharta setacea Nees subsp. uniflora (Burch, ex Stapf)

Gibbs Russell

( -E . uniflora Burch, ex
Stapf) 2.

Delicate perennial; rhizomat-
ous; trailing, forming dense
masses. Leaf blades 50-80 mm
long; to 2 mm wide (soft, flat, not
distichous). Spikelets 4. 5-6. 5 mm
long. Inflorescence a raceme of 1—4 spikelets; glumes
usually slightly longer than lemmas; 1st sterile lemma
short, glumelike, 2nd with a canoe-shaped tip.

Flowering September to December (occasionally to
March). Wet places and forest margins, alt. 10-500 m.
Rare. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (670), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig 4(5)). Voucher:
Esterhuysen 34039. PRECIS code 9901600-02620.

Ehrharta thunhergii Gibbs Russell

(=£. gigantea Thunb.) 5;

(=£. virgata Launert) 5.

Perennial; long-rhizomatous
and tufted (erect); to 1500 mm
tall. Leaf blades 30-100 mm
long; to 5 mm wide (rolled, often
deciduous). Spikelets 8-10 mm
long; 2-3 mm wide. Rhizomes
sub-bulbous, with overlapping hairy cataphylls; glumes 6-8
mm long, translucent; sterile lemmas similar, profusely
hairy.

Flowering September to December. Hillslopes in sandy
or gravelly soil, occasionally in coastal sand. Infrequent to
locally common. Biome: Fynbos and Succulent Karoo.
Endemic.

Description: Stapf 1898-1900 (680), Chippindall 1955
(45). Voucher: Acocks 23393. PRECIS code 9901600-
02750.

Ehrharta triandra Nees ex Trin.

Leafy annual; 60-A50 mm tall.

Leafblades30-120mm long; 2-6
mm wide (flat, thin). Spikelets
6 — 1 1 (—14) mm long (including
awns). Sterile lemmas similar,
subglabrous, tips long-awned,
bases not bearded, sides with 4-8
strong corrugations; stamens 3.

Flowering July to October.

Hillsides in shade of rocks and shrubs and in wet places,
sometimes in disturbed places and roadsides. Locally com-
mon. Biome: Succulent Karoo. Endemic.

Description: Stapf 1898-1900 (663), Chippindall 1955
(37). Illustration: Chippindall 1955 (fig. 4(7)). Voucher:
Goldblatt 2819. PRECIS code 9901600-02800.

Ehrharta villosa Schult. f. var. maxima Stapf

Robust perennial; long-rhizo-
matous and tufted (erect); to 1500
mm tall. Leaf blades 15-130 mm
long; to 8 mm wide (rolled, often
deciduous). Spikelets ( 1 0-) 12-18
mm long; to 4 mm wide. Culms
to 5 mm across; rhizomes naked;
inflorescence subtended by
inflated leaf sheath; glumes
13-18 mm long; sterile lemmas similar, profusely hairy.

Flowering September to March (sporadically). Sea
dunes. Rare. Locally dominant. Endemic. Erosion control.

Description: Stapf 1898-1900 (681), Chippindall 1955
(45). Illustration: Chippindall 1955 (fig. 4(12)). Voucher:
Boucher 1689. PRECIS code 9901600-03200.

Ehrharta villosa Schult. f. var. villosa

Robust perennial; long-rhizo-
matous and tufted (erect); to 1 500
mm tall. Leaf blades 30-130 mm
long; to 8 mm wide (rolled, often
deciduous). Spikelets 11-14 mm
long; to 3 mm wide. Culms to 3
mm across; rhizomes naked;
inflorescence exserted from
uppermost leaf sheath; glumes

129

9-13 mm long; sterile lemmas similar, profusely hairy.

Flowering October to December. Seaside dunes, to 1 km
inland. Rare. Locally dominant. Endemic. Erosion control.

Description: Stapf 1898-1900 (681), Chippindall 1955
(45). Illustration: Chippindall 1955 (fig. 4(12)). Voucher:
Cleghorn 3122. PRECIS code 9901600-03300.

Eleusine Gaertn.

Annual, or perennial (the culms flattened); caespitose
(or mat-forming). Culms 100-1500 mm high; herbaceous.
Sheath margins free (the sheaths keeled). Leaf blades
linear, flat, or folded. Ligule a fringed membrane.

Inflorescence of spike-like main branches', open, or
contracted (sometimes forming a capitulum); digitate or
subdigitate (or shortly racemose, but clustered at the top
of the culm)', espatheate. Spikelet-bearing axes persistent.

Spikelets biseriate; 3.5-1 1 mm long; compressed
laterally; disarticulating above the glumes, or not disarticu-
lating (E. coracana ); disarticulating between the florets
(except in E. coracana). Glumes two; very unequal;
markedly shorter than the spikelets; awnless. Upper glume
3-5 nerved. All florets female-fertile, or distal incomplete
florets also present; proximal incomplete florets absent.

Female-fertile florets 3-15 . Lemmas 3 nerved; entire;
awnless to mucronate. Palea present. Lodicules 2; fleshy,
or membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small (0.9-2 mm); ellipsoid to subglobose; hilum short;
pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
NAD-ME (2 species); XyMS+. PCR sheath outlines even.

Fig. 75. Eleusine coracana subsp. africana

PCR sheath extensions absent. PCR cell chloroplasts with
well developed grana; centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 9. Chloridoideae; Chlorideae sensu lato. 9
species. Tropical and subtropical. Mesophytic, or xero-
phytic; in open habitats (savanna, grassland, weedy places).
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. Indigenous
species (1), naturalized species (3?).

References. 1. Phillips. 1971. Kew Bull. 27: 252. 2.
Clayton et al. 1974. FTEA. 3. De Wet et al. 1984. Amer.
J. Bot. 71(4): 550.

Species treatment by M. Koekemoer.

1(0). Spikes 2-8, alternately placed on a short axis,
sometimes in a compact cluster, 15-30 mm long,

8-15 mm wide E. multiflora

Spikes 2-13, digitate or subdigitate, 20-170 mm long,

3-10 mm wide 2

2(1). Spikes 1 — 4, stout, 20-30 mm long, 5-1 1 mm wide .

E. tristachya

Spikes more than four, slender, 20-170 mm long, 3-6

mm wide 3

3(2). Spikelets 4-5 mm long, 2. 0-2. 5 mm wide, 3-9-
flowered, disarticulating above the glumes and
between the florets at maturity; grains oblong . . .

E. indica subsp. indica

Spikelets 5-8 mm long, 3-4 mm wide, 2-6-flowered,
not disarticulating at maturity; grains globose . . .

E. coracana subsp. africana

Eleusine coracana (L.) Gaertn. subsp. africana (K -

O’Byrne) Hilu & De Wet

Fig. 75. PI. 68.

{=E. africana K. -O'Byrne) I;

(=£. indica (L.) Gaertn. subsp.
africana (K. -O'Byrne) S.M.

Phillips) 3.

African finger millet, osgras.

Annual; tufted; 210-620 mm
tall. Leaf blades 220-500 mm
long; 6-10 mm wide. Spikelets 5-8 mm long; 3-4 mm
wide. Spikes 3-13, 60-170 mm long, 4-10 mm wide; spike-
lets 2-6-flowered, not disarticulating at maturity; grains
globose.

Flowering October to May. Ruderal, on many soil types.
Common. Biome: Fynbos, Savanna, and Grassland. Warm
and temperate regions from Africa to Japan and in
Australia. Food and drink (grains ground up for porridge
or left to germinate for beer; eaten as a vegetable in
Indonesia), or domestic use (for plaiting bracelets), or tradi-
tional medicine (internal remedy for leprosy or liver
diseases), or chemicals (hydrocyanic acid), or weed (in
cultivated lands and disturbed places). In east Africa, where
it is cultivated as cereal, five races are distinguished on the
inflorescence morphology. It also has a long historical
record, for it is present in archaeological records of early
African agriculture that date back 5000 years, and was
introduced into India some 3000 years ago.

Description: De Wet et al. 1984, Kennedy-O’Byme
1957 Kew Bull. 12,1 (65-72), Stapf 1898-1900 (645),
Hitchcock & Chase 1950 (481), Chippindall 1955 (129),
Clayton et al. 1970-1982 (260). Illustration: Chippindall
1955 (fig. 103). Voucher: Smook 5427. PRECIS code
9903310-00150.

130

Eleusine indica (L.) Gaertn. subsp. indica

Osgras, goose grass.

Annual; stoloniferous and
tufted; 230^400 mm tall. Leaf
blades 50-350 mm long; 2. 5-6.0
mm wide. Spikelets 4-5 mm
long; 2-3 mm wide. Spikes
slender, 20-120 mm long,

3. 0-5. 5 mm wide; spikelets 4-9-
flowered, disarticulating above the glumes and between the
florets at maturity; grains oblong.

Flowering November to February. Ruderal; on rocky or
turf soils. Infrequent to locally common. Naturalized from
India. Biome: Savanna and Grassland. Worldwide. Food
and drink (grown occasionally as grain), or poisonous
(reported from Australia and elsewhere that young plants
sometimes contain hydrogen cyanide and are responsible
for deaths of calves and sheep), or traditional medicine
(cough remedy), or chemicals (hydrocyanic acid), or weed
(serious worldwide, and a host for numerous fungi,
nematodes and viruses).

Description: Kennedy-O’Byrne 1957 Kew Bull. 12,1
(65), Stapf 1898-1900 (645), Flitchcock & Chase 1950
(481), Chippindall 1955 (129), Clayton et al. 1970-1982
(262). Illustration: Chippindall 1955 (fig. 102), Hitchcock
& Chase 1950 (fig. 1027). Voucher: Stirton 8781. PRECIS
code 9903310-00300.

Eleusine multiflora Rich.

Annual; tufted (culms fairly
slender and ascending); 120-400
mm tall. Leaf blades 60-260 mm
long; 3-6 mm wide. Spikelets
7-1 1 mm long. Spikes 2-8, stout,

15-25 mm long, 8-15 mm wide,
alternating on a short axis.

Flowering February to April.

Disturbed places in bush- or
grassveld. Rare. Biome: Grassland. Tropical east Africa to
Ethiopia. Weed (in cultivated lands).

Description: Chippindall 1955 (129), Clayton et al.
1970-1982 (261). Voucher: Smook 5050. PRECIS code
9903310-00400.

Eleusine tristachya (Lam.) Lam.

Goose grass.

Perennial; tufted; 70-180 mm
tall. Leaf blades 50-180 mm
long; 3-5 mm wide. Spikelets 4-7
mm long. Inflorescence digitate;
spikes 1-4, stout, 10-25 mm
long, 4-11 mm wide; spikelets
2-3-flowered.

Flowering February to April. Disturbed weedy places.
Infrequent. Biome: Fynbos and Grassland. Tropical Africa,
South America, eastern North America.

Description: Hitchcock & Chase 1950 (481), Chippin-
dall 1955 (129). Voucher: Acocks 23824. PRECIS code
9903310-00500.

Elionurus Kunth ex Willd.

Callichloea Steud., Habrurus Hochst.

Annual, or perennial; caespitose. Culms 100-1500 mm
high; herbaceous; unbranched above. The shoots aromatic
(with a bitter taste), or not aromatic . Leaf blades sometimes
flat, or folded (tightly). Ligule a fringed membrane (very
short), or a fringe of hairs. Plants bisexual, with bisexual

spikelets; with hermaphrodite florets. The spikelets of
sexually distinct forms on the same plant ; homomorphic.

Inflorescence a single raceme, or paniculate (of single
‘racemes’ , terminal or sometimes axillary and gathered
into false panicles)-, spatheate; not comprising ‘partial inflo-
rescences’ and foliar organs. Spikelet-bearing axes spike-
like (flexuous)', solitary; with substantial rachides
(flattened); disarticulating at the joints.

131

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only. Female-fertile spikelets compressed
dorsiventrally; falling with the glumes. Glumes two; rela-
tively large; very unequal, or more or less equal; awned (G1
often cuspidate to a bifid tip, the tails several mm long), or
awnless; very dissimilar (the lower tougher, carinate on the
edges, the keels generally glandular or with tufts of hairs;
the upper membranous, lanceolate, not 2-keeled), or similar
(rarely, both subulate). Proximal incomplete florets /;
epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present, or absent;
when present very reduced. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Rottboelliinae. 15 species. Tropical and
subtropical. Mesophytic to xerophytic; in open habitats
(savanna, often on dry soils); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 2 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Leaves basal; lower glume of sessile spikelets with
dense long hairs on back; inflorescence

conspicuously hairy E. muticus

Leaves mostly cauline; lower glume of sessile spikelet
glabrous or with a few sparse hairs on back;
inflorescence with bare sessile spikelets outlined by
short hairs of rachis and pedicels . E. tripsacoides

Elionurus muticus (Spreng.) Kunth

Fig. 76. PI. 69.

(=E. argenteus Nees) 1; ( -E .
glaber Phill.) 1; ( =E . glaber
Phill. var. villosus Phill.) 1; (=£.
pretoriensis Phill.) 1.

Koperdraad, silky grass, suur-
pol, wildebeestegras, wire grass.

Perennial; densely tufted;

200-1200 mm tall. Leaf blades 10-150 mm long; setaceous
or to 1-2 mm wide. Spikelets (sessile) 6-14 mm long (pedi-
cellate somewhat shorter). Leaves basal; lower glume of
sessile spikelets with long dense, silky, white hairs, tip
bidentate.

Flowering September to May. In open grassland,
especially sourveld. Common to dominant. Biome: Savanna
and Grassland. Tropical and subtropical Africa and
America. Indicator (of veld mismanagement), or weed
(ruderal). Other superficially similar species having single
spikelike inflorescences with silky white hairs include
Schizachyrium jeffreysii, Digitaria monodactyla and
Anthephora argentea.

Description: Chippindall 1955 (518), Clayton et al.
1970-1982 (837). Illustration: Chippindall 1955 (pi. 27),
Clayton et al. 1970-1982 (fig. 195). Voucher: De Winter
2551. PRECIS code 9900280-00100.

Elionurus tripsacoides Willd.

Perennial; tufted; 600-1500
mm tall. Leaf blades 200-300
mm long; 1-3 mm wide. Spike-
lets (sessile) 5-8 mm long (pedi-
cellate shorter). Leaves mostly
cauline; lower glume of sess ’e
spikelets glabrous or with fe\
hairs, tip entire or shortly
bidentate.

Open places in savanna. Infrequent. Biome: Savanna.
Tropical Africa and America. Occasionally intergrades with
E. muticus.

Description: Clayton et al. 1970-1982 (838). Voucher:
Ellis 2997. PRECIS code 9900280-00200.

Elymandra Stapf

Annual, or perennial (coarse); caespitose. Culms
500-2500 mm high; herbaceous; branched above. Ligule an
unfringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant (hermaphrodite, male-only and/or sterile );
overtly heteromorphic (only the female-fertile spikelets
awned).

Fig. 77. Elymandra grallata

Inflorescence of spike-like main branches, or paniculate
(of long-exserted ‘racemes’ gathered into a false panicle)',
spatheate; a complex of ‘partial inflorescences’ and inter-
vening foliar organs (the spathes and spatheoles narrow,
subulate or setaceous at tips). Spikelet-bearing axes
‘racemes' (elongated)', paired (two per spatheole, each with
1-6 or more male-only or sterile pairs at the base, then one
or more heterogamous pairs above and a heterogamous
terminal triad); with substantial rac hides', disarticulating at
the joints.

Spikelets in triplets and in pairs (with a terminal
heterogamous triad); consistently in ‘long-and-short’ com-
binations, these pedicellate/sessile (except in the upper part
of the raceme, the lower homogamous pairs are all sessile).
Pedicels free of the rachis. The sessile spikelets hermaphro-
dite (i.e., in the heterogamous combinations). The
pedicellate spikelets male-only, or sterile (rarely). Female-
fertile spikelets not noticeably compressed to compressed
dorsiventrally; falling with the glumes. Glumes two; more
or less equal; awned (G2); very dissimilar (leathery. G1

132

obtuse or truncate; G2 pointed or with a subule, dorsally
rounded and grooved). Lower glume not two-keeled.
Proximal incomplete florets 1; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes; incised; awned. Awns 1; median; from the sinus;
geniculate; much longer than the body of the lemma. Palea
absent. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous.

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Andropogoninae. 4
species. Tropical Africa. Mesophytic; in shade, or in open
habitats (savanna woodland); glycophytic. Namibia and
Botswana. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

Elymandra grallata (Stapf) Clayton

Fig. 77. PI. 70.

Perennial; tufted; 500-2000
mm tall. Leaf blades to 300 mm
long; to 6 mm wide. Spikelets
(sessile) 6.5-12.0 mm long (pedi-
cellate somewhat longer).

Homogamous spikelets olive-
green; sessile spikelets dark
brown, awns 30-50 mm long.

Flowering February to May.

Sandy soil in woodland. Rare and conservation status not
known. Biome: Savanna. Mozambique and north to central
Africa.

Description: Clayton et al. 1970-1982 (823).

Illustration: Clayton et al. 1970-1982 (fig. 189). Voucher:
De Winter & Marais 4721. PRECIS code 9900801-00100.

Elytrigia Desv.

Sometimes included in Agropyron, Elymus.

Perennial; long-rhizomatous (or densely turf-forming).
Culms 200-1500 mm high: herbaceous: unbranched above.
Sheath margins joined (often, on vegetative shoots), or free.
Leaf blades linear; flat, or rolled (convolute). Ligule an
unfringed membrane . The spikelets of sexually distinct
forms on the same plant (with sterile spikelets localised at
the tip of the rachis), or all alike in sexuality.

Inflorescence a single spike ( erect or drooping, linear)-,
espatheate. Spikelet-bearing axes persistent.

Female-fertile spikelets solitary; distichous; 7-23 mm
long; compressed laterally to not noticeably compressed;
disarticulating above the glumes, or falling with the glumes.
Glumes present; two\ very unequal to more or less equal;
decidedly shorter than the adjacent lemmas; awned, or
awnless; non-carinate (or only slightly keeled towards the
tip)\ similar (ovate, oblongate or lanceolate, not awnlike).
Incomplete florets distal to the female-fertile florets, merely
underdeveloped; proximal incomplete florets absent.

Female-fertile florets 3-10 (but rarely more than 7).
Lemmas similar in texture to the glumes (leathery,
lanceolate); 5 nerved; entire, or incised; awnless, or
mucronate, or awned. Awns when present 1; from the sinus,
or apical; non-geniculate; much shorter than the body of the
lemma to much longer than the body of the lemma (to 20
mm). Palea present; relatively long. Lodicules 2;
membranous; ciliate. Stamens 3 (the anthers relatively
long). Ovary hairy. Fruit medium sized (4-6 mm); hilum
long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x -1 . Pooideae; Triticodae; Triticeae. 8 species.
North and south temperate. Mesophytic, or xerophytic;
maritime-arenicolous, or glycophytic. Cape Province. 1
naturalized species.

Intergeneric hybrids with Agropyron (A Agrotrigia
Tsvel ev), Hordeum (X Elytrohordeum Hylander), Aegilops,
Leymus (X Leymotrigia Tsvelev), Lophopyrum, Secale,
Triticum (X Trititrigia Tsvelev), Thinopyrum.

References. 1. Chippindall. 1955. Gr. & Past. 2. Dewey.
1984. Genomic classification in Gustafson, Gene
manipulation: 209.

Species treatment by M. Koekemoer.

Elvtrigia repens (L.) Nevski

Fig. 78. PI. 71.

( =Agropyron repens (L.)

Beauv.) 1.

Perennial; stoloniferous and
tufted (culms erect or geniculate-
ly ascending); 500-1 000(-l 200)
mm tall. Leaf blades 150-240
mm long; 6-12 mm wide. Spike-
lets 10-20 mm long. Spike
1 00-200(-300) mm long, erect, rachis not breaking up;
spikelets alternately arranged, usually overlapping, 3-8-
flowered; lemmas 8-13 mm long, blunt, sharp pointed or
shortly awned.

Flowering December and March. In waste places,
gardens and cultivated lands. Rare. Naturalized from
Europe. Biome: Fynbos and Grassland. Europe and the
Mediterranean area, introduced elsewhere. Weed (of
cultivation in many temperate countries).

Description: Bor 1985 (1817), Hitchcock & Chase 1950
(231). Illustration: Hitchcock & Chase 1950 (fig. 442).
Voucher: Acocks 17852. PRECIS code 9904345—00300.

133

Elytrophorus P. Beauv.

Echinalysium Trin.

Annual; caespitose. Culms 100-500 mm high; herba-
ceous (hydrophytic). Leaf blades linear; flat. Ligule an
unfringed membrane to a fringed membrane. The spikelets
of sexually distinct forms on the same plant (reduced,
sterile spikelets often present at the bases of the clusters).

Inflorescence a false spike, with clusters of spikelets on
reduced axes (the glomerules sometimes confluent to form
a cylinder); espatheate (but the glomerules and the clusters
within them subtended by the enlarged, spreading glumes
of the lower spikelets). Spikelet-bearing axes persistent.
Female-fertile spikelets associated with bractiform involu-
cres (constituted by the enlarged glumes of the lower

Spikelets compressed laterally; disarticulating above the
glumes. Glumes two; more or less equal; markedly shorter
than the spikelets, or about equalling the spikelets; awned
(shortly aristulate), or awnless (muticous); similar
(narrowly lanceolate, persistent, membranous). Incomplete
florets distal to the female-fertile florets; proximal incom-
plete florets absent.

Female-fertile florets 2-6. Lemmas similar in texture to
the glumes (membranous, granular, ovate); hairless (or
scabrid ciliate on keel and margins); 3 nerved; entire;
awned. Awns 1 ; median; apical (lemma becoming setaceous
at the summit); non-geniculate; much shorter than the body
of the lemma. Palea present; conspicuous but relatively
short; 2-nerved (or more?). Lodicules 1, or 2; fleshy;
glabrous. Stamens 1-3. Ovary glabrous. Fruit small; hilum
short; pericarp free; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, jc = 13. Arundinoideae; Danthonieae (?). 2-4
species. Tropical Africa, tropical Asia, Australia. Helo-
phytic. Namibia, Botswana, and Transvaal. 2 indigenous
species.

References. 1. Clayton. 1970. FTEA.

Species treatment by N.P. Barker.

1(0). Spikelets in globose clusters 10 mm wide and spaced
from 10 to 25 mm apart; clusters of spikelets
subtended by 2 to many acuminate bracts, up to

12 mm long; plants up to 500 mm tall

E. globularis

Spikelets clustered but clusters usually confluent,
about 8 mm wide, subtending bracts are usually
absent, but if present then shorter than spikelets;
plants up to 350 mm tall E. spicatus

Elytrophorus globularis Hack.

Fig. 79. PI. 72.

Annual; hydrophyte; to 500
mm tall. Leaf blades 30-500 mm
long; 3-8 mm wide. Spikelets 4-7
mm long (including lemma
awns). Leaves often overtopping
the inflorescence; panicle narrow,
interrupted, to 200 mm long;
spikelets clustered into dense,
globose aggregations 8-1 2 mm in
diameter and spaced at intervals of 10-25 mm up the axis,
sometimes confluent near apex; clusters subtended by 2 to
many bracts, each to 12 mm long; glumes 4. 5-6.0 mm long;
lemmas 3. 5-5.0 mm long, including stiff awn; anthers 1-3,
1 .5-2.0 mm long.

Flowering October to June. Vleis, pans, in shallow water
or damp places. Common. Biome: Savanna and Desert.
Tropical Africa. Pasture (while soft and green).

Description: Launert 1970 (160:76), Chippindall 1955
(187), Clayton et al. 1970-1982 (135-136). Illustration:
Chippindall 1955 (fig. 163). Voucher: Smith 2010. PRECIS
code 9903700-00100.

Elytrophorus spicatus (Willd.) A. Camus

Annual; hydrophyte; to 350
mm tall. Leaf blades to 250 mm
long; 2-4 mm wide. Spikelets
2. 0-3. 5 mm long (including
awns). Leaves generally shorter
than inflorescence; panicle
narrow, cylindrical, 20-250 mm
long, sometimes shortly branch-
ed; spikelets not densely clus-
tered; clusters 5-8 mm in diameter, often confluent, usually
lacking subtending bracts, or when present the bracts are

134

shorter than the spikelets; glumes 1.5-2. 5 mm long; lemmas
2. 0-2. 5 mm long, including slender awn; anthers 1-3, 0.3
mm long.

Flowering throughout the year. Vleis and pans. Com-
mon. Biome: Savanna. Northwards to the Congo and
Tanzania, and also from Australia.

Description: Launert 1970 (160:77), Chippindall 1955
(188), Clayton et al. 1970-1982 (135). Illustration: Chip-
pindall 1955 (fig. 162 - inflorescence only), Clayton et al.
1970-1982 (fig. 45). Voucher: Schweickerdt 2089. PRECIS
code 9903700-00200.

Enneapogon P. Beauv.

Calotheria Steud.

Annual (rarely), or perennial; caespitose. Culms
(30— )50— 1 000(— 1 100) mm high; herbaceous. Leaf blades
linear; flat, or rolled. Ligule a fringe of hairs.

Inflorescence paniculate ; contracted (feathery); espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 3.5-1 1 mm long; compressed laterally, or not
noticeably compressed, or compressed dorsiventrally; dis-
articulating above the glumes; not disarticulating between
the florets. Glumes two; very unequal to more or less equal;
about equalling the spikelets ; awnless; similar
(membranous). Lower glume 5-21 nerved. Incomplete
florets distal to the female-fertile florets, sterile or
rudimentary, sometimes reduced to awns; proximal incom-
plete florets absent.

Female-fertile florets 1-3. Lemmas decidedly firmer
than the glumes (coriaceous); 9 nerved; incised; awned.
Awns 9; median and lateral. The median awn similar in form
to the laterals; apical; non-geniculate; about as long as the
body of the lemma to much longer than the body of the
lemma. Palea present; relatively long (longer than the body
of the lemma). Lodicules 2; fleshy, or membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit small; hilum
short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
NAD-ME (7 species); XyMS+. PCR sheath outlines
uneven. PCR sheath extensions present. Maximum number
of extension cells 1 (usually). PCR cell chloroplasts elon-
gated; with well developed grana; centrifugal/peripheral to
centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 10. Chloridoideae; Pappophoreae. 30
species. In warm regions. Xerophytic; in open habitats
(bushland and semidesert); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 7 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Awns of lemma glabrous or scabrous, not plumose;

panicle open, not spikelike 2

Awns of lemma plumose, or at least with a fringe of
hairs on the margins in the lower half; panicle open

or spikelike 3

2(1). Spikelets 4-5 mm long; plant perennial; widespread

distribution E. scaber var. scaber

Spikelets 3 mm long; plant annual; Namibia

E. scaber var. (=De Winter & Hardy 8051)
3(1). Plant reedlike; lower leaves with deciduous blades;
restricted to limestone soils in the northeastern

Transvaal E. sp. (=Ellis 3208)

Plant not reedlike; all leaf blades persistent;

distributions various 4

4(3). Glume tips reddish brown, often shining; culm bases
erect, clad in hard shiny yellow sheaths; culms not

branched E. pretoriensis

Glume tips not reddish brown, not shining; culm bases
various; culms often branched above the base . . 5
5(4). Plant usually taller than 500 mm; leaf blades flat,
wider than 3 mm; panicles branched at least in the

lower half at maturity 6

Plant usually shorter than 500 mm (but E. scoparius
sometimes to 650 mm); leaf blades rolled, narrower
than 3 mm; panicles spikelike and unbranched . 7
6(5). Plant annual; panicle branches barely spreading, so
the central axis is hidden; plant with dense gland-
tipped hairs on leaves and culms . E. cenchroides
Plant perennial, with a woody rootstock clad by hairy
cataphylls; panicle branches spreading so the
central axis is exposed; plant with sparse gland-

135

tipped hairs or nearly glabrous . . . E. spathaceus
7(5). Culms often strongly geniculate or decumbent and
rooting at lower nodes; plant usually shorter than
250 mm; plant densely hairy, nodes usually with
a conspicuous ring of hairs; anthers 0.2— 0.7( — 1 .2)

mm long E. desvauxii

Culms erect; plant usually taller than 300 mm; plant
sparsely hairy, nodes lacking a conspicuous ring of
hairs; anthers 1.0-2. 5 mm long . . . . E. scoparius

Enneapogon cenchroides (Roem. & Schult.) C.E. Hubb.

Fig. 80. PI. 73.

Short-lived perennial, or an-
nual; tufted; to 1000 mm tall.

Leaf blades 30-250 mm long; 3-8
mm wide. Spikelets 3-5 mm long
Plant densely glandular-hairy;
panicle branched but contracted,
often dense and spikelike, central
axis hidden; lemma awns hair-
fringed; anthers 1.0-1. 5 mm long.

Flowering throughout the year (usually in summer, but
occasionally in winter in the north). Sandy soils, in
disturbed places and overgrazed veld. Common. Biome: Sa-
vanna, Grassland, and Nama-Karoo. North to Sudan,
through Arabia to India.

Description: Chippindall 1955 (236), Clayton et al.
1970-1982 (169). Illustration: Chippindall 1955 (fig. 211),
Clayton et al. 1970-1982 (fig. 55). Voucher: De Winter &
Wiss 4434. PRECIS code 9903570-00100.

Enneapogon desvauxii Beauv.

(-E. brachystachyum (Jaub.

& Spach) Stapf) 2.

Kalkgras, wondergras.

Possibly perennial, or annual;
densely tufted; 30-300 mm tall.

Leaf blades 25-250 mm long;
filiform or to 7 mm wide. Spike-
lets 3. 0-5. 5 mm long. Plant densely glandular-hairy, nodes
with a ring of hairs; culms often decumbent; panicle spike-
like, dense, unbranched; lemma awns hair-fringed; anthers
0.2-0.7(-1.2) mm long.

Flowering throughout the year (usually in summer but
rarely in winter in the north). Many habitats and soil types,
often in overgrazed veld. Common. Biome: Savanna,
Grassland, Nama-Karoo, Succulent Karoo, and Desert.
Throughout Africa, and in southern Asia, Central and South
America. Cleistogenes in basal leaf sheaths germinate in
place so that seedlings grow out of the old plant.

Description: Chippindall 1955 (237), Clayton et al.
1970-1982 (167). Illustration: Chippindall 1955 (fig. 212).
Voucher: Pole Evans 2066. PRECIS code 9903570-00200.

Enneapogon pretoriensis Stent

Perennial; densely tufted;

300-650 mm tall. Leaf blades
50-250 mm long; setaceous or to
3 mm wide. Spikelets 5. 5-7.0 mm
long. Plant wiry, culms
unbranched, bases erect, clad in
hard, shiny, yellowish sheaths;
panicle contracted, rarely open,
branched; glume tips reddish
brown, often shining; lemma awns hair-fringed; anthers 2.5
mm long.

Flowering November to May. Rocky hillsides, often on
northern aspect. Infrequent. Biome: Savanna and Grass-
land. Endemic.

Description: Chippindall 1955 (235). Illustration: Chip-
pindall 1955 (fig. 209). Voucher: Wasserfall 23-11-1944.
PRECIS code 9903570-00300.

Fig. 81.

Enneapogon scaber Lehm. var. scaber

Perennial; tufted; 70-350 mm
tall. Leaf blades 50-115 mm
long; 2-5 mm wide. Spikelets 4-5
mm long. Panicle open, branched,
not spikelike; lemma awns
glabrous or scabrous; anthers
0.8-1 .0 mm long.

Flowering throughout the year
(most commonly in summer but
in winter in winter rainfall areas). Hillsides among rocks.
Infrequent. Biome: Fynbos, Savanna, Nama-Karoo, Succu-
lent Karoo, and Desert. Apparently endemic.

Description: Chippindall 1955 (235). Illustration: Chip-
pindall 1955 (fig. 208). Voucher: Compton 23907. PRECIS
code 9903570-00400.

Enneapogon scaber var. (=De Winter & Hardy 8051)

Annual; to 200 mm tall
(usually less). Leaf blades to 170
mm long; 4 mm wide. Spikelets
3 mm long. Differs from the
typical variety in its annual habit,
smaller size and shorter spikelets.

Flowering March to June.

Rock crevices, gravel plains and
dry sandy riverbeds. Conserva-
tion status not known. Biome: Nama-Karoo. ?Endemic.

Voucher: De Winter & Hardy 8051. PRECIS code

136

Enneapogon scoparius Stapf

(= E.filifolius (Pilg.) Stapf ex
Garabedian) 2.

Perennial; densely tufted;

300-650 mm tall. Leaf blades
50-200(-250) mm long; filiform
or to 3 mm wide. Spikelets

3. 5- 4. 5 mm long. Plant wiry,
sparsely hairy; culms erect; pani-
cle dense, spikelike, unbranched; lemma awns hair-fringed;
anthers 1.0-2. 5 mm long.

Flowering throughout the year (in summer, but in winter
in northernNamibia). Dry grassland and among rocks on
hillsides. Common. Biome: Savanna, Grassland, Nama-
Karoo, and Desert. Southern tropical Africa.

Description: Chippindall 1955 (235). Illustration: Chip-
pindall 1955 (fig. 210). Voucher: Theron 586. PRECIS
code 9903570-00500.

Enneapogon spathaceus Goossens

Perennial; short-rhizomatous
and tufted (rootstock with hairy
cataphylls); 650-900 mm tall.

Leaf blades to 200 mm long; 3-4
mm wide. Spikelets 6-7 mm long.

Plant with sparse gland-tipped
hairs or nearly glabrous; panicle
open, branched, central axis
exposed; lemma awns hair-
fringed; anthers 2.5 mm long.

Flowering November to March. Sandveld. Conservation
status not known. Biome: Savanna. Endemic. Possibly a
hybrid between Enneapogon cenchroides and Schmidtia
pappophoroides, because of its intermediate characters and
restricted distribution.

Description: Chippindall 1955 (235). Voucher: Fisher
& Schweickerdt 543. PRECIS code 9903570-00600.

Enneapogon sp. (=EUis 3208)

Perennial; tufted; 550-1100
mm tall. Leaf blades to 170 mm
long; 3^f mm wide. Spikelets

5. 5- 6. 5 mm long. Plant reed-like,
culms stiffly erect; leaf blades de-
ciduous from lower nodes; pani-
cle open, branched, but narrow;
lemma awns hair-fringed.

Flowering January to Feb-
ruary. Restricted to soil overlying Malvernia limestone
formation. Rare. Biome: Savanna. ?Endemic, possibly also
in Zimbabwe. Vegetatively very similar to E. spathaceus
but remarkable because of its distinctive height and habit.

Voucher: Ellis 3208. PRECIS code 9903570-99999.

Enteropogon Nees

Macrostachya A. Rich.

Perennial; caespitose. Culms 200-1200 mm high ; herba-
ceous. Leaf blades linear; flat, or rolled (then involute-
filiform). Ligule a fringed membrane (short).

Inflorescence a single spike, or of spike-like main
branches, or a single raceme (with short pedicels)', digitate
or subdigitate (often), or non-digitate; espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary; biseriate; sessile to subsessile\ 5-8
mm long; compressed dor siventr ally, disarticulating above
the glumes; disarticulating between the florets. Glumes two
(hyaline); very unequal ; long relative to the adjacent
lemmas (i.e., the longer glumes); awned (G2, often), or

awnless; minutely bidentate, membranous. Incomplete
distal florets 1-2, sterile, stipitate, awned; proximal incom-
plete florets absent.

Female-fertile florets 1 , or 2 (L2 male or with a
hermaphrodite floret). Lemmas decidedly firmer than the
glumes (coriaceous, rigid); 3 nerved; entire, or incised;
awned. Awns 1 ; median; from the sinus, or apical; non-gen-
iculate; about as long as the body of the lemma to much
longer than the body of the lemma. Palea present; relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small; ellipsoid; hilum short; pericarp fused;
embryo small to large (up to 1/3 of the grain length).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
present, or absent. Maximum number of extension cells
when present 1-2. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 6
species. Africa, Seychelles, India, Formosa, Australia,
Pacific. Mesophytic to xerophytic; in shade, or in open

Fig. 82. Enteropogon macrostachyus

137

habitats (savanna on sand or clay); glycophytic. Namibia,
Botswana, Transvaal, and Natal. 4 indigenous species.

References. 1. Clayton et al. 1974. FTEA. 2. Clayton.
Kew Bull. 1967. 21: 105.

Species treatment by M. Koekemoer.

1(0). Spikelets 4-6-flowered; plants annual; spikes digitate,
4-9, 40-120 mm long; lemmas 3-5 mm long,
ciliate on margins only; awn 7-25 mm long ....

E. prieurii

Spikelets 2-3-flowered; plants perennial; spikes
solitary (rarely two), 60-200 mm long; lemmas
4.5-10.0 mm long, scabrid; awn 1-18 mm long . 2
2(1). Leaf sheaths strongly keeled, margins ciliate; awn of

lowest lemma 2. 5-8.0 mm long

E. monostachyos

Leaf sheaths not keeled, margins glabrous; awn of

lowest lemma 1-18 mm long 3

3(2). Awn of lowest lemma 10-18 mm long

E. macrostachyus

Awn of lowest lemma 1-5 mm long . . E. rupestris

Enteropogon macrostachyus (A. Rich.) Benth.

(-E. simplex (Schumach. &

Thonn.) A. Chev.) 1 .

Perennial; tufted; 500-1200
mm tall. Leaf blades 100-600
mm long; 3-7 mm wide. Spike-
lets 8-10 mm long. Spikelets 3-
flowered; lemma of lower floret
7-10 mm long, awn 10-18 mm
long.

Flowering November to June. Disturbed places or light
shade under trees. Locally common. Biome: Savanna.
Tropical Africa. Similar to E. rupestris, which has much
shorter lemma awns.

Description: Chippindall & Crook 1976 (233), Clayton
1967 (105), Clayton et al. 1970-1982 (332). Voucher:
Dinter 5702. PRECIS code 9903000-00100.

Enteropogon monostachyos (Vahl) K. Schum. subsp.

africanus Clayton

Perennial; tufted; 400-1000
mm tall. Leaf blades 150-300
mm long; 2-4 mm wide. Basal
leaf sheaths laterally flattened
and strongly keeled; lemma of
lower floret 6-8 mm long with an
awn 2. 5-8.0 mm long.

Flowering November to April.

Grey granite flats and sandy soil
near rivers, often in the shade. Locally common. Biome: Sa-
vanna. Southern tropical Africa. Related to E.
macrostachyus and E. rupestris, which have leaf .sheaths
rounded and margins glabrous.

Description: Clayton 1967 (105), Clayton et al.
1970-1982 (333). Voucher: Ward 3666. PRECIS code
9903000-00200.

Enteropogon prieurii (Kunth) Clayton

( =Chloris prieurii Kunth) 2.

Annual; tufted; 200-500 mm
tall. Leaf blades to 300 mm long;

2-5 mm wide. Spikelets 3-5 mm
long. Spikes 2-8, 50— 80( — 1 20)
mm long; spikelets 4-6-flowered,

4-6-awned; lemma awn 7-10 mm
long.

Fig. 82. PI. 74.

Flowering February. Deep white sand on palm flats.
Rare (in Namibia). Biome: Savanna. Tropical Africa to
Arabia. Very different from other species in this genus,
which have solitary spikes, are perennial and have spikelets
2-3-flowered. This species resembles Chloris superficially,
from which it is distinguished by dorsally compressed
lemma and a different grain shape.

Description: Hitchcock & Chase 1950 (504), Clayton et
al. 1970-1982 (342). Voucher: De Winter & Marais 4724.
PRECIS code 9903000-00300.

Enteropogon rupestris (J.A. Schmidt) A. Chev.

Bushy perennial; tufted;

500-1000 mm tall. Leaf blades
50-250 mm long. Spikelets 4-8
mm long. Culms branched; spike-
lets mostly 2-flowered; lemma of
lowest floret 4. 5-8.0 mm long
with awn 1-5 mm long.

Flowering March to May.

Black clay or humiferous loam,
among rocks and often on north-facing slopes. Locally
common. Biome: Savanna. Central Africa, Cape Verde
Islands. Similar to E. macrostachyus, which has much
longer lemma awns.

Description: Clayton 1967 (105), Clayton et al.
1970-1982 (332). Voucher: Giess & Loutit 14142. PRECIS
code 9903000-00400.

Entolasia Stapf

Perennial; long-rhizomatous, or caespitose. Culms
200-1200 mm high (sometimes straggling/climbing);
woody and persistent (wiry, bushy), or herbaceous;
branched above, or unbranched above. Ligule a fringe of
hairs.

Inflorescence of spike-like main branches, or paniculate
(but usually with sessile spiciform racemes, appressed to
the common axis); open, or contracted; espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary, or in pairs; secund\ consistently in
‘long-and-short’ combinations (rarely), or not in distinct
‘long-and-short’ combinations. Spikelets 2.5-6 mm long;
adaxiab, usually compressed dorsiventrally, falling with the
glumes. Glumes present; two; very unequal; awnless; very
dissimilar (lower tiny, hyaline; upper membranous,
equalling the spikelet). Proximal incomplete florets 7;
epaleate; sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes (membranous
to coriaceous); smooth to striate; not becoming indurated;
hairy ( densely silky -hairy)-, having the margins tucked in
onto the palea; with a clear germination flap; 3-5 nerved;
entire; awnless. Palea present (hairy between the keels); rel-
atively long. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small; hilum short; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 5 species. Tropical Africa, eastern
Australia. Helophytic, mesophytic, and xerophytic; in shade
and in open habitats (marshy places, damp grassland and
dry forest); glycophytic. Namibia, Botswana, and
Transvaal. 2 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

1(0). Spikelets 2. 2-2. 5 mm long; leaves linear-lanceolate

E. olivacea

Spikelets 4. 5-6. 5 mm long; leaves linear

E. imbricata

138

Fig. 83. Entolasia imbricata

Entolasia imbricata Stapf

Fig. 83. PI. 75.

Perennial; tufted; to 1500 mm
tall. Leaf blades to 500 mm long;

2-8 mm wide. Spikelets 4. 5-6. 5
mm long; 1.5 mm wide. Culms
erect, simple and stout at base;
leaf blades linear, tapering to an
acute point; inflorescence
100^150 mm long; spikelets pale
straw-coloured; upper glume 5
mm long.

Flowering January to March. Flood plain, seasonally
flooded to one meter. Infrequent. Biome: Savanna. Tropical

Description: Clayton et al. 1970-1982 (573). Voucher:
P.A. Smith 1876. PRECIS code 9901021-00100.

Entolasia olivacea Stapf

Perennial; rhizomatous; to
1000 mm tall. Leaf blades 50-100
mm long; 5-15 mm wide. Spike-
lets 2. 2-2. 5 mm long; 1 mm wide.

Culms erect, geniculate and
branched; leaf blades linear-lan-
ceolate, constricted at base, point
acutely acuminate; inflorescence
70-150 mm long; spikelets dull
green colour; upper glume 2.5 mm long.

Flowering January to March. In shade, moist places. In-
frequent. Biome: Savanna. Tropical Africa.

Description: Clayton et al. 1970—1982 (573). Voucher:
Johannsmeier 372. PRECIS code 9901021-00300.

Entoplocamia Stapf

Annual (robust). Culms (200 — )400— 1 100 mm high; her-
baceous; unbranched above. Leaf blades linear-lanceolate;
flat, or rolled. Ligule a fringe of hairs.

Inflorescence a single spike, or a single raceme, or pa-
niculate; contracted (the spikelets solitary or in clusters or
secondary spikes on the rachis of a simple or compound
spike); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 9-20 mm long; compressed laterally
(becoming twisted when mature); falling with the glumes;
not disarticulating between the florets (the spikelets falling
whole). Glumes two; very unequal to more or less equal;
markedly shorter than the spikelets; awnless; similar (thin,
membranous, ovate). Incomplete florets both distal and
proximal to the female-fertile florets ; distal incomplete
florets merely underdeveloped; proximal incomplete florets
2.

Fig. 84. Entoplocamia aristulata

139

Female-fertile florets 4-20. Lemmas decidedly firmer
than the glumes (cartilaginous at the base, chartaceous
above, hyaline at the margins); without a germination flap;
9-1 1 nerved; entire; mucronate to awned. Awns 1; median;
apical (the midnerve excurrent into a short, stout mucro or
awn); much shorter than the body of the lemma. Palea
present; relatively long. Stamens 3. Ovary glabrous. Fruit
small (2 mm); ellipsoid; hilum short; pericarp free; embryo
large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 1
species. Angola, southwest and southern Africa. Xerophytic
(but often grows in depressions where moisture collects);
in open habitats; halophytic (sometimes), or glycophytic
(usually). Namibia. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

Entoplocamia aristulata (Hack. & Rendle) Stapf

Fig. 84. PI. 76.

Robust annual; tufted;

(200— )400— 1 100 mm tall. Leaf
blades 75-150 mm long; 2-3 mm
wide. Spikelets 9-17 mm long.

Spikelets robust, spiny, laterally
compressed, sometimes twisted,
often in glomerate racemes on a
central axis; lemma 9-1 1 -nerved,
chartaceous, with a short, spiny,
deflexed awn.

Flowering February to May. Rocky outcrops or open
plains on brackish or calcareous soil. Infrequent (but
occasionally in dense stands in moist depressions). Biome:
Savanna, Nama-Karoo, and Desert. Angola.

Description: Stapf 1898-1900 (711), Chippindall 1955
(189). Illustration: Chippindall 1955 (fig. 165). Voucher:
Du Toit 258. PRECIS code 9903280-00100.

Eragrostis N. M. Wolf

Boriskerella Terekhov, Erochloe Raf., Erosion Lunell,
Exagrosis Steud., Neeragrostis Nicora, Macroblepharus
Philippi, Psilantha (K. Koch) Tzvelev, Roshevitzia
Tsvelev, Triphlebia Stapf, Vilfagrostis Doell.

Annual, or perennial; caespitose (sometimes shrubby),
or decumbent. Culms 100-3000 mm high; herbaceous
(usually), or woody and persistent (occasionally); branched
above, or unbranched above. Leaf blades linear ; flat, or
folded, or rolled; not disarticulating. Ligule a fringed
membrane to a fringe of hairs.

Inflorescence a false spike, with clusters of spikelets on
reduced axes (occasionally), or paniculate (often
glandular, characteristically scented ); open, or contracted;
espatheate. Spikelet-bearing axes persistent.

Spikelets 1-25 mm long; compressed laterally (usually
strongly so), or not noticeably compressed (rarely — Section
Cylindrostachya); usually disarticulating above the glumes,
or falling with the glumes (in some species); not disarticu-
lating between the florets (with persistent paleas), or disar-
ticulating between the florets. Glumes two (persistent or
deciduous); very unequal, or more or less equal; markedly
shorter than the spikelets; decidedly shorter than the adja-
cent lemmas', awnless; similar (membranous). Upper glume
1 nerved. All florets female-fertile, or with distal incom-
plete florets also present; proximal incomplete florets
nearly always absent (a very few species with 1-3
incomplete lower florets).

Female-fertile florets 2 (rarely), or 3^)5. Lemmas
similar in texture to the glumes, or decidedly firmer than

Fig. 85. Eragrostis curvula

140

the glumes (narrow, membranous to papery); hairless
(usually glabrous )\ without a germination flap; 1-3 nerved;
entire, or incised; awnless, or mucronate (very rarely almost
awned). Palea present (often persistent); relatively long (but
shorter than the lemma), or conspicuous but relatively short.
Lodicules when present 2; fleshy; glabrous. Stamens 1-3.
Ovary glabrous. Fruit small; hilum short; pericarp usually
fused (but rather readily detachable in some species), or free
(e.g. in E. megalosperma, E. stapfiana)', embryo large.

Photosynthetic pathway and related features. C4 (with
the startling exception of E. walteri : see Ellis 1984);
NAD-ME (14 species); XyMS+. PCR sheath outlines
uneven, or even, or uneven to even. PCR sheath extensions
present, or absent. PCR cell chloroplasts ovoid, or elon-
gated; with well developed grana; centrifugal/peripheral, or
centripetal, or centrifugal/peripheral to centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 350
species. Cosmopolitan, mostly subtropical. Helophytic, or
mesophytic, or xerophytic; mostly in open habitats (often
on poor soils or disturbed ground); maritime-arenicolous,
or halophytic, or glycophytic. Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. Indigenous species (79), naturalized
species (4).

References. 1. De Winter in Chippindall. 1955. Gr. &
Past. 2. De Winter. 1960. Bothalia 7: 387. 3. De Winter.
1961. Bothalia 7: 467. 4. De Winter. 1961. Kirkia 1: 100.
5. De Winter. 1966. Bothalia 9: 137. 6. De Winter. 1969.
Bothalia 10: 72. 7. Launert. 1970. FSWA. 8. Gordon-Gray.
1972. FI. Natal. 9. Clayton et al. 1974. FTEA. 10. Phillips.
1982. Kew Bull. 37:133.

Species treatment by L. Smook.

1(0). Spikelets disarticulating below the glumes and falling

entire as a unit 2

Spikelets breaking up in various places above the

glumes at maturity 3

2( 1 ). Plants perennial; palea wings broad and entire: lemma

narrowly ovate in profile E. superba

Plants annual; palea wings very broad and usually

lacerate; lemma lanceolate in profile

E. pilgeriana

3(1). Lemma deeply trilobed at the apex, with a median
awn 0.8-1. 5 mm long and the lateral nerves
excurrent into a short distinct mucro or awn ....

E. aristata

Lemma entire or shallowly lobed at the apex, with or
without a median awn, lateral nerves not excurrent

into minute mucros or awns 4

4(3). Palea keels with hairs 0.3-1. 3 mm long for most of
its length which are usually exserted beyond the
lemma (note: long hairs are found at the base of the
palea keels in E. capensis and E. cimicina but these

are not exserted) 5

Palea keels glabrous or scabrid, or ciliate with hairs
shorter than 0.3 mm and not exserted beyond the

lemma 10

5(4). Rachilla persistent, the lemmas and/or paleas
breaking up from the base upwards; plants

perennial 6

Rachilla fragile, the lemmas, paleas and part of the
rachilla intemode breaking off as units from the

apex downwards; plants annual 7

6(5). Lemma with lateral nerves and sometimes the keel

with hairs 0.3-1. 2 mm long E. lappula

Lemma with lateral nerves and keel glabrous or with

hairs less than 0.3 mm long E. hierniana

7(5). Lemma keel (at least in the upper lemmas of a
spikelet) with stiff hairs 0. 2-0.4 mm long,

especially towards the base E. ciliaris

Lemma keel smooth, scabrid or scaberulous 8

8(7). Inflorescence contracted, dense, branches appressed
to the main axis; spikelets crowded; anthers 0. 3-0.4

mm long E. arenicola

Inflorescence open, branches spreading; spikelets

distant; anthers to 0.3 mm long 9

9(8). Inflorescence usually dense with many spikelets and
with sticky glands (noticable due to particles

adhering to the glandular area) E. viscosa

Inflorescence not dense, with fewer spikelets,
eglandular or with non-sticky glands . . E. tenella
10(4). Vegetative parts of plants with swollen-tipped

glandular hairs 11

Vegetative parts of plants lacking swollen-tipped

glandular hairs 12

1 1(10). Spikelets 1.5-2. 5 mm wide; anthers 0.6-1. 0 mm
long; pedicels slender, flexible and with an

annular gland E. annulata

Spikelets 1.0-1. 5 mm wide; anthers 0.2 mm long;
pedicels stout, rigid, lacking annular glands . .

E. pygmaea

12(10). Lemma with distinct elongated dark patches on or

next to the lateral nerves E. caesia

Lemma without distinct elongated dark patches, or
these only occasionally scattered on the lemmas

but not confined to the lateral nerves 13

13(12). Spikelets to 2.1 mm wide 14

Spikelets 2.2 mm and wider 95

14(13). Inflorescence consisting of 2-8 dense globose to
ovoid clusters of spikelets distant from each other
along the main axis; spikelets elliptic to narrowly

oblong E. congesta

Inflorescence not as above, if spikelets in clusters
these clusters coalescent with only the lower

clusters sometimes distant 15

15(14). Culms with sticky glandular patches below the
nodes and on the leaf sheaths below the collar
(noticable due to particles adhering to the

glandular areas) E. gummiflua

Culms and sheaths eglandular or with non-sticky

glandular patches 16

16(15). Plants annual 17

Plants perennial 45

17(16). Anthers to 0.5 mm long 18

Anthers 0.6-1. 5 mm long 40

18(17). Spikelets sessile, in wedge-shaped clusters, these

coalescent into a spikelike inflorescence

E. patens

Spikelets pedicellate or sessile, distant or crowded,
or clustered but the clusters not wedge-shaped;

inflorescence open or contracted 19

19(18). Rachilla fragile, spikelets breaking up from the

apex downwards 20

Rachilla persistent or the upper part of the rachilla
eventually becoming fragile, spikelets with

lemmas and/or paleas breaking up from the base

upwards (persistent in E. tef) 21

20(19). Lemma obtuse to truncate; caryopsis subglobose

E. aspera

Lemma acute to acuminate; caryopsis ovate-elliptic
E. leersiiformis

21(19). Anthers 2 E. gangetica

Anthers 3 22

22(21). Spikelets to 1.2 mm wide 23

Spikelets 1.3-2. 1 mm wide 33

23(22). Lowest lemma 2. 0-2. 7 mm long 24

Lowest lemma 0.5-1 .9 mm long 25

24(23). Upper glume 1/2-2/3 the length of the lemma
directly above in the intact spikelet .... E. tef
Upper glume barely reaching or just covering the
base of the lemma directly above in the intact
spikelet E. tenuifolia

25(23). Spikelets narrowly elliptic when young to broadly
ovate at maturity; 1— 2(— 3) florets per spikelet .

E. biflora

Spikelets oblong, linear to lanceolate; usually with
more than 3 florets per spikelet 26

141

26(25). Lower glume to 1/3 the length of the lemma above
in the intact spikelet, weakly keeled and loosely

folded 27

Lower glume 1/3 the length to as long as the lemma
above in the intact spikelet, strongly keeled and

tightly folded 31

27(26). Lemmas on the same side of the rachilla barely
reaching the lemma above in the intact spikelet

E. remotiflora

Lemmas on the same side of the rachilla distinctly
overlapping the lemma above in the intact

spikelet 28

28(27). Lower lemma 1.8 mm and longer . . E. tenuif'olia

Lower lemma to 1.7 mm long 29

29(28). Inflorescence robust, spikelets irregular and

densely condensed along the primary branches,
these either spreading or appressed to the main
axis, branches stout and rigid; lemmas ovate-

elliptic E. homomalla

Inflorescence delicate, loose with spikelets distant,
branches slender, usually flexible; lemmas

broadly ovate 30

30(29). Lowest lemma 0. 7-1.0 mm long, lateral veins
indistinct; lemmas hardly diminishing in length
towards the apex of the spikelet; inflorescence
branches not bearded in the axils . E. aethiopica
Lowest lemma 1.0-1. 6 mm long, lateral veins
distinct; lemmas conspicuously becoming shorter
towards the apex of the spikelet; inflorescence

branches bearded in the axils E. pilosa

31(26). Caryopsis subglobose; basal leaf sheaths densely
covered for the whole length with bulbous-based

hairs E. pygmaea

Caryopsis oblong-elliptic; basal leaf sheaths
glabrous or with a few scattered bulbous-based

hairs 32

32(31). Leaf blade with margins smooth, with raised
glands; leaf blade with midrib with glandular

dots E. kingesii

Leaf blade with margins scabrid, eglandular; leaf
blade with midrib eglandular .... E. virescens

33(22). Caryopsis subglobose 34

Caryopsis oblong to elliptic 35

34(33). Lowest lemma 1.7-2. 8 mm long, obtuse

E. cilianensis

Lowest lemma 1.0- 1.6 mm long, acute

E. pygmaea

35(33). Upper glume barely reaching or just covering the
base of the lemma directly above in the intact

spikelet E. tenuifolia

Upper glume 1/3-2/3 the length of the lemma

directly above in the intact spikelet 36

36(35). Inflorescence branches usually more than 40 mm

long, flexible, pedicels slender E. tef

Inflorescence branches usually less than 40 mm

long, rigid, pedicels usually stout 37

37(36). Lemma acute 38

Lemma obtuse 39

38(37). Inflorescence sparsely branched, with a few
spikelets not too densely crowded; lowest lemma
1 .4— 1 ,6(— 1 .8) mm long; spikelets 1 .0—1 .5(— 1 .8)

mm wide E. kingesii

Inflorescence much branched, with many spikelets
densely crowded; lowest lemma ( 1 .8— )2.0— 2.5
mm long; spikelets ( 1 .7— )2.0— 2.5 mm wide . . .

E. procumbens

39(37). Glumes unequal; leaf blade margins scabrid ....

E. barrelieri

Glumes subequal; leaf blade margins with raised

glands E. minor

40(17). Lateral nerves of lemma with glandular dots ....

E. laevissima

Lateral nerves of lemma without glandular dots .

41

41(40). Lemma very broadly ovate to almost oblate, glossy,
coriaceous, with a broad, clear membranous
margin in the upper part .... E. membranacea

Lemma broadly elliptic or oblong to lanceolate,
dull, chartaceous to membranous, without a

distinctly different upper margin 42

42(41). Lowest lemma 1.8-2. 7 mm long . E. omahekensis

Lowest lemma to 1.7 mm long 43

43(42). Lowest lemma obovate-elliptic, 1.0-1. 5 mm long,
apex truncate to broadly rounded . . . E. porosa

Lowest lemma broadly elliptic to broadly oblong-
ovate, 1.5- 1.7 mm long, apex obtuse to subacute
44

44(43). Lateral nerves of lemma obscure; palea
oblanceolate to narrowly obovate, margins

touching or overlapping at the apex

E. cylindriflora

Lateral nerves of lemma prominent; palea obovate,
margins close but not touching or overlapping at

the apex E. glandulosipedata

45(16). Inflorescence with spikelets directly on the main
axis or with branches closely appressed to the
main axis or spreading but then sparsely
branched with only primary branches or very
short secondary branches and spikelets appressed

to the branches 46

Inflorescence moderately to much branched,
branches spreading, spikelets usually spreading

46(45). Spikelets reddish brown; anthers 2 . E. chapelieri
Spikelets various shades and combinations of

green, grey, purple or red; anthers 3 47

47(46). Culms wiry and matted 48

Culms not as above 49

48(47). Glumes acuminate; lemmas lanceolate in profile,

lateral nerves distinct E. walteri

Glumes obtuse; lemmas broadly ovate in profile,

lateral nerves indistinct E. volkensii

49(47). Lateral nerves of lemma reaching to the upper
margins and usually excurrent into minute
mucros, small glandular dots present; caryopsis

subglobose E. crassinervis

Lateral nerves of lemma not reaching to the upper
margins, not excurrent into mucros, eglandular;

caryopsis ovate, elliptic to oblong 50

50(49). Plants densely tufted; leaves mainly basal .... 51
Plants loosely tufted or creeping; leaves mainly

cauline 58

51(50). Palea keels flat, usually 0.1 mm or wider .... 52
Palea keels a raised ridge or narrow line to 0.1 mm

wide 53

52(51). Plants robust, culms 2-4 mm wide; lowest lemma
1 .4-2.0 mm long, broadly obtuse . . . E. pallens
Plants slender, culms to 2 mm wide; lowest lemma
2-3 mm long, acute to acuminate . E. nindensis

53(51). Glumes membranous to chartaceous 54

Glumes cartilaginous 55

54(53). Lemmas on the same side of the rachilla

overlapping the lemma above up to just over 1/2;

spikelets linear to oblong E. curvula

Lemmas on the same side of the rachilla

overlapping the lemma above by 2/3 or more;

spikelets lanceolate to narrowly ovate

E. stenothyrsa

55(53). Upper glume lanceolate-oblong; lateral nerves of

lemma distinct E. elatior

Upper glume ovate, boat-shaped; lateral nerves of

lemma indistinct 56

56(55). Basal sheaths glabrous or thinly hairy with pale,

silky hairs E. racemosa

Basal sheaths with dense yellowish woolly hairs .
57

57(56). Inflorescence branches spreading from the main

axis E. sclerantha subsp. sclerantha

Inflorescence branches erect and appressed to the
main axis .... E. sclerantha subsp. villosipes
58(50). Upper glume tapering into a long, thick-textured

acuminate, awn-like apex E. walteri

Upper glume acute or obtuse to broadly obtuse, not
awn-like 59

142

59(58). Anthers 0.2-0. 3 mm long; lowest lemma 1 .5( — 1 .7)

mm long E. sarmentosa

Anthers 0.6-1. 3 mm long; lowest lemma 1. 6-3.0

mm long 60

60(59). Inflorescence usually shorter than 40 mm; spikelets
with lemmas conspicuously diminishing in

length towards the apex E. sabulosa

Inflorescence usually 50 mm or longer; spikelets
with lemmas hardly diminishing in length

towards the apex 61

61(60). Inflorescence slender, unbranched or sparsely
branched; spikelets solitary on main axis or 2-6
per branch closely appressed to the main axis,
spikelets or branches distant, not overlapping

each other E. elatior

Inflorescence robust, moderately branched;
spikelets more than 6 per branch, branches
appressed to main axis, and overlapping each

other E. inamoena

62(45). Spikelets very broadly ovate E. habrantha

Spikelets linear to oblong, to ovate or elliptic . 63
63(62). Lower glume to 1/3 the length of the lemma above
in intact spikelet, upper glume barely reaching
to just overlapping the base of the lemma above

64

Characters not occuring in the above combination

65

64(63). Lateral lemma nerves with glandular dots; plants

robust E. plana

Lateral lemma nerves eglandular; plants slender .

E. tenuifolia

65(63). Palea keels thickened, either broad or flat or a
prominent ridge along the entire length .... 66
Palea keels thickened into a narrow line or obscure
and apparently the palea folded only, without any
thickening, occasionally keels slightly wider

towards the base 68

66(65). Palea obovate, usually protruding from the lemma;

lateral nerves of lemma indistinct or a faint line;
spikelets glossy, rachilla fragile; plants lacking a

creeping rhizome E. pallens

Palea oblanceolate to narrowly obovate; lateral
nerves of the lemma distinct; spikelets dull,
rachilla persistent; plants with creeping rhizomes

67

67(66). Palea narrowly obovate, membranous except for the

keels, apex usually truncate to obtuse

E. inamoena

Palea oblanceolate, thick-textured, apex acute to

subacute E. patentissima

68(65). Palea margins not meeting or overlapping except

occasionally at the base 69

Palea margins meeting or overlapping along their

entire lengths or at the apex only 77

69(68). Basal sheaths densely covered with woolly hairs

E. sclerantha subsp. sclerantha

Basal sheaths glabrous or with scattered hairs, or
densely hairy at the very base only, hairs not

woolly 70

70(69). Spikelets narrowly elliptic . E. pseudosclerantha

Spikelets linear-oblong 71

71(70). Culm nodes (those nodes without branches) hairy

72

Culm nodes (those nodes without branches)

glabrous 73

72(71). Plants sprawling, stoloniferous and rooting at the

nodes; leaf blades 2-6 mm wide

E. barbinodis

Plants geniculate or straight, but erect, not
sprawling, not stoloniferous or rooting at the

nodes; leaf blades 1 .5-2.0 mm wide

E. lehmanniana var. chaunantha

73(71). Pedicels with an annular gland

E. moggii var. moggii

Pedicels eglandular or with glandular patches or

dots, but no annular gland 74

74(73). Basal sheaths with nerves very close together at the

base, forming prominent squarish ridges usually
with long hairs in the deep furrows between the

nerves E. curvula

Basal sheaths with the nerves wide apart at the base,
forming obscure or roundish ridges with shallow
furrows or furrows absent, glabrous or only hairy

at the very base 75

75(74). Leaf blades usually 3-5 mm wide, narrowing
abruptly into a long, thin apex, curling when dry

E. rigidior

Leaf blades usually to 3 mm wide, narrowing
gradually to the apex, not curling when dry . 76
76(75). Inflorescence with lowest branches whorled, and
with long hairs in the axils; spikelets untidy and
whitish in appearance because the apical margins
of the lemmas and glumes are whitish,
membranous and usually torn . E. trichophora
Inflorescence with lowest branches not whorled and
the axils glabrous; spikelets appearing tidy
because the glumes and lemmas are firm around
the apices . E. lehmanniana var. lehmanniana
77(68). Inflorescences with lowest branches whorled,
pseudo-whorled or clustered around the main

axis 78

Inflorescences with lowest branches not arranged

as above 87

78(77). Basal sheaths densely hairy at the base 79

Basal sheaths glabrous or obscurely hairy .... 82

79(78). Lemmas greyish green to near the apex, then
yellowish grading into white at the margins, often
flushed purple below the yellow; inflorescence
branches (excluding main axis) thickly covered

with minute prickles E. rotifer

Lemmas variously coloured, the apex the same
colour as the rest of the lemma, or white with the
yellowish patch absent; inflorescence branches
smooth, or if scabrid then the prickles large and

not densely packed 80

80(79). Plants slender, wiry; culms usually much branched
and geniculate, often rooting at the nodes; collar
of leaf sheath often with round, usually purple
glandular dots; glumes as long as the lemmas
directly above, lower glume wide and covering
most of the lemma above in the intact spikelet

E. trichophora

Plants robust, culms usually unbranched,
occasionally geniculate, not rooting at the nodes;
collar of leaf sheath often lacking round
glandular dots; glumes variable in length, lower
glume usually not so broad that it covers most of

the lemma above in the intact spikelet 81

81(80). Culms easily compressed; leaf blades usually 4-10

mm wide, flat E. jeffreysii

Culms not easily compressed; leaf blades to 3 mm
wide, usually rolled or appearing setaceous . . .

E. curvula

82(78). Lemma strongly keeled, keel prominent and
obvious for the entire length of the lemma . 83
Lemma not strongly keeled, keel obscure or only
prominent in the upper part of the lemma . . 85
83(82). Inflorescence effuse, much branched, the shortest
pedicel of the spikelet pair as long as or longer
than the spikelet; spikelets spreading on the

branches E. micrantha

Inflorescence moderately branched; shortest
pedicel of spikelet pair shorter than the spikelet;

spikelets condensed on the branches 84

84(83). Spikelets to 1.5 mm wide; leaf blades 2-3 mm wide;

plants moderately slender; lower glume
translucent, smooth or scabrid only on the keel

or at the apex E. heteromera

Spikelets 1.5-2. 5 mm wide; leaf blades 5-10 mm
wide; plants robust; lower glume opaque for most

of its surface, rough E. acraea

85(82). Lower glume 4/5 to slightly longer than the lemma
above in the intact spikelet; leaf sheaths at the
collar with small round glandular dots, usually

143

flushed purple; leaves mainly cauline, narrowing
abruptly to a point at the apex; plants erect, culms

usually branched and geniculate

E. trichophora

Lower glume to 3/4 the length of the lemma above
in the intact spikelet; leaf sheaths at the collar
without small round glandular dots; leaves

mainly a dense basal tuft 86

86(85). Plants tall, robust; spikelets 5-1 1 -flowered

E. planiculmis

Plants short to moderately tall; spikelets 2— 3(— 5)-

flowered E. stapfii

87(77). Lemma nerves with small glandular dots, which
often give the nerves a lumpy appearance ....

E. laevissima

Lemma nerves without glandular dots 88

88(87). Lemma bicoloured, usually deep purple to violet,
yellowish at the apex with whitish margins,
occasionally pallid with yellow; rhizome oblique

E. bicolor

Characters not present in the above combination .

89

89(88). Leaf blades 5-10 mm wide; glumes opaque, usually

rough E. acraea

Leaf blades to 4.5 mm wide; glumes translucent,
usually smooth, occasionally scaberulous
towards the apex 90

90(89). Inflorescence 25-80 mm long; leaf blades only
slightly tapered to the apex; culm nodes usually
with long spreading white hairs; plants
stoloniferous and often rooting at the nodes . .

E. sabinae

Inflorescence usually longer than 80 mm, if shorter,
leaf blades tapering into a very long filiform
apex; culm nodes glabrous; plants not

stoloniferous or rooting at the nodes 91

91(90). Basal sheaths hairy for quite a way up from the
base, with long hairs especially in the deep
furrows between the prominent ridges formed by
the nerves; inflorescence extremely variable . .

E. curvula

Basal sheaths glabrous or obscurely hairy or
densely hairy only at the extreme base, furrows

shallow or absent between the nerves 92

92(91). Plant base with culms not densely compacted,

easily separable to individual culms 93

Plant base with culms strongly and densely
compacted, not easily separable into individual

culms 94

93(92). Weak perennial; inflorescence effuse, spikelets
spreading from each other and from the branches

E. micrantha

Moderately strong perennial; inflorescence open,
spikelets appressed to the branchlets and close to

one another E. heteromera

94(92). Leaf blades straight or drooping; spikelets linear

E. planiculmis

Leaf blades very curly; spikelets narrowly obovate

E. chloromelas

95(13). Basal sheaths with long, dense, woolly hairs at the
base (these sometimes only visible on the inner

sheaths) 96

Basal sheaths glabrous or hairy but not with long
woolly hairs, or hairy only at the extreme base

of the sheaths 98

96(95). Spikelets dark olive-green; lemma apex acute,
lateral nerves conspicuous; rachilla persistent .

E. sclerantha subsp. sclerantha

Spikelets pallid to dark purple; lemma apex broadly
obtuse to broadly truncate, lateral nerves

conspicuous, rachilla fragile 97

97(96). Lowest lemma truncate E. truncata

Lowest lemma broadly obtuse E. bergiana

98(95). Plants annual 99

Plants perennial 107

99(98). Upper glume barely reaching the base of the lemma

above in the intact spikelet E. tenuifolia

Upper glume 1/4 as long to longer than the lemma

above in the intact spikelet 100

100(99). Lemma narrowly to broadly obtuse in profile .

101

Lemma acute to acuminate in profile 103

101(1 00). Spikelets glossy; lemma margins differing in
texture from the rest of the lemma, being clear,

thinly membranous and often torn

E. membranacea

Spikelets dull; lemma margins of similar texture

to the rest of the lemma 102

102( 101 ). Spikelets ovate to very broadly oblong, rachilla
fragile, breaking up from the apex downwards;
anthers 1.0-1. 2 mm long; caryopsis obovate

• E. brizantha

Spikelets narrowly ovate to narrowly oblong,
rachilla persistent, lemmas and/or paleas
breaking up from the base upwards, rachilla
often breaking off above the glumes before all
the lemmas have fallen; anthers 0.2-0. 3 mm

long; caryopsis subglobose . . E. cilianensis
103(100). Lemma acuminate in profile, often awned ....

E. dinteri

Lemma acute in profile, unawned, but a mucro
present or absent 104

104( 103). Inflorescence with the branches slightly
spreading and the spikelets few and distant .

E. rogersii

Inflorescence with the branches usually
appressed to the main axis, occasionally
spreading in the lower part and the spikelets

many and densely crowded 105

105( 104). Glumes longer than the lemmas directly above

them in the intact spikelet

.... E. macrochlamys var. macrochlamys
Glumes shorter to as long as the lemmas directly

above them in the intact spikelet 106

106( 105). Spikelets oblong to elliptic, rachilla usually
persistent or breaking off above the glumes
before all the lemmas have fallen; lemmas
and/or paleas breaking up from the base

upwards E. procumbens

Spikelets broadly oblong to broadly ovate,
rachilla fragile; lemmas and/or paleas breaking

up from the apex downwards

E. macrochlamys var. wilmaniae

107(98). Palea keels with lower portion very broad and
projecting from the rest of the keel (Fig. 91)

108

Palea keels of variable width, entire, the widest
part not projecting from the rest of the keel .

109

1 08( 1 07). Palea keels with lower broad portion ending in a
deep notch at the top; palea apex acute ....

E. echinochloidea

Palea keels with lower broad portion rounded or
only shallowly notched at the top; palea apex

rounded E. x pseud-obtusa

109( 107). Lemma with lateral nerves indistinct 110

Lemma with lateral nerves clearly distinct . 1 14
1 10( 109). Culms straggling and matted; cauline leaves
usually becoming reflexed .... E. volkensii
Culms erect or sometimes geniculate but not
matted or straggling; cauline leaves never

reflexed Ill

111(110). Lower glume lanceolate 112

Lower glume ovate 113

1 12(1 1 1). Plants 500-800 mm tall, not sprawling, dense

basal tuft of leaves absent; inflorescence
80-100 mm long .... E. pseudosclerantha
Plants 300-400 mm tall, sprawling, dense basal
tuft of leaves present; inflorescence to 75 mm
long E. lamprospicula

144

1 13(1 1 1). Rachilla persistent, lemmas and/or paleas
breaking up from the base upwards; palea keels
with a narrow thickened ridge, not winged

E. racemosa

Rachilla fragile, spikelets breaking up from the
apex downwards; palea keels broad and

winged E. nindensis

1 14( 109). Lemma obtuse to subobtuse or rounded ... 115
Lemma acuminate, acute to subacute, or truncate
to concave between the keel and the lateral

nerves which reach the margin 118

1 15(1 14). Palea broadly elliptic to round; rachilla very

fragile E. obtusa

Palea ovate, elliptic to obovate; rachilla
persistent, or sometimes with the upper portion

fragile 116

1 16(1 15). Spikelets with the opposite rows of the florets
hardly overlapping at their bases, rachilla
usually visible between the florets; leaves
mainly cauline, culms often branched in the

upper parts E. scopelophila

Spikelets with opposite rows of florets closely
packed and overlapping at the bases, rachilla

not visible; leaves mainly basal 117

1 17(1 16). Lemma dull, granular; palea narrowly obovate,
membranous to subcartilaginous between the
margin and keel, margins close together at the
base becoming farther apart towards the apex

E. capensis

Lemma shiny; palea elliptic, thickly cartilaginous
especially between the margins and the keels,
margins nearly meeting along the entire length

of the palea E. cimicina

1 18(1 14). Inflorescence with most pedicels more than 3

times the length of the spikelets

E. patentissima

Inflorescence with most pedicels (excluding the
terminal ones) shorter than or just as long as

the spikelets 119

1 19(1 18). Palea margins almost meeting to overlapping
along the entire length of the palea .... 120
Palea margins far apart for most of the length of

the palea 122

120(1 19). Inflorescence slender, sparsely branched,
spikelets either solitary or 2-6 clustered
together and distant along the main axis . . .

E. elatior

Inflorescence open or contracted but then the
branches overlapping; moderately to well

branched 121

121(120). Palea long and narrow, margins touching along
their entire length and overlapping at the apex,
keels a thin obscure line; spikelets 5-7-

flowered; leaf blades 5-10 mm wide

E. acraea

Palea narrowly obovate, margins slightly apart or
just touching along the entire length, keels
narrow but distinct; spikelets 7^f0-flowered;

leaf blades 2-4 mm wide E. inamoena

122(1 19). Palea keels narrow, rounded . E. scopelophila

Palea keels broad and flattened 123

123(122). Palea keels very wide in lower 2/3, distinctly
narrowing in the upper 1/3, excurrent to a soft

mucro at the apex E. walteri

Palea keels the same width throughout or only

slightly narrower at the apex 124

124(123). Anthers 3; lemma with lateral nerves raised and
prominent, excurrent into minute mucros,
usually gland-dotted; caryopsis subglobose

E. crassinervis

Anthers 2; lemma with lateral nerves usually not
raised and prominent, not excurrent into
minute mucros, eglandular; caryopsis broadly
ellipsoid E. chapelieri

Eragrostis acraea De Winter

Robust perennial; densely
tufted; to 2000 mm tall. Leaf
blades 200-300(-600) mm long;

5-10 mm wide. Spikelets 5-7 mm
long; 1.5-2. 5 mm wide. Basal
sheaths glabrous or obscurely
hairy at the base; inflorescence
moderately branched, spreading
or contracted, spikelets condens-
ed on the branches, most pedicels shorter than the spikelets;
spikelets oblong to oblong-elliptic, 5-7-flowered, rachilla
persistent, lemmas and/or paleas breaking up from the base
upwards; glumes usually rough, opaque; lemma acute to
subacute, strongly keeled, keel prominent for the entire
length of the lemma, lateral nerves prominent; palea long
and narrow, keels a thin line, entire, scabrid to smooth, mar-
gins touching along the entire length and overlapping at the
apex; anthers 3, 1 .2-1 .7 mm long, caryopsis oblong-elliptic.

Flowering November to April. Mountainous areas, often
between rocks and in disturbed places. Locally common.
Biome: Grassland and Savanna. Zimbabwe. Domestic use
(temporary thatching that does not last), or pasture (grazed
when young, unpalatable when old, green in winter).

Description: De Winter 1961 (100), Chippindall &
Crook 1976 (151). Voucher: Codd & Dyer 9074. PRECIS
code 9902860-00100.

Eragrostis aethiopica Chiov.

Annual; loosely tufted (erect);
to 600 mm tall. Leaf blades
30-200 mm long; 1-3 mm wide.

Spikelets 1.7-5. 0 mm long;

0.7-1.0 mm wide. Inflorescence
delicate, loose, branches and ped-
icels slender and flexible, axils of
the branches not bearded, the
spikelets distant; spikelets linear
to oblong, lemmas on the same side of the rachilla overlap-
ping the lemma above and hardly diminishing in length
towards the apex of the spikelet, rachilla persistent, the lem-
mas and/or paleas breaking up from the base upwards; low-
er glume to 1/3 the length of the lemma above in the intact
spikelet, weakly keeled; lemma broadly ovate with lateral
veins not visible, lowest lemma 0. 7-1.0 mm long; palea
keels usually smooth; anthers 3, 0. 1-0.2 mm long; caryop-
sis ellipsoid.

Flowering January to May. Damp sand or black clay in
small vleis, pan edges and riverbeds. Biome: Savanna.
Northwards to east Africa and Ethiopia. Resembles E.
pilosa, which has the lemmas conspicuously shorter
towards the apex of the spikelet, and E. remotiflora, in
which the lemmas on the same side of the rachilla do not
overlap the base of the lemma above it.

Description: De Winter in Chippindall 1955 (153), Clay-
ton et al. 1970-1982 (215). Voucher: Theron 2955. PRECIS
code 9902860-00200.

Eragrostis annulata Rendle ex Scott Elliot

Annual; tufted; to 350 mm
tall. Leaf blades 50-100 mm
long; to 3 mm wide. Spikelets
5-15 mm long; 1.5-2. 5 mm wide.

Vegetative parts with glandular
hairs with swollen tips; inflores-
cence open with spikelets spread-
ing, pedicels long, flexible and
with a single annular gland;
spikelets with the upper part of the rachilla fragile and the
lower part persistent; lemma and palea keels glabrous or
scabrid; anthers 3, 0. 6-1.0 mm long; caryopsis broadly ob-
long to broadly ovate.

145

Flowering February to May. On a variety of soils,
especially sandy or stony ground and calcareous soil where
the water table is high, and in disturbed areas. Locally com-
mon. Biome: Savanna, Nama-Karoo, and Desert. Angola.
Resembles E. cilianensis , which lacks swollen-tipped
glandular hairs and has a subglobose caryopsis.

Description: Stapf 1898-1900 (619), De Winter in Chip-
pindall 1955 (178). Illustration: De Winter in Chippindall
1955 (fig. 150). Voucher: Acocks 12639; Theron 1967.
PRECIS code 9902860-00300.

Eragrostis arenicola C.E. Hubb.

Annual; loosely tufted (erect);
to 350 mm tall. Leaf blades to 100
mm long; to 4 mm wide. Spike-
lets 2-4 mm long; 1. 0-1.8 mm
wide. Inflorescence contracted,
dense, branches appressed to the
main axis, spikelets close to one
another, glands if present not
sticky; spikelets with rachilla fra-
gile, lemmas and/or paleas breaking up from the apex
downwards; lemma keels smooth or scaberulous; palea
keels with hairs 0.3-0. 6 mm long, which are exserted from
the lemma; anthers 2-3 (number variable in the same inflo-
rescence), 0.3-0. 4 mm long.

Flowering April. Sandy soil in disturbed areas such as
cultivated lands and roadsides. Infrequent (in FSA area).
Biome: Savanna. Throughout tropical Africa but mainly in
the south. Weed (of cultivated lands). Centre of a cluster
of closely related species including E. tenella , which has
the inflorescence open with the spikelets spreading. E.
ciliaris, which has the lemmas with long stiff hairs on the
keels, and E. viscosa , which has sticky glands on the inflo-
rescence.

Description: Chippindall & Crook 1976 (149), Clayton
et al. 1970-1982 (207). Voucher: Scheepers 630. PRECIS
code 9902860-00400.

Flowering February to June. On sandy soil in dolomite
areas and in disturbed places and old cultivated areas. Lo-
cally common. Biome: Savanna. Tropical Africa and India.
Similar toE. leersiiformis, which has an ovate-elliptic cary-
opsis.

Description: Chippindall & Crook 1976 (7), Stapf
1898-1900 (628), De Winter in Chippindall 1955 (160),
Clayton et al. 1970-1982 (209). Illustration: De Winter in
Chippindall 1955 (fig. 128). Voucher: Killick 1714.
PRECIS code 9902860-00600.

Eragrostis barbinodis Hack.

Perennial; stoloniferous and
tufted; culms to 1000 mm long,
geniculate to decumbent, often
rooting at the nodes. Leaf blades
1 00(— 1 50) mm long; 2-6 mm
wide. Spikelets to 7 mm long;

1.2-1. 5 mm wide. Culm nodes
(without branches at the nodes)
with long spreading hairs; inflo-
rescence open, branches spreading, lowest branches not
whorled; spikelets linear to oblong, rachilla persistent,
sometimes becoming fragile, lemmas and/or paleas break-
ing up from the base upwards; glumes 1/2-3/4 the length
of the lemmas directly above in the intact spikelet; palea
margins wide apart, not touching, keels a narrow line, sca-
berulous; anthers 3, 0. 8-1.0 mm long.

Flowering December to May. Red sandy loam, gritty to
sandy soils and black turf. Locally common. Biome: Savan-
na. Introduced to east Africa as a forage grass. Hybridizes
with £. rigidior, which is more erect and has the unbranch-
ed nodes glabrous.

Description: Stapf 1898-1900 (621), De Winter in Chip-
pindall 1955 (147). Voucher: Smook 4454. PRECIS code
9902860-00800.

Eragrostis barrelieri Dav.

Eragrostis aristata De Winter

Annual; tufted (erect and geni-
culate); to 750 mm tall. Leaf blad-
es to 200 mm long (possibly
longer); to 6.5 mm wide. Spike-
lets to 6 mm long; 1 .5-3 mm wide
(excluding awns). Spikelet rachil-
la persistent at first, becoming
fragile, lemmas and/or paleas
breaking up from the base up-
wards; lemma deeply two-lobed at the apex with the central
awn 0. 8-1.5 mm long, the lateral nerves shortly awned or
sometimes mucronate; palea keels scabrid; anthers 3,
0.6-0. 8 mm long; caryopsis oblong.

Flowering August, and April to May. Moist places. Lo-
cally common (Brandberg and Unjab mouth in Namibia).
Biome: Nama-Karoo and Desert. Endemic.

Description: De Winter 1961 (468). Voucher: Oliver,
Muller & Steenkamp 6688. PRECIS code 9902860-00500.

Eragrostis aspera (Jacq.) Nees

Grootpluimeragrostis.

Annual; tufted (erect); to 800
mm tall. Leaf blades to 300 mm
long; to 10 mm wide. Spikelets
3-10mm long; 1.0-1. 5 mm wide
Inflorescence open, branches
ascending at 45 degree angles,
bearded in the axils, pedicels long
and slender; spikelet rachilla fragile, lemmas and/or paleas
breaking up from the apex downwards; lemma obtuse to
truncate; palea keels scabrid; anthers 3, 0.2-0. 3 mm long;
caryopsis subglobose.

Annual; laxly tufted (erect to
geniculate); to 300 mm tall. Leaf
blades to 100 mm long; to 3.5 mm
wide. Spikelets 5-15 mm long;

1.5-1. 8 mm wide. Inflorescence
open, primary branches usually
not longer than 40 mm, spread-
ing, pedicels stout; spikelet ra-
chilla persistent, lemmas and/or
paleas breaking up from the base upwards; glumes unequal;
lemma obtuse; palea keels scabrid; anthers 3, 0.2-0. 3 mm
long; caryopsis oblong-elliptic.

Flowering December to January. On sand in disturbed
areas such as road verges and in gardens. Locally common.
Naturalized and invader from southern Europe. Biome; Sa-
vanna and Grassland. North Africa through the Middle East
to India, and southern Europe. Weed. Similar to E.
cilianensis, which has a subglobose caryopsis and wider
spikelets, and £. minor, which has subequal glumes.

Description: Hitchcock & Chase 1950 (156), De Winter
in Chippindall 1955 (158), Clayton et al. 1970-1982 (239).
Illustration: De Winter in Chippindall 1955 (fig. 125).
Voucher: De Winter 267. PRECIS code 9902860-00900.

Eragrostis bergiana (Kunth) Trin.

Kalkkweek.

Mat-forming perennial; long
rhizomatous and tufted; to 400
mm tall. Leaf blades 4-8 mm
long; 1 .0-1.5 mm wide. Spikelets
4-8 mm long; 2. 2-3. 8 mm wide.
Basal sheaths with dense, long,
woolly hairs; inflorescence

146

sparsely branched, spikelets densely clustered on the side
branches, which are slightly spreading from the main axis;
spikelets completely pallid or flushed with dark purple, ra-
chilla fragile, lemmas and/or paleas breaking up from the
apex downwards; lowest lemma broadly obtuse, lateral
nerves conspicuous; palea scaberulous; anthers 3, 1.5 mm
long.

Flowering September, December, and February.
Limestone soils, especially in pans and eroded places. Lo-
cally common. Biome: Nama-Karoo. Drought and frost
resistant pasture and erosion control (soil binder). Barely
distinguishable from E. truncata and a detailed study is
needed in this group. Resembles E. annulata, which has
glandular hairs with swollen tips and a broadly oblong to
broadly ovate caryopsis, and E. barrelieri and E. minor,
which have oblong caryopsis and narrower spikelets.

Description: Stapf 1898-1900 (624), De Winter in Chip-
pindall 1955 (178). Illustration: De Winter in Chippindall
1955 (fig. 149). Voucher: Smook 3923. PRECIS code
9902860-01000.

Eragrostis bicolor Nees

Fyn vleigras.

Perennial; usually hydrophyte,
or tufted (densely), or rhizomat-
ous (rhizome oblique); to 600 mm
tall. Leaf blades to 200 mm long;
to 1.5 mm wide. Spikelets to 8
mm long; 1-2 mm wide. Leaves
mainly basal, flat, usually glau-
cous; inflorescence open, lax, branches and spikelets
spreading, lowest branches solitary or 2-3, not whorled;
spikelets linear to narrowly oblong, rachilla persistent or
upper part fragile and florets breaking off in groups; glumes
1/2-2/3 the length of the lemmas above them in the intact
spikelet; lemmas deep purple or violet with a yellowish
apex, sometimes mostly yellowish but always bicoloured;
palea margins very close for the entire length, just touching
to overlapping at the apex, keels a thin line, entire, smooth
or scaberulous; anthers 3, 0.8-1. 2 mm long.

Flowering October to May. Often in brack areas, in wat-
er or on wet soil around seasonal pans, and in dry riverbeds.
Locally common. Biome: Savanna and Nama-Karoo.
Zimbabwe. Pasture (grazed by game).

Description: Stapf 1898-1900 (605), De Winter in Chip-
pindall 1955 (141). Illustration: De Winter in Chippindall
1955 (fig. 109). Voucher: Smook 3385, Bryant 645a.
PRECIS code 9902860-0 1100.

Fig. 86. Eragrostis biflora

Eragrostis biflora Hack, ex Schinz

Fig. 86.

Annual; tufted; to 700 mm
tall. Leaf blades to 300 mm long;
to 8 mm wide. Spikelets 1.5-2. 5
mm long; 0.5-1. 2 mm wide. In-
florescence open, delicate, much
branched, pedicels long and
slender; spikelets narrowly el-
liptic when young to broadly
ovate when mature, with 1 — 2(— 3 )
florets, rachilla persistent, becoming fragile in the upper
part, lemmas and/or paleas breaking up from the base up-
wards; lemma 1.0-1. 5 mm long; palea keels glabrous to
scabrid; anthers 3, 0.2-0. 3 mm long.

Flowering September to May. Moist disturbed areas,
especially under trees. Locally common. Biome: Savanna,
Grassland, and Nama-Karoo. Endemic. Weed. Can be
confused with the genus Sporobolus, which has 1 -flowered
spikelets, and resembles E. habrantha, which is perennial.

Description: Stapf 1898-1900 (610), De Winter in Chip-
pindall 1955 (151). Illustration: De Winter in Chippindall
1955 (fig. 118). Voucher: Ellis 2617. PRECIS code
9902860-01200.

Eragrostis brizantha Nees

Annual; tufted (erect to geni-
culate); to 500 mm tall. Leaf blad-
es to 150 mm long; to 4.5 mm
wide. Spikelets to 5 mm long;

2. 2-4.0 mm wide. Basal sheaths
glabrous or with long scattered
hairs; inflorescence with spikelets
densely crowded on slightly
spreading to spreading branches;
spikelets ovate to very broadly oblong, with rachilla fragile
and breaking up from the apex downwards; upper glume
2/3— 3/4 the length of the lemma above in the intact spikelet;
lemma dull, of the same texture throughout, often flushed
with purple, broadly obtuse (in profile), lateral nerves dis-
tinct with small glandular dots; palea keels entire, glabrous,
with small glandular dots; anthers 3, 1.0-1. 2 mm long;
caryopsis obovate.

Flowering February to May (also July to November).
Sandy and calcareous soils around rivers and in disturbed
areas. Locally common. Biome: Savanna and Nama-Karoo.
Endemic. Resembles E. echinochloidea, which has the low-
er glume acuminate and the palea keels very broad and
protruding in the lower part and ending in a deep notch on
top.

147

Description: Stapf 1898-1900 (626), De Winter in Chip-
pindall 1955 (174). Illustration: De Winter in Chippindall
1955 (fig. 146). Voucher: Giess & Mueller 12268. PRECIS
code 9902860-01300.

Eragrostis caesia Stapf

Perennial; densely tufted;

450-600 mm tall. Leaf blades to
200 mm long; to 3 mm wide.

Spikelets 4-7 mm long; 1.5-2. 4
mm wide. Inflorescence dense
and contracted, branches usually
appressed to the main axis, occas-
ionally spreading; spikelets
tardily breaking up between the
florets; lemma with distinct elongated black patches along
the lateral nerves; palea keels with hairs less than 0.1 mm
long; anthers 3, 1.0-1. 3 mm long.

Flowering November to June. Moist areas on shallow
soil, Cave sandstone, and seepage areas in mountainous
grassland. Locally common. Biome: Grassland and Afro-
montane. Zimbabwe. Eagerly grazed pasture (remains
green in winter).

Description: Stapf 1898-1900 (599), De Winter in Chip-
pindall 1955 (141). Voucher: Smook 1060, Schmitz 4167A.
PRECIS code 9902860-01400.

Eragrostis capensis (Thunb.) Trin.

Fig. 87. PI. 77.

Hartjie-eragrostis.

Perennial; tufted; to 900 mm
tall. Leaf blades 70-350 mm
long; 2-5 mm wide. Spikelets
3.5-15.0 mm long; 3-7 mm wide.

Basal sheaths glabrous to hairy
but not woolly-hairy at the base;
leaves mainly basal; inflores-
cence sparsely branched or unbranched, spikelets appressed
to the main axis or branches; spikelets plump, with the op-
posite row of florets overlapping and closely packed with
the rachilla not visible, rachilla persistent, lemmas and/or
paleas breaking up from the base upwards; lemma obtuse
to subobtuse, dull and granular, greenish to greenish brown,
strongly flushed with purple, lateral nerves distinct; palea
narrowly obovate, keels entire, membranous to
subcartilaginous between the keel and the margins, margins
nearly touching to touching in the lower parts to widely
separated in the upper parts; anthers 3,1-2 mm long; cary-
opsis elliptic.

Flowering September to April. Sandy to clayey soils in
moist areas on slopes, rocky and disturbed places. Widely
common (especially after fire). Biome: Savanna, Grassland,
and Fynbos. Northwards to Zaire, Kenya and Tanzania,
Madagascar and Thailand. Only useful as early spring pas-
ture. Similar to E. cimicina, which has smooth and shiny
lemmas and the palea margins nearly touching to touching
along the entire length. Sometimes confused with E.
superba, in which the spikelets are strongly flattened and
the entire spikelet disarticulates as a unit below the glumes.

Description: Chippindall & Crook 1976 (51), Clayton et
al. 1970-1982 (221). Illustration: De Winter in Chippindall
1955 (fig. 140). Voucher: Kluge 1119, Balsinhas 3201,
Smook 2077. PRECIS code 9902860-01500.

Eragrostis chapelieri (Kunth) Nees

Bruinsaadgras.

Perennial; erect and densely
tufted; to 900 mm tall. Leaf blad-
es to 400 mm long; to 5 mm wide.

Spikelets 6-24 mm long; 2.0-2. 5
mm wide. Basal sheaths glabrous
to obscurely hairy at the base; in-
florescence narrow, dense, bran-

ches appressed to the main axis and overlapping but often
distant in the lower part; spikelets usually reddish brown,
rachilla persistent, lemmas and/or paleas breaking up from
the base upwards; lemma acute-acuminate, lateral nerves
distinct but not raised; palea nearly the same width between
the keel and the margins throughout, narrowing upwards
and running into the keel near the apex, margins not touch-
ing, at least not in the upper part, keels entire, broad and
flattened; anthers 2, 0.3-0. 7 mm long; caryopsis broadly
ellipsoid.

148

Flowering March to August. Poor sandy soils or occas-
ionally on clays, in disturbed areas such as old lands and
pathsides. Rare (in southern Africa). Locally common.
Biome: Savanna. Tropical Africa to Sudan and in
Madagascar. Weed (in cultivated lands). Similar to E.
patens, which is annual, and E. elatior, which has 2-6 dark
olive green spikelets in clusters distant from one another on
the main axis, and E. inamoena , which has the palea mar-
gins slightly apart to touching along the entire length and
spikelets greyish-green, often flushed with purple.

Description: Chippindall & Crook 1976 (157), Stapf
1898-1900 (614), Clayton et al. 1970-1982 (225).
Illustration: De Winter in Chippindall 1955 (fig. 142).
Voucher: Codd 5456. PRECIS code 9902860-01600.

Eragrostis chloromelas Steud.

Perennial; tufted; to 800 mm
tall. Leaf blades to 300 mm long;
setaceous. Spikelets 4-6 mm
long; 1. 0-1.5 mm wide. Culms
strongly compacted and not easily
separated individually, nodes gla-
brous; basal sheaths glabrous or
obscurely hairy at the very base;
leaf blades tapering to very long
filiform tips, very curly especially when older; inflores-
cence open and much branched, branches and pedicels
spreading, long hairs in the axils, lowest branches usually
solitary, never whorled; spikelets narrowly obovate to ob-
long, pale greenish-grey to dark green, rachilla persistent
in the lower part, usually becoming fragile in the upper part,
lemmas and/or paleas breaking up from the base upwards;
glumes translucent, usually smooth, longer than 1/3 the
length of the lemmas directly above; palea margins nearly
touching to touching along the entire length of the palea;
anthers 3, 0.6-0. 8 mm long; caryopsis ellipsoid.

Flowering December to May. Hillslopes, rocky ridges,
in disturbed areas on sandy soil, loam and dolomite. Com-
mon. Biome: Savanna and Grassland. Endemic. Reasonably
palatable pasture. Intergrades with forms off. curvula.

Description: Stapf 1898-1900 (602), Hitchcock &
Chase 1950 (168), De Winter in Chippindall 1955 (145).
Illustration: De Winter in Chippindall 1955 (fig. 112).
Voucher: Smook 5825, De Winter 626. PRECIS code
9902860-01700.

Eragrostis cilianensis (All.) F.T. Hubb.

Stinkgras.

Annual; loosely tufted (often
geniculate); to 900 mm tall. Leaf
blades to 250 mm long; to 10 mm
wide. Spikelets 3-20 mm long;

1. 5-4.0 mm wide. Basal sheaths
glabrous, if hairy not densely
woolly-hairy; inflorescence with
side branches usually 40 mm or shorter, spreading to ap-
pressed near the apex of the inflorescence, branches and
pedicels stout; spikelets narrowly ovate to narrowly oblong,
rachilla persistent, lemmas and/or paleas breaking up from
the base upwards; lemma narrowly to broadly obtuse in
profile, 1.7-2. 8 mm long; palea keels entire, scabrid; an-
thers 3, 0.2-0. 3 mm long; caryopsis subglobose.

Flowering October to June. Sandy soils, often in moist
places, also disturbed areas such as cultivated lands,
pathsides and overgrazed places. Locally common. Biome:
Grassland, Savanna, and Nama-Karoo. Throughout Africa
and in tropical and warm temperate regions of the Old
World. Introduced to the New World. Ruderal weed.
Resembles E. annulata, which has glandular hairs with
swollen tips and broadly oblong to broadly ovate caryopsis,
and E. banelieri and E. minor, which have the caryopsis
oblong and the spikelets somewhat narrower.

Description: Chippindall & Crook 1976 (8), Hitchcock
& Chase 1950 (154), Clayton et al. 1970-1982 (232).

Illustration: De Winter in Chippindall 1955 (fig. 126), Clay-
ton et al. 1970-1982. Voucher: Ross 1916. PRECIS code
9902860-01800.

Eragrostis ciliaris (L.) R. Br.

Woolly love grass.

Annual; tufted (erect); to 600
mm tall. Leaf blades to 120 mm
long; to 5 mm wide. Spikelets
2.0— 4.5 mm long; 1.5-2. 2 mm
wide. Inflorescence contracted,
often interrupted, spikelets dense-
ly clustered; spikelets usually
flushed with purple, rachilla fragile, breaking up from the
apex downwards; lemma keels (at least the upper ones in
the spikelets) with stiff hairs 0. 2-0.4 mm long, especially
towards the base; palea keels with hairs 0.5-0. 7 mm long,
which are exserted from the lemma; anthers 2, 0.20-0.25
mm long.

Flowering throughout the year. Moist sandy soils in dis-
turbed places such as human habitation, cultivated lands,
overgrazed and trodden places. Locally common. Biome:
Savanna. Extending through tropical Africa, Arabia and
Mascarene Islands to India, also tropical America. Pasture
(eaten by stock in Mozambique), or weed (ruderal). Belongs
to a group of related species, including E. tenella, which
has an open inflorescence with spreading branches, E.
viscosa, which has sticky, glandular patches on the inflores-
cence, and E. arenicola, which has the lemma keel smooth
or scabrid.

Description: Chippindall & Crook 1976 (149), Stapf
1898-1900 (629), Hitchcock & Chase 1950 (145), Clayton
et al. 1970—1982 (204). Illustration: De Winter in Chippin-
dall 1955 (fig. 153). Voucher: Smook 5737. PRECIS code
9902860-01900.

Eragrostis cimicina Launert

Perennial; occasionally rhizo-
matous and tufted (densely); to
1 500 mm tall. Leaf blades 50-320
mm long; to 5 mm wide. Spike-
lets 3-6(-9) mm long; 3-5 mm
wide. Basal sheaths glabrous or
hairy at the base but not woolly-
hairy; inflorescence open: spike-
lets with the opposite rows of flo-
rets closely packed and overlapping at the base, rachilla not
visible, spikelet rachilla persistent, lemmas and/or paleas
breaking up from the base upwards; lemma smooth and
shiny, obtuse to subobtuse, lateral nerves distinct, median
keel with long cilia on the lower 1/3; palea elliptic, thickly
cartilaginous, especially between the keel and the margins,
margins nearly touching along the entire length, keels
smooth to scaberulous; anthers 3, 1 .5-2.0 mm long; caryop-
sis broadly oblong with a deep pit dorsally.

Flowering January to March. Sandy loam on flood-
plains. Infrequent. Biome: Savanna Southern Angola,
Zambia and Zimbabwe. Similar to E. capensis, which has
the inflorescence contracted rather than open and the spike-
lets dull and granular.

Description: Launert 1970 (221). Voucher: De Winter
9204. PRECIS code 9902860-02000.

Eragrostis congesta Oliv.

Weak perennial; tufted (erect
or geniculate); to 800 mm tall.

Leaf blades 100-200 mm long;

3-4 mm wide. Spikelets 3-10 mm
long; 1. 2-2.0 mm wide. Inflores-
cences consisting of 2-8 dense,
distant clusters; spikelets elliptic
to narrowly oblong, breaking up
from the base upwards, rachilla

149

breaking off above the glumes soon after the lemmas begin
to fall; palea keels scaberulous; anthers 3, 0. 3-0.4 mm long.

Flowering May to July. Moist areas or disturbed places,
such as roadsides. Infrequent. To east Africa. Said to be
locally common in Zimbabwe. Up to this time, only a few
specimens have been collected from one small area in the
FSA region.

Description: Chippindall & Crook 1976 (156), Clayton
et al. 1970-1982 (224). Illustration: De Winter in Chippin-
dall 1955 (fig. 106). Voucher: Strey 10947. PRECIS code
9902860-02100.

Eragrostis crassinervis Hack.

Perennial; occasionally hydro-
phyte and stoloniferous, tufted
(densely); to 600 mm tall. Leaf
blades to 1 00 mm long; to 2.5 mm
wide. Spikelets 4.5-8.0(-15.0)
mm long; 2-3 mm wide. Inflores-
cence narrow, unbranched or only
sparsely branched, spikelets ap-
pressed to the main axis orbran-
ches, pedicels short and stout; spikelets light green to straw
coloured, or purple or at least flushed with purple, rachilla
persistent, lemmas and/or paleas breaking up from the base
upwards; lemma usually truncate to concave between the
keel and the lateral nerves, which are raised and excurrent
into minute mucros and have small glandular pits; palea
moderately wide between the margins and the keels, mar-
gins not touching, keels entire, flattened, equally broad for
the entire length; anthers 3, 0.4-0. 6 mm long; caryopsis
subglobose.

Flowering January to April. Moist places such as river
beds and vleis, sometimes on brackish soils. Infrequent.
Biome: Savanna and Nama-Karoo. Zimbabwe. Resembles
E. walteri, which has the palea keels very broad in the lower
2/3, narrowing sharply to the apex where they are excurrent
into a small, soft mucro.

Description: De Winter in Chippindall 1955 (178).
Voucher: Giess 8108, Giess 9566. PRECIS code 9902860-
02200.

Eragrostis curvula (Schrad.) Nees

(=£. robusta Stent) 8.

Oulandsgras, weeping love
grass.

Wiry perennial; erect and
densely tufted; to 1200 mm tall.

Leaf blades to 500 mm long; to 3
mm wide. Spikelets 4-10 mm
long; 1.0-1 .5 mm wide. Plants variable; culms unbranched,
not easily compressed, nodes glabrous; basal sheaths dense-
ly hairy for quite a way up from the base, with long hairs
in the deep furrows between the prominent, squarish ridges
formed by the closely packed nerves; leaf blades rolled or
flat, appearing setaceous, with long tapering, filiform tips;
inflorescence much branched, variable, being open and
spreading, or contracted with the branches appressed to the
main axis, the lowest branches whorled or not whorled, the
pedicels smooth or with prickles distant from one another,
the spikelets appressed to the branches; spikelets linear to
oblong, rachilla persistent or upper part often fragile, lem-
mas and/or paleas breaking up from the base upwards;
glumes translucent, of variable length but longer than 1/3
the length of the lemmas above in the intact spikelet,
smooth or scaberulous at the apex and along the keels; lem-
ma pale green to dark green to greyish; palea margins meet-
ing or overlapping for the entire length, keels a thin narrow
line or palea apparently only folded; anthers 3, 0.6-1. 0 mm
long; caryopsis ellipsoid.

Flowering August to June. In high rainfall areas on
sandy or acid to loamy soils, often in disturbed or badly

Fig. 85. PI. 78.

managed areas. Biome; Fynbos, Savanna, Grassland,
Nama-Karoo, and Succulent Karoo. Northwards to east
Africa, introduced throughout the tropics mainly as a
fodder. Widely cultivated pasture, or erosion control
(rehabilitation of roadverges and ground cover), or weed
(where it has escaped from cultivation and in some areas
of the world where it has been introduced). A very variable
grass, with several ploidy levels, which appears to grade in-
to other species such as E. chloromelas , E. barbinodis, E.
caesia,E. lehmanniana,E. planiculmis and£. rigidior. This
species is currently under investigation.

Description; Chippindall & Crook 1976 (17), Stapf
1 898-1900 (599), Hitchcock & Chase 1950 (168), De Win-
ter in Chippindall 1955 (142), Clayton et al. 1970-1982
(243). Illustration: De Winter in Chippindall 1955 (fig. 1 10
& 111). Voucher: Smook 3839, Smook 3254, Retief 129,
S. van Wyk 1 14. PRECIS code 9902860-02300.

Eragrostis cylindriflora Hochst.

(=£. horizontalis Peter) 9.

Annual; loosely tufted (usual-
ly erect); to 800 mm tall. Leaf
blades 30-150 mm long; 2^1 mm
wide. Spikelets 3-8 mm long;

0.5-1. 5 mm wide. Leaf sheaths
nearly always dotted with oblong
glands; inflorescences with the
lowest branches whorled; spikelet with rachilla persistent
in the lower part and fragile above, lemmas and/or paleas
breaking up from the base upwards; lower glume 4/5 as long
as to longer than the lemma above in the intact spikelet;
lowest lemma 1.5-1. 7 mm long, chartaceous to membran-
ous, broadly elliptic, obtuse to subacute, lateral nerves ob-
scure; palea keels minutely scaberulous; anthers 3, 0. 8-1.0
mm long; caryopsis ellipsoid.

Llowering January to August. Sand, clayey loam or
black turf in river beds, depressions and in disturbed areas
such as overgrazed places. Locally common. Biome: Savan-
na. Tropical Africa. Resembles £. glandidosipedata , which
has lemmas with distinct lateral nerves, and £.
omahekensis, which has the lowest lemma 1.8-2. 2 mm
long, and £. trichophora, a perennial species. Tends to in-
tergrade with £. porosa, which has obovate-elliptic lem-
mas, with truncate to broadly rounded apices.

Description: De Winter in Chippindall 1955 ( 1 50), Clay-
ton et al. 1970-1982 (239). Illustration: Clayton et al.
1970-1982 (fig. 66). Voucher: De Winter 2731, De Winter
2284. PRECIS code 9902860-02400.

Eragrostis dinteri Stapf

Annual; tufted (erect; occas-
ionally geniculate and rooting at
the nodes); to 500 mm tall. Leaf
blades to 150 mm long; to 8 mm
wide. Spikelets 7-17 mm long;

3-5 mm wide. Basal sheaths gla-
brous or with bulbous-based hairs
on the margins; spikelet rachilla
fragile in upper part and subper-
sistent below, breaking up from the apex downwards; upper
glume 2/3— 4/5 as long as the lemma above in the intact
spikelet; lemmas of the same texture throughout, acumin-
ate, usually with a median awn; palea keels entire, scabrid;
anthers 3, 1.2-1. 5 mm long; caryopsis subglobose.

Flowering February to June. Deep, red sandy soils and
disturbed areas. Locally common. Biome: Savanna.
Angola. Similar to £. rogersii , which has acute lemmas
with or without a short mucro. A strong smell has been
recorded for the plant.

Description: De Winter in Chippindall 1955 (172).
Voucher: Smook 5220; Biegel, Muller & Gibbs Russell
4999. PRECIS code 9902860-02700.

150

Eragrostis echinochloidea Stapf

Tick grass, bosluisgras.

Perennial; tufted (erect or gen-
iculate); to 900 mm tall. Leaf
blades to 500 mm long; to 6 mm
wide. Spikelets 2-6 mm long;

2.2-3. 5 mm wide. Basal sheaths
glabrous or long-hairy at the
extreme base only; inflorescence
sparsely branched, spikelets densely congested and secund
on the branches, pedicels short and stout; spikelet rachilla
very fragile, easily disarticulating between the florets from
the apex downwards; lower glume acuminate; palea acute,
keels scaberulous, with the lower portion very broad and
projecting from the upper portion, top of broad portion ends
in deep notch or tooth; anthers 3, 0. 5-0.7 mm long; caryop-
sis elliptic.

Flowering November to May. Prefers shallow moist
calcrete soils especially around pans, also disturbed sandy
places such as cultivated lands. Infrequent to locally com-
mon. Biome: Savanna and Nama-Karoo. Endemic; has been
naturalized in Arizona (USA). Drought resistant palatable
pasture (especially when green), or indicator (denuded
veld). Resembles E. brizantha, which has glumes obtuse to
subacute in profile and palea keels entire.

Description: Stapf 1898-1900 (627), De Winter in Chip-
pindall 1955 (174). Illustration: Muller 1984 (fig. 74), De
Winter in Chippindall 1955 (fig. 145). Voucher: De Winter
& Wiss 4427, Relief 1530. PRECIS code 9902860-02800.

Eragrostis elatior Stapf

Throughout tropical Africa and India. Resembles the more
slender forms of E. membranacea, which has anthers 3,
0.8-1 .3 mm long.

Description: Launert 1970 (160:96), Stapf 1898-1900
(617), Clayton et al. 1970-1982 (217). Voucher:

Schweickerdt 2194. PRECIS code 9902860-03000.

Eragrostis glandulosipedata De Winter

Annual; tufted (geniculate); to
1000 mm tall. Leaf blades to 300
mm long; to 5.5 mm wide. Spike-
lets 3-5 mm long; 1.0- 1.8 mm
wide. Inflorescence with the low-
est branches whorled; spikelets
with the rachilla persistent, some-
times fragile in the upper part,
lemmas and/or paleas breaking up
from the base upwards; lowest lemma broadly elliptic to
broadly oblong-ovate, obtuse to subacute, 1 .5-1 .7 mm long,
lateral nerves conspicuous; palea keels scaberulous; anthers
3, 0.8-1.0 mm long; caryopsis oblong to broadly oblong.

Flowering February to June. Sand, gravel, turf and
calcareous soils and in areas of high moisture and dis-
turbance. Locally common. De Winter (1961, Bothalia 7)
cites a single specimen, Bogan 3119, from Kenya. Pasture
(probably good as fodder). Resembles E. cylindriflora,
which has lemmas with obscure lateral nerves, and E.
omahekensis , which has the lowest lemma 1.8-2. 2 mm
long.

Description: De Winter 1961 (469). Voucher: De
Winter 2290, Giess, Volk & Bleissner 6413. PRECIS code
9902860-03100.

Perennial; rhizomatous and
tufted (densely); to 500 mm tali.

Leaf blades to 200 mm long; to
3.5 mm wide. Spikelets 5-8 mm
long; 2.0-2. 5 mm wide. Leaves
mainly cauline; inflorescence
slender, 50-200 mm long,
unbranched or sparsely branched,
branches and spikelets appressed
to the main axis, spikelets solitary or in distant clusters of

2- 6 spikelets along the main axis, not overlapping each
other; spikelets dark olive green, lemmas hardly diminish-
ing in length towards the apex, rachilla subpersistent, fra-
gile in the upper part; glumes cartilaginous, upper glume
lanceolate to oblong; lemma acute to subacute, lateral
nerves distinct, lowest lemma 2. 5-3.0 mm long; palea mar-
gins nearly touching or touching for the entire length, keels
a raised ridge, scabrid; anthers 3, 0.6—1 .0 mm long; caryop-
sis oblong-elliptic.

Flowering December and March. Rocky banks of rivers
and periodically inudated areas. Locally common (coastal
areas of the southwestern Cape). Biome: Fynbos. Endemic.
Resembles E. chapelieri , which has many spikelets and the
spikelets and branches overlapping except the lower
branches which are sometimes distant.

Description: Stapf 1898-1900 (617), De Winter in De
Winter in Chippindall 1955 (161). Voucher: Kruger 1177.
PRECIS code 9902860-02900.

Eragrostis gangetica (Roxb.) Steud.

Annual; tufted (erect); to 800
mm tall. Leaf blades to 600 mm
long; 1-3 mm wide. Spikelets

3- 10 mm long; 1 .0-1 .7 mm wide.

Inflorescence usually open, pedi-
cels long and slender; spikelets
with rachilla persistent, lemmas
and/or paleas breaking up from
the base upwards; palea keels sca-
berulous; anthers 2, 0. 1-0.3 mm long; caryopsis

subglobose.

Flowering January, April, and May. Open areas near
marshes or temporary vleis. Infrequent. Biome: Savanna.

Eragrostis gummiflua Nees

Gum grass, gomgras.

Perennial; densely tufted; to
900 mm tall. Leaf blades to 500
mm long; to 4.5 mm wide. Spike-
lets 2. 5^1.0 mm long; 1.0-1. 8
mm wide. Culms with sticky
glandular patches below the nod-
es and on the leaf sheaths below
the collar, noticeable because soil particles adhere to the
patches; spikelet rachilla fragile, lemmas and/or paleas
breaking up from the apex downwards; palea keels scaberu-
lous; anthers 3, 0. 5-0.8 mm long; caryopsis oblong-elliptic.

Flowering September, and November to April. Locally
common. Biome: Fynbos, Savanna, and Grassland.

Mozambique. Domestic use (brooms in Lesotho).

Description: Chippindall & Crook 1976 (152), Stapl
1898-1900 (629), De Winter in Chippindall 1955 (178).
Illustration: De Winter in Chippindall 1955 (fig. 152).
Voucher: Smook 3009. PRECIS code 9902860-03200.

Eragrostis habrantha Rendle

Perennial; tufted (erect); to
1000 mm tall. Leaf blades to 100
mm long; to 5 mm wide. Spike-
lets 1-2 mm long; to 1.2 mm
wide. Inflorescence linear,
delicate, much branched, branch-
es fine and flexible; mature spike-
lets very broadly ovate, 2-3-
flowered, rachilla subpersistent,
fragile in the upper part, lowest floret persistent; palea keels
smooth to scaberulous; anthers 3, 0.6-0. 8 mm long.

Flowering January to May. Sandy and clayey soils in
open damp areas along rivers and around vleis. Locally
common (limited to a few sites in the FSA area, becoming
more widespread northwards). Biome: Savanna and Grass-
land. Central tropical Africa. Resembles E. micrantha,
which has a linear to oblong spikelet, and the inflorescence
resembles E. biflora, which is an annual plant.

Description: Chippindall & Crook 1976 (158), De Win-

151

ter in Chippindall 1955 (154). Voucher: Gertenbach 7027,
Volk A39. PRECIS code 9902860-03300.

Eragrostis heteromera Stapf

Moderately slender perennial;
tufted (erect); to 1000 mm tall.

Leaf blades to 150 mm long; 2^1
mm wide. Spikelets 4-9 mm
long; to 1.5 mm wide. Culms not
densely compacted at the base
and easily separable, nodes gla-
brous; basal sheaths glabrous or
obscurely hairy; leaf blades long-
tapering at the tip; inflorescence open, moderately branch-
ed, lowest branches 1-8, whorled or not whorled, spikelets
appressed to the branches, pedicels (except terminal ones)
shorter than or as long as the spikelets; spikelets linear to
oblong, rachilla persistent, lemmas and/or paleas breaking
up from the base upwards; glumes translucent, smooth or
scaberulous on the keels and apex, 1/3-3/4 the length of the
lemmas directly above in the intact spikelet; lemma purple
to violet to green, often with yellow especially near the
apex, nerves usually prominent, strongly keeled with keel
prominent along the entire length; palea margins meeting
or overlapping along the entire length, keels distinct but a
narrow line; anthers 3, 0. 8-1.0 mm long; caryopsis narrow-
ly oblong.

Flowering December to May. On moist sand or black
clay in depressions, seasonal pan margins and in disturbed
areas. Locally common. Biome: Savanna and Grassland.
Northwards to Ethiopia. Pasture (of average fodder value).
Similar to£. rotifer, which has densely hairy basal sheaths.

Description: Chippindall & Crook 1976 (171), Stapf
1898-1900 (610), De Winter in Chippindall 1955 (156),
Clayton et al. 1970-1982 (215). Illustration: De Winter in
Chippindall 1955 (fig. 123). Voucher: De Winter & Codd
498. PRECIS code 9902860-03400.

Eragrostis hierniana Rendle

(=£. uniglumis Hack.) 9.

Perennial; tufted (sometimes
geniculate); to 1000 mm tall. Leaf
blades 100-250 mm long; 2-4
mm wide. Spikelets 5-12 mm
long; 1.0-2. 5 mm wide. Spikelets
with the rachilla persistent, the
lemmas and/or paleas breaking up
from the base upwards; lemma glabrous or with hairs less
than 0.3 mm long; palea keels with hairs 0. 3-2.0 mm long,
extending beyond the lemma; anthers 3, 0.8-1. 2 mm long.

Flowering August to April. Moist sandy soils in hollows
on hills, along rivers and in disturbed areas such as old
cultivated lands. Locally common (in Natal, only in Muzi
swamps). Biome: Savanna. Northwards to Tanzania.
Resembles E. lappula, which has hairs longer than 0.3 mm
on the lateral nerves of the lemma, and E. inamoena, which
has scabrid palea keels.

Description: De Winter in Chippindall 1955 (164), Clay-
ton et al. 1970-1982 (213). Voucher: Godfrey & Acocks
SH 1601. PRECIS code 9902860-03450.

Eragrostis homomalla Nees

(-E. hygrophila C.E. Hubb.

& Schweick.) 1 .

Reengrassie.

Annual; tufted (erect to de-
cumbent); to 500 mm tall. Leaf
blades 20-100 mm long; 1^1 mm
wide. Spikelets 2-7 mm long;

0. 7-1.0 mm wide. Inflorescence heavy, branches spreading
or appressed to the main axis, rigid, spikelets irregularly
and densely condensed along the primary branches, pedi-

cels stout; spikelets linear to oblong, lemmas on the same
side of the rachilla distinctly overlapping the lemma above,
rachilla subpersistent, lemmas and/or paleas breaking up
from the base upwards; lower glume to 1/3 the length of
the lemma above in the intact spikelet; lemma ovate-
elliptic, 1.0- 1.5 mm long; palea keels smooth to scabrid;
anthers 3, 0.3 mm long; caryopsis ellipsoid.

Flowering January to May. Moist, sandy loam or clay
in brackish depressions or seasonally wet pans. Locally
common. Biome: Savanna and Nama-Karoo. Endemic,
except for a single specimen, Estes 29, recorded from
Kenya.

Description: Stapf 1898-1900 (631), De Winter in Chip-
pindall 1955 (153), Clayton et al. 1970-1982 (214).
Illustration: De Winter in Chippindall 1955 (fig. 121).
Voucher: Smook & Gibbs Russell 2454a, Leistner 1762.
PRECIS code 9902860-03500.

Eragrostis inamoena K. Schum.

( -E . atrovirens auctt., non
Trin. ex Steud.).

Perennial; loosely tufted and
rhizomatous (rhizome short and
oblique to long and branched); to
1000mm tall. Leaf blades 40-250
mm long; 2— A mm wide. Spike-
lets 5-20 mm long; 2. 0-3. 5 mm
wide. Basal sheaths glabrous at the base; leaves mainly
cauline; inflorescence 50-200 mm long, variable, grading
from open with spreading branches to contracted with bran-
ches overlapping one another and appressed to the main
axis, pedicels (except the terminal ones) shorter than, or as
long as the spikelets; spikelets dull, greyish-green to dark
green, frequently Hushed with purple, 7-40-flowered, lem-
mas hardly becoming shorter towards the apex, rachilla per-
sistent, lemmas and/or paleas breaking up from the base up-
wards; upper glume acute; lowest lemma 1 .6-2.5 mm long,
acute, lateral nerves distinct; palea narrowly obovate, mem-
branous except for the keel, apex truncate to obtuse, mar-
gins nearly touching to touching along the entire length,
keels narrow, Hat, but distinct, entire, scaberulous; anthers
3, 0.6-1. 3 mm long; caryopsis narrowly elliptic.

Flowering November to May. Sandy to organically rich
soils on seasonally flooded areas and marshy places. Local-
ly common. Biome: Savanna. Northwards into east Africa.
Resembles E. lappula and£. hierniana, which have the pa-
lea keels with hairs longer than 0.3 mm, and E. chapelieri,
which has the palea margins not touching and reddish
brown spikelets.

Description: Chippindall & Crook 1976( 154), De Winter
in Chippindall 1955 ( 163), Clayton et al. 1970-1982 (218).
Illustration: De Winter in Chippindall 1955 (fig. 132).
Voucher: Smook 5708, Smook 5710. PRECIS code
9902860-03550.

Eragrostis jeffreysii Hack,

Geelhoutpluimgras.

Robust perennial; tufted; to
2000 mm tall. Leaf blades to 1 000
mm long; to 10 mm wide. Spike-
lets 5-8 mm long; 1.0-1. 5 mm
wide. Culms easily compressed;
basal sheaths hairy at the base,
leaves flat and mainly basal; in-
florescence open, branches spreading, lowest branches
whorled, branches and pedicels yellow, scabrid, not thickly
covered with prickles; spikelets linear to oblong, rachilla
persistent in lower portion, often fragile in upper part, lem-
mas and/or paleas breaking up from the base upwards;
glumes 1/2-2/3 the length of the lemmas directly above in
the intact spikelet and the lower glume not wide enough to
cover the lemma; lemma pallid to pale greyish-green; palea
margins almost touching the entire length, touching and
overlapping at the apex, keels entire, obscure, smooth to

152

scaberulous; anthers 3, 0.8 mm long.

Flowering February and June. Sandy moist areas. Infre-
quent. Biome: Savanna. To Zimbabwe. Close to £. curvula,
which has culms that are not easily compressed and leaves
to 3 mm wide, and a detailed study is needed in this group.

Description: Launert 1970 (160:107), Hackel 1909
Feddes Rep. 6 (322). Voucher: Volk 1019. PRECIS code
9902860-03570.

Eragrostis kingesii De Winter

Annual; tufted (erect to de-
cumbent); to 100 mm tall. Leaf
blades to 20 mm long; to 3 mm
wide. Spikelets to 5 mm long;

1.0-1. 5 mm wide. Basal leaf
sheaths glabrous or with a few
scattered, bulbous-based hairs
near the apex; leaf blade margins
with raised glands, midrib with
glandular dots; inflorescence moderately to densely con-
tracted, side branches less than 40 mm long, rigid, pedicels
stout; spikelets lanceolate, rachilla persistent, lemmas
and/or paleas breaking up from the base upwards; lower
glume 1/2-2/3 the length of the lemma above in the intact
spikelet, strongly keeled; lemma acute, 1.5-1. 8 mm long;
palea keels scabrid; anthers 3, 0. 1-0.3 mm long; caryopsis
oblong-elliptic.

Flowering February to May. Disturbed soils along road-
sides and in farmyards. Locally common (Luderitz area).
Biome: Desert. Endemic. Weed. Similar to E. procumbens,
which has spikelets 2.0-2. 5 mm wide, and E. pygmaea ,
which has many bulbous-based hairs on the lower leaf
sheaths and a subglobose caryopsis.

Description: De Winter 1961 (470). Voucher: De Winter
& Giess 6083. PRECIS code 9902860-03600.

Eragrostis laevissima Hack.

Perennial, or annual (occas-
ionally); densely tufted (erect or
occasionally geniculate), or rhi-
zomatous (rhizome oblique); to
800 mm tall. Leaf blades to 150
mm long; 1-2 mm wide. Spike-
lets 2-8 mm long; 1. 3-2.0 mm
wide. Inflorescence open, spike-
lets condensed on the branches,
lowest branches 1-2, not whorled, pedicels short and stout;
spikelets oblong to ovate, rachilla persistent, lemmas and/or
paleas breaking up from the base upwards; glumes 1/2-2/3
the length of the lemmas directly above; lemma with lateral
nerves with small round glands, which sometimes give a
lumpy appearance to the nerves and keel; palea margins
touching along the entire length, keels a thin line, glandular
and scabrid; anthers 3. 0.6-1. 2 mm long.

Flowering February to March. Sandy and brackish
calcareous soils around edges of pans and vleis. Infrequent.
Biome: Savanna. Endemic. Resembles forms of£. sabinae,
which have lemmas with the lateral nerves eglandular.

Description: Launert 1970 (160:101), De Winter in
Chippindall 1955 (152). Voucher: Giess & Mueller 13961,
Giess & Loutit 14127, De Winter 2940. PRECIS code
9902860-03700.

Eragrostis lamprospicula De Winter

Perennial; tufted (geniculate);
to 800 mm tall. Leaf blades to 100
mm long; to 3 mm wide. Spike-
lets 5-15 mm long; 2. 2-3. 5 mm
wide. Plants not sprawling; basal
sheaths glabrous; leaves not
forming a dense basal tuft; inflo-
rescence 80-100 mm long; spike-
lets greyish yellow, rachilla per-

sistent, lemmas and/or paleas breaking up from the base up-
wards; lower glume lanceolate; lemma with lateral nerves
indistinct; palea keels entire; anthers 3, 0.9-1. 2 mm long;
caryopsis ovate.

Flowering January. Open places on brackish flats. Infre-
quent. Biome: Savanna. Zimbabwe. Similar to the annual
E. membranacea. Clayton et al. (1974)place the species in
synonymy with E. pseudosclerantha. This is unacceptable
because the latter species has a sprawling habit with a dense
basal tuft of leaves.

Description: De Winter 1961 (471). Voucher: De Winter
734. PRECIS code 9902860-03800.

Eragrostis lappula Nees

Perennial; shortly rhizomatous
and tufted (densely, erect); to
1 200 mm tall. Leaf blades to 250
mm long; 2-4 mm wide. Spike-
lets 5-10 mm long; 1.9-4.0 mm
wide. Spikelets with the rachilla
persistent, lemmas and/or paleas
breaking up from the base up-
wards; lemma with hairs 0.3-1. 2
mm long on the lateral nerves and sometimes on the median
keel: palea keels with hairs 0.3-1. 2 mm long which extend
beyond the lemma; anthers 3, 1.2 mm long.

Flowering December to May. Moist sandy soils of
annually flooded areas and river beds. Locally common.
Biome: Savanna. East Africa. Palatable pasture (when
young). Two varieties have been recognized based on
whether the inflorescence branches are appressed to the
central axis or lax. However intermediates have been found
and therefore the varieties have not been upheld in this
treatment. Similar to£. hierniana , which has glabrous later-
al nerves on the lemma, and£. inamoena , which has scabrid
palea keels.

Description: Chippindall & Crook 1976 (155), De Win-
ter in Chippindall 1955 (164), Clayton et al. 1970-1982
(212). Voucher: Smook 1935: Smook & Gibbs Russell
1956; Smith 2675. PRECIS code 9902860-04000.

Eragrostis leersiiformis Launert

Annual; tufted (erect to geni-
culate); to 500 mm tall. Leaf blad-
es to 170 mm long; to 2.7 mm
wide. Spikelets 3-5 mm long;
about 1 mm wide. Inflorescence
open, branches spreading, pedi-
cels slender; spikelets with the ra-
chilla fragile, the lemmas and/or
paleas breaking up from the apex
downwards; palea keels shortly ciliate; anthers 3, 0.16 mm
long; caryopsis ovate-elliptic.

Flowering February. Edges of vleis. Infrequent. Biome:
Savanna. Zambia. Note: no specimens were available for
this study and all the information was obtained from
literature. Resembles £. micrantha , which is a perennial
with the spikelet rachilla persistent, and £. aspera, which
has a subglobose caryopsis.

Description: Launert 1970 (160:224). Voucher: Van
Vuuren 1035 (WIND). PRECIS code 9902860-04100.

Eragrostis lehmanniana Nees var. chaunantha (Pilg.)

De Winter

Perennial; tufted (erect); to
600 mm tall. Leaf blades to 250
mm long; 1. 5-2.0 mm wide.

Spikelets 4-8 mm long; to 1 mm
wide. Culm nodes and internodes
hairy; inflorescence open, lax,
branches 1-2 at the base, not
whorled; spikelets linear to ob-
long, rachilla subpersistent usually fragile in the upper part;

153

glumes 1/2-2/3 the length of the lemmas directly above in
the intact spikelet; palea margins wide apart (except some-
times at the base), keels a narrow thin line, scaberulous; an-
thers 3, 1.0 mm long; caryopsis oblong.

Flowering December to April. Kalahari sand. Infre-
quent. Biome: Savanna. Possibly Zimbabwe. Said to be
separated from var. lehmanniana , which has glabrous culm
internodes, but a detailed study is needed.

Description: De Winter in Chippindall 1955 (145).
Voucher: Barker 198. PRECIS code 9902860-04200.

Eragrostis lehmanniana Nees var. lehmanniana

Fig. 88.

Knietjiesgras, Lehmann’s love
grass.

Perennial; tufted (erect, geni-
culate, sometimes rooting at the
lower nodes); to 600 mm tall.

Leaf blades to 100 mm long;

1.5-2. 8 mm wide. Spikelets 4-8
mm long; 1.0- 1.5 mm wide.

Culm nodes (those without branches) glabrous; basal
sheaths papery, nerves round, not close together or forming
prominent ridges, glabrous or slightly hairy at the extreme
base only; leaf blades gradually narrowing to the apex;
spikelets linear to oblong, dark green, grey-green to red with
yellow, rachilla subpersistent, fragile in the upper portion;
glumes 1/2-2/3 the length of the lemmas directly above in

the intact spikelet; palea margins wide apart, keels thin and
narrow, glabrous or scaberulous; anthers 3, 0.7 mm long;
caryopsis oblong.

Flowering November to June. Sand or sandy loam usual-
ly over limestone, in disturbed areas. Locally common.
Biome: Savanna and Nama-Karoo. Zimbabwe and Angola.
Introduced into east Africa and India as fodder. Hardy,
palatable pasture (especially when young), or erosion con-
trol (colonizes bare, eroded or denuded ground), or indica-
tor (denuded veld). Said to be separated from var.
chaunantha by the hairy culm internodes. A detailed study
is needed in this taxon.

Description: Stapf 1 898-1900 (601 ), De Winter in Chip-
pindall 1955 (145). Illustration: De Winter in Chippindall
1955 (fig. 113). Voucher: Smook 2928, Van Vuuren &
Giess 1 127. PRECIS code 9902860-04300.

Eragrostis macrochlamys Pilg. var. macrochlamys

Annual; tufted (erect and geni-
culate to procumbent); to 300 mm
tall. Leaf blades to 200 mm long;
to 3 mm wide. Spikelets 4-5 mm
long; 3-4 mm wide. Basal sheaths
glabrous or with a few scattered
hairs, mainly along the margins;
inflorescence branches usually
appressed to the main axis, occas-
ionally spreading in the lower part, spikelets many and
crowded; spikelets with the rachilla fragile, breaking up
from the apex downwards; glumes longer than the lemmas
directly above in the intact spikelet, glands present on the
keels; lemma acute (in profile), mucro present or absent; pa-
lea keels entire, scabrid; anthers 3, 0. 2-0.4 mm long; cary-
opsis oblong-lanceolate to oblong-elliptic.

Flowering October to May. Sandy soils in river courses
or calcrete soils, also disturbed places like roadsides. Local-
ly common. Biome: Savanna, Nama-Karoo, and Desert.
Endemic. Resembles var. wilmaniae , which has the inflo-
rescence eglandular, glumes shorter than or equal to the
lemma directly above in the intact spikelet, and E.
procumbens, in which the rachilla is persistent and becomes
fragile in the upper part and breaks up after the lower lem-
mas have started to fall.

Description: De Winter in Chippindall 1955 (175).
Illustration: De Winter in Chippindall 1955. Voucher: Giess
1743, Mueller 218. PRECIS code 9902860-04400.

Eragrostis macrochlamys Pilg. var. wilmaniae (C.E.

Hubb. & Schweick.) De Winter

Annual; tufted (erect, genicu-
late to procumbent); to 300 mm
tall. Leaf blades to 1 50 mm long;
to 2.5 mm wide. Spikelets 4-5
mm long; 3^1 mm wide. Basal
sheaths glabrous or with scattered
hairs; inflorescence branches usu-
ally appressed to the main axis,
spikelets many and usually densely crowded; spikelets
broadly oblong to broadly ovate, rachilla fragile, breaking
up from the apex downwards; glumes 2/3 — 4/5 the length of
the lemmas directly above in the intact spikelet, eglandular;
lemma acute to subacute (in profile); palea keels entire, sca-
brid; anthers 3, 0. 2-0.4 mm long; caryopsis oblong-
lanceolate to oblong-elliptic.

Flowering February to April. Moist areas in disturbed
places and on calcrete soils especially around pans. Locally
common. Biome: Savanna and Nama-Karoo. Endemic.
Differs from var. macrochlamys , which has glands on the
inflorescence, glumes equal to or longer than the lemmas
directly above in the intact spikelet. Resembles E.
procumbens , which has the spikelet oblong to elliptic and
the rachilla persistent, though the upper portion often
becomes fragile, lemmas and/or paleas breaking up from
the base upwards.

Description: De Winter in Chippindall 1955 (175).
Illustration: De Winter in Chippindall 1955. Voucher:
Henri 32. PRECIS code 9902860-04500.

154

Eragrostis membranacea Hack, ex Schinz

Annual; hydrophyte (occas-
ionally), tufted (erect); to 1 100
mm tall. Leaf blades to 400 mm
long; to 5 mm wide. Spikelets
3-1 5 mm long; 1 .9—4.0 mm wide.

Basal sheaths glabrous; inflores-
cence open; spikelets with rachil-
la persistent, the lemmas and/or
paleas breaking up from the base
upwards; upper glume 1/4—1/2 the length of the lemma
above in the intact spikelet; lemma very broadly ovate to
almost oblate, narrowly obtuse (in profile), coriaceous,
glossy with a clear broad membranous margin in the upper
part; palea keels entire, glabrous to minutely scaberulous;
anthers 3, 0.8-1. 2 mm long.

Flowering January to March. Sandy soils in moist areas
around pans and water courses, occasionally in shallow
water. Infrequent. Biome: Savanna. Zimbabwe & Zambia.
Similar to E. lamprospicula, which is said to be perennial
and to show a small difference in caryopsis structure. The
more delicate forms resemble E. gangetica , which has an-
thers 2, 0.1 -0.2 mm long.

Description: De Winter in Chippindall 1955 (170).
Voucher: De Winter9153; Soini PRE 56810. PRECIS code
9902860-04600.

Eragrostis micrantha Hack.

Weak perennial; tufted (erect);
to 1000 mm tall. Leaf blades to
600 mm long; to 3 mm wide.

Spikelets 2-4 mm long; to 1.2
mm wide. Culms not densely
compacted at base and easily
separated into individual culms;
basal sheaths glabrous; leaf blad-
es tapering to a filiform tip; inflo-
rescence 100-300 mm long, effuse, much branched, bran-
ches and spikelets spreading, the shortest pedicel of the
spikelet pair as long as or longer than the spikelet; spikelets
with rachilla persistent in the lower portion, usually fragile
in the upper portion, the lemmas and/or paleas breaking up
from the base upwards; glumes translucent, smooth or sca-
berulous around the apex and along the keel, 1/2-2/3 the
length of the lemmas directly above in the intact spikelet;
lemma light greenish to green-grey, strongly keeled along
the entire length; palea margins meeting or overlapping
along the entire length or at least at the apex, keels a thin
line or apparently palea only folded; anthers 3, 0.6-1 .0 mm
long; caryopsis lanceolate.

Flowering January to May. Sands, loams and calcareous
soils in disturbed areas and moist places around vleis and
pans, in semi-shade. Locally common. Biome: Savanna,
Grassland, and Nama-Karoo. Endemic. Could be confused
with an undescribed species from Swaziland, which has a
robust rhizome and a more montane habitat (eg. Compton
26766).

Description: Launert 1970 (160:92), Stapf 1898-1900
(608), De Winter in Chippindall 1955 (151). Voucher:
Smook & Gibbs Russell 2425. PRECIS code 9902860-
04700.

Eragrostis minor Host

(=£. poaeoides Beauv. ex
Roem. & Schult.) 1.

Little love grass.

Annual; loosely tufted (often
geniculate); to 600 mm tall. Leaf
blades to 120 mm long; to 5 mm
wide. Spikelets 3-19 mm long;

1. 3-2.0 mm wide. Leaf margins with raised glands; inflo-

rescence open, side branches usually less than 40 mm long,
pedicels stout; spikelet with the rachilla persistent, lemmas
and/or paleas breaking up from the base upwards; glumes
subequal; lemma obtuse; palea keels scabrid; anthers 3, 0.3
mm long; caryopsis broadly oblong.

Flowering November. Disturbed and weedy places. In-
frequent. Naturalized from southern Europe. Biome: Savan-
na. Warm temperate and subtropical regions of the Old
World, occasionally found as an introduction in the New
World tropics. Weed. Intergrades withf. cilianensis and E.
procumbens , and resembles E. barrelieri, which has
unequal glumes in the intact spikelet.

Description: De Winter in Chippindall 1955 (153), Clay-
ton et al. 1970-1982 (234). Voucher: Smith 6166. PRECIS
code 9902860-04750.

Eragrostis moggii De Winter var. moggii

Perennial; tufted (erect, geni-
culate to decumbent and rooting
at the nodes); 400-900 mm tall.

Leaf blades to 200 mm long; to 3
mm wide. Spikelets 4-7 mm
long; 1.0-1. 5 mm wide. Culm
nodes glabrous; inflorescence
open, branches spreading, pedi-
cels long to very long, with an an-
nular gland; spikelets linear to oblong, green to greyish-
green, rachilla persistent, lemmas and/or paleas tardily
breaking up from the base upwards; glumes to 2/3 the
length of the lemmas directly above in the intact spikelet,
often flushed purple; palea margins apart and not touching
for the entire length, sometimes touching at the base, keels
a thin line, entire, scaberulous; anthers 3, 0.5-0. 7 mm long;
caryopsis oblong.

Flowering December and April. Sandy soils, especially
in open forests. Infrequent. Biome: Savanna and Forest.
Mozambique.

Description: De Winter 1966 (137). Voucher: Smook
5720. PRECIS code 9902860-04900.

Eragrostis nindensis Fical. & Hiern

Fig. 89.

(=£. denudata Hack, ex
Schinz) 1.

Agtdaepluimgras.

Slender perennial; densely
tufted; to 900 mm tall. Leaf blad-
es 50-300 mm long; 2-3 mm
wide. Spikelets 4—20 mm long;

1 .5^4.0 mm wide. Culms to 2 mm wide; basal sheaths gla-
brous or hairy, but not densely woolly-hairy; leaves mainly
forming a basal tuft; inflorescence sparsely branched,
spikelets solitary or in clusters subsessile on the main axis
or side branches; spikelets yellowish-green with a serrated
outline, rachilla fragile, breaking up from the apex down-
wards; lower glume ovate; lowest lemma 2-3 mm long,
acute to acuminate, lateral nerves indistinct and not reach-
ing the margin; palea keels entire, flat, winged and scabrid;
anthers 3, 1.0-1. 4 mm long; caryopsis ellipsoid.

Flowering October to June. Prefers bare exposed areas
and stony sandy soils. Locally common. Biome: Savanna
and Nama-Karoo. Northwards to Tanzania, Zaire and
Angola. Palatable and drought resistant pasture. A
polymorphic species varying in shape and size of inflores-
cence and spikelets, often confused with E. racemosa ,
which has olive to dark green spikelets with margin outline
usually entire, rachilla persistent and lemmas and/or paleas
breaking up from the base upwards.

Description: Chippindall & Crook 1976 (161), De Win-
ter in Chippindall 1955 (167), Clayton et al. 1970-1982
(21 1). Illustration: De Winter in Chippindall 1955 (fig. 108
& 137). Voucher: Skarpe 539; Smook 2824. PRECIS code
9902860-05000.

155

Eragrostis obtusa Munro ex Fical. & Hiern

Perennial; tufted (geniculate);
to 400 mm tall. Leaf blades to 150
mm long; to 4.5 mm wide. Spike-
lets 3-5 mm long; 3-4 mm wide.

Basal sheaths glabrous or hairy
but not woolly-hairy at the base;
inflorescence open and lax to
somewhat contracted; spikelets
pallid, green to dark grey, broadly
ovate to oblong, rachilla extremely fragile, breaking up
from the apex downwards; lemma obtuse to rounded, lateral
nerves distinct; palea broadly elliptic to round, keels entire,
shortly ciliolate; anthers 3, 0.8-1. 3 mm long; caryopsis
broadly elliptic.

Flowering July to May. Sandy or limestone soils in dis-
turbed places such as roadsides and overgrazed areas. Lo-
cally common. Biome: Savanna, Grassland, and Nama-
Karoo. Endemic. Indicator (heavily grazed areas). Similar
to E. x pseud-obtusa, which has the lower part of the palea
keels broader and projecting from the upper part and ending
in a round or shallow notch at the top. It is said to be in-
termediate between E. obtusa and£. echinochloidea, which
have the lower part of the palea keel broader and projecting

from the upper part, with a very deep notch at the top.

Description: Stapf 1898-1900 (625), Hitchcock &
Chase 1950 (168), De Winter in Chippindall 1955 (173).
Illustration: De Winter in Chippindall 1955 (fig. 144).
Voucher: Smook & Gibbs Russell 2412; Smook 3015.
PRECIS code 9902860-05100.

Eragrostis omahekensis De Winter

Sandveldpluimgras.

Annual; tufted (erect and geni-
culate); 600-800 mm tall. Leaf
blades to 300 mm long. Spikelets
5-7 mm long; 0.8-1. 5 mm wide.

Inflorescence with the spikelets
densely clustered on the side
branches; spikelets with the ra-
chilla persistent, the lemmas and/or paleas breaking up from
the base upwards; lowest lemma chartaceous, broadly
elliptic to broadly oblong-ovate, 1.8-2. 2 mm long; palea
keels scaberulous; anthers 3, 1.0- 1.3 mm long; caryopsis
obovate-oblong.

Flowering February to May. On sand in disturbed places
such as roadsides and cultivated lands. Locally common.
Biome: Savanna and Nama-Karoo. Endemic. Pasture
(possibly good hay when grown in quantity). Resembles E.
cylindriflora and E. glandulosipedata, which have the low-
est lemma up to 1.7 mm long.

Description: De Winter 1961 (473). Illustration: De
Winter 1961 (fig. 2). Voucher: De Winter 2498, Liebenberg
4663. PRECIS code 9902860-05200.

Eragrostis pallens Hack.

Besemgras.

Robust perennial; densely
tufted (plants with stolons col-
lected at Mkuzi Game Reserve,

Natal); to 2000 mm tall. Leaf
blades to 1000 mm long; to 8 mm
wide. Spikelets 5— 1 5(— 25 ) mm
long; 1. 5-2.0 mm wide. Culms
erect, 2-4 mm wide; leaves mainly basal; inflorescence var-
iable, open with branches spreading or grading to contract-
ed with branches appressed to the main axis; spikelets
glossy, greenish-grey to yellowish, rachilla fragile, breaking
up from the apex downwards, the upper portion sometimes
breaking off as a whole; lowest lemma 1. 4-2.0 mm long,
broadly obtuse, lateral nerves obscure; palea obovate, usu-
ally protruding from the lemma, keels broad and flat, sca-
brid; anthers 3, 1.2 mm long.

Flowering December to May. Sandy soils, especially
with a high moisture content such as around seasonal pans.
Locally common. Biome: Savanna. Mozambique. Domestic
use (for playing musical instruments in Owamboland), or
timber (thatching grass by Owambos).

Description: Stapf 1 898- 1 900 (616). De Winter in Chip-
pindall 1955 (169). Illustration: Muller 1984 (fig. 79), De
Winter in Chippindall 1955 (fig. 138). Voucher: De Winter
7380. PRECIS code 9902860-05300.

Eragrostis patens Oliv.

Annual; tufted (erect to
procumbent); to 400 mm tall.

Leaf blades 30- 1 00 mm long; 2-3
mm wide. Spikelets 7-40 mm
long; 1.0-1. 5 mm wide. Inflores-
cence spikelike, with the spikelets
in wedge-shaped clusters; spike-
lets with the rachilla becoming
fragile soon after the lower lem-
mas start to fall, lemmas and/or paleas breaking up from the
base upwards; lemma with a mucro or awn to 0.5 mm long,

Fig. 90.

156

lateral nerves not awned or mucronate; palea keels with
hairs to 0.2 mm long; anthers 3, 0. 1-0.2 mm long.

Flowering February to June. In disturbed places such as
paths and overgrazed veld, usually on sandy soils but also
recorded on dolerite and clayey loams. Common (rare in
Namibia). Biome: Savanna. Northwards into east Africa
and Congo (Brazzaville). Indicator (overgrazed veld).

Description: Chippindall & Crook 1976 (159), Clayton
et al. 1970-1982 (225). Illustration: De Winter in Chippin-
dall 1955 (fig. 143), Clayton et al. 1970-1982 (fig. 64).
Voucher: Dahlstrand 881. PRECIS code 9902860-05400.

Eragrostis patentissima Hack.

Perennial; rhizomatous (rhi-
zomes stout), tufted (genicu-
late at base); 500-700 mm tall.

Leaf blades to 250 mm long; to 4
mm wide. Spikelets 6— 8(— 15) mm
long; 1. 5-3.0 mm wide. Basal
sheaths glabrous at the base; in-
florescence ovate to orbicular,
branches fairly rigid, spreading,
pedicels 3 times the length of the spikelets, spreading;
spikelets grey-green, rachilla persistent, becoming fragile
in the upper portion; lemma acuminate, lateral nerves dis-
tinct; palea oblanceolate, thick textured, apex acute to sub-
acute, keels entire, forming a prominent ridge, shortly
ciliolate; anthers 3, 0. 8-1.0 mm long; caryopsis elliptic.

Flowering November to March. Sandy to loamy soils of
open areas in damp places and disturbed areas. Infrequent
to locally common. Biome: Grassland. Endemic. Pasture
(eagerly grazed by cattle).

Description: Stapf 1898-1900 (613), De Winter in Chip-
pindall 1955 (162). Illustration: De Winter in Chippindall
1955 (fig. 131). Voucher: Acocks 9515. PRECIS code
9902860-05500.

Eragrostis pilgeriana Dinter ex Pilg.

Annual; tufted; to 400 mm
tall. Leaf blades to 150 mm long;
to 4 mm wide. Spikelets to 8 mm
long; 6-7 mm wide. Spikelets
strongly flattened, with the sides
appearing jagged, disarticulating
below the glumes at maturity and
falling entire as an unit; lemma
lanceolate in profile, keel winged
and scabrid; palea keels broadly winged, usually lacerate,
protruding laterally from the lemmas; anthers 3, 0.5 mm
long.

Flowering February to May. Soil specific, growing on
disturbed ground with calcrete and usually a high moisture
content. Locally common (on suitable soils). Biome: Savan-
na. Endemic. This species and E. superba , which is
perennial, are the only species in Eragrostis in the FSA
region in which the spikelets disarticulate below the glumes
at maturity and fall as entire units.

Description: Launert 1970 (1 60:83), De Winter in Chip-
pindall 1955 (171). Voucher: Giess 12542. PRECIS code
9902860-05600.

Eragrostis pilosa (L.) Beauv.

Annual; loosely tufted (erect,
occasionally geniculate); to 700
mm tall. Leaf blades 20-200 mm
long; 1 -4 mm wide. Spikelets 3-7
mm long; 0.7-1. 2 mm wide. In-
florescence delicate, open, bran-
ches and pedicels slender and
usually flexible, spikelets distant,
axils of branches bearded; spike-
lets linear to oblong, lemmas on the same side of the rachil-

la distinctly overlapping the lemma above, lemmas becom-
ing conspicuously shorter towards the apex of the spikelet,
with the rachilla persistent, lemmas and/or paleas breaking
up from the base upwards; lower glume to 1/3 the length
of the lemma above in the intact spikelet, weakly keeled;
lemma broadly ovate, lateral veins visible, lowest lemma
1.0-1. 6 mm long; palea keels glabrous to scabrid: anthers
3, 0.2-0. 3 mm long; caryopsis ellipsoid.

Flowering October to May. Sandy soils in wet areas such
as pan edges, vleis and river banks, disturbed places, often
in the shade. Infrequent to locally common. Biome: Savan-
na. Tropical and warm temperate regions of Old World, in-
troduced to New World. Weed. Similar to E. aethiopica,
which has the lowest lemma 0.7-1 .0 mm long and the later-
al veins not visible, and E. remotiflora , which has the lem-
mas on the same side of the rachilla not overlapping the
lemma directly above.

Description: Chippindall & Crook 1976 (164), Hitch-
cock & Chase 1950 (150), De Winter in Chippindall 1955
(154), Clayton et al. 1970-1982 (214). Voucher: Smook
1765, Hilliard 5383. PRECIS code 9902860-05700.

Fig. 90. Eragrostis pollens

157

Eragrostis plana Nees

Eragrostis porosa Nees

Taaipoleragrostis.

Perennial; densely tufted; to
1000 mm tall. Leaf blades to 800
mm long; to 4 mm wide. Spike-
lets 6-10 mm long; 0. 5-2.0 mm
wide. Basal sheaths strongly
compressed, smooth and shiny;
inflorescence branches usually
spreading, spikelets appressed to the branches; spikelets lin-
ear-oblong, with the rachilla persistent, lemmas and/or pa-
leas breaking up from the base upwards; lower glume scale-
like, reaching up to 1/3 the length of the lemma above and
the upper glume barely reaching or just touching the base
of the lemma above in the intact spikelet; lemmas with lat-
eral nerves prominent and with glandular dots; palea keels
entire, glabrous to scabrid, glandular dots present or absent;
anthers 3, 1. 6-2.0 mm long; caryopsis oblong.

Flowering November to May. In high rainfall regions in
waterlogged, overgrazed, burnt or disturbed areas. In dry
areas it favours wet soils around vleis and rivers. Locally
common to locally dominant. Biome: Savanna and Grass-
land. Zimbabwe, Zambia, Malawi, introduced to India. Do-
mestic use (weaving of hats, baskets, necklaces and
bangles), or pasture (occasionally grazed in autumn), or in-
dicator (overgrazed and burnt areas), or traditional medi-
cine (in Lesotho). Similar to E. tenuifolia, which has an
identical caryopsis but no glands on the lateral nerves of the
lemmas and is a weak perennial or an annual. Vegetatively
very similar to Sporobolus pyramidalis, which has one flo-
ret per spikelet.

Description: Chippindall & Crook 1976 (162), Stapf
1898-1900 (609), De Winter in Chippindall 1955 (157).
Voucher: Smook 4714, Hanekom 1701. PRECIS code
9902860-05800.

Eragrostis planiculmis Nees

( =E . nebulosa Stapf) 1.

Besemeragrostis.

Perennial; tufted (erect); to
1200 mm tall. Leaf blades
setaceous, 100-900 mm long; to
1.5 mm wide. Spikelets to 8 mm
long; 0.5-2.0 mm wide. Base
with culms densely and strongly compacted, not easily
separated, nodes glabrous; basal sheaths glabrous, inner
sheaths often yellow; leaves mainly in a dense basal tuft,
leaf blades long-tapering at apex, straight or drooping; in-
florescence 100-700 mm long, open, much branched, low-
est branches whorled or not whorled, pedicels long; spike-
lets linear, 5-1 1 -flowered, rachilla persistent in the lower
portion, fragile in the upper part, lemmas and/or paleas
breaking up from the base upwards; glumes translucent,
smooth or scaberulous around the apex and along the keels,
lower glume up to 3/4 the length of the lemma above in the
intact spikelet; lemma dark green to greenish -grey, not
strongly keeled, keel obscure in lower part; palea margin
nearly touching to touching along the entire length, over-
lapping at the apex, keel a narrow line; anthers 3, 0.6-1. 2
mm long.

Flowering November to April. Clay or dolorite soils in
depressions, vlei margins and disturbed areas. Infrequent to
locally common. Biome: Fynbos, Savanna, and Grassland.
Endemic. Resembles forms of£. curvula , which has dense-
ly hairy basal sheaths.

Description: Stapf 1898-1900 (63 1 ), De Winter in Chip-
pindall 1955 (142). Voucher: Dieterlen 317, Smook &
Gibbs Russell 2169a. PRECIS code 9902860-05900.

Annual; loosely tufted (usual-
ly erect); to 800 mm tall. Leaf
blades 40-150 mm long; 2-5 mm
wide. Spikelets 3-5 mm long;

1.0-1. 5 mm wide. Leaf sheaths
densely pilose with bulbous-based
hairs; inflorescence with the low-
est branches whorled; spikelets
with the rachilla persistent in the
lower part, fragile above, lemmas and/or paleas breaking up
from the base upwards; lowest lemma obovate-elliptic,
truncate to broadly rounded, 1.0-1. 5 mm long; palea keels
minutely scaberulous; anthers 3, 0.6-0. 9 mm long;
caryposis ellipsoid.

Flowering January to July. Stony or sandy soils often on
limestone around rivers and pans, also in disturbed areas.
Infrequent to common (widespread). Biome: Savanna and
Nama-Karoo. Zimbabwe to Kenya, with a few records from
Chad and Ethiopia. Reported to intergrade with E.
cylindriflora, which has the lowest lemma broadly elliptic,
obtuse to subacute and 1.5-1. 7 mm long.

Description: Stapf 1898-1900 (604), De Winter in Chip-
pindall 1955 (150), Clayton et al. 1970-1982 (240).
Voucher: De Winter 7472, De Winter & Codd 284, Mostert
1637. PRECIS code 9902860-06200.

Eragrostis procumbens Nees

Annual; tufted (geniculate or
procumbent); to 500 mm tall.

Leaf blades to 200 mm long; to
3.5 mm wide. Spikelets about 7
mm long; ( 1 .7— )2.0— 2.5 mm
wide. Basal sheaths glabrous or
hairy, but not with woolly hairs;
inflorescence with the side bran-
ches usually appressed to the
main axis, though the lower branches sometimes spreading,
with many spikelets densely congested and appressed to the
branches; spikelets oblong to elliptic, rachilla persistent,
sometimes becoming fragile in the upper part, lemmas
and/or paleas breaking up from the base upwards; glumes
shorter than the lemmas directly above in the intact spikelet;
lemmas acute ( 1 .7— )2.0— 2.5 mm long, with a minute mucro
present or absent; palea keels entire, scabrid; anthers 3,
0.2-0. 3 mm long; caryopsis oblong.

Flowering October to June. Moist gravel or sandy soils
in depressions, along water courses and disturbed areas. Lo-
cally common. Biome: Savanna and Nama-Karoo.

Endemic. Weed. Similar to E. kingesii , which is a smaller
plant with the spikelets 1.0-1. 5 mm wide, and E.
macrochlamys var. macrochlamys, in which the rachilla is
fragile and the spikelet breaks up from the apex downwards.

Description: Stapf 1898-1900 (620), De Winter in Chip-
pindall 1955 (159). Illustration: De Winter in Chippindall
1955 (fig. 127). Voucher: Zietsman 1666; Smook 3262.
PRECIS code 9902860-06300.

Eragrostis x pseud-obtusa De Winter

Fig. 91.

Perennial; tufted (erect); to
600 mm tall. Leaf blades to 150
mm long; 2-3 mm wide. Spike-
lets 3-5 mm long; 2. 5-3. 5 mm
wide. Basal sheaths glabrous, or
if hairy not with long-woolly
hairs for a distance up along the
sheaths; inflorescence lax to
dense, sparsely branched, spike-
lets congested; spikelets with rachilla fragile, lemmas
and/or paleas breaking up from the apex downwards;

158

glumes boat-shaped, acute to subacute; palea obtuse to
rounded, keels with the lower portion very broad and
protruding from the rest of the keel, top of the projecting
portion rounded or shallowly notched, ciliolate; anthers 3,
0.7-0. 8 mm long; caryopsis elliptic.

Flowering November to May. Sandy loam, shallow
sandy soils, sand over limestone, in moist areas such as
ditches, along streambeds and dams, also in disturbed
places. Locally common. Biome: Savanna and Nama-
Karoo. Endemic. Intermediate between E. obtusa , which
has the paleas entire, and E. echinochloidea, which has the
palea acute, and the lower projecting portion of the keels
deeply notched at the top.

Description: De Winter 1961 (474). Illustration: De
Winter 1961 (476). Voucher: Smook 2785. PRECIS code
9902860-06400.

Fig. 91 . Eragrostis x pseud-obtusa

Eragrostis pseudosclerantha Chiov.

Footpath love grass.

Short-lived perennial; some-
times stoloniferous (geniculate,
rooting at the nodes); 300^)00
mm tall. Leaf blades to 90 mm
long; to 4.5 mm wide. Spikelets
4-10 mm long; 2-3 mm wide.

Plants sprawling; leaves mainly
from a dense basal tuft; inflorescence to 75 mm long; spike-
lets narrowly elliptic, greyish -green, rachilla persistent,
lemmas and/or paleas breaking up from the base upwards;
lower glume lanceolate, to 1/2 the length of the lemma di-
rectly above and upper glume 1/2-2/3 the length of the lem-

ma above in the intact spikelet; lemma with lateral nerves
indistinct; palea narrow between the keel and the margins,
the margins far apart, keels a narrow line, entire, scabrid;
anthers 3, 0.8-1 .0 mm long; caryopsis ellipsoid.

Flowering September to April. Stony ground in open
places in short grassland or under trees, in disturbed places.
Locally common. Biome: Savanna and Grassland.

Northwards through east Africa to Ethiopia. Weed.

Description: Chippindall & Crook 1976(169), De Win-
ter in Chippindall 1955 (167), Clayton et al. 1970-1982
(231). Illustration: De Winter in Chippindall 1955 (fig.

1 38). Voucher: Smook 3 1 58, Chippindall 1 8. PRECIS code
9902860-06500.

Eragrostis pygmaea De Winter

Annual; tufted (erect or de-
cumbent); to 70 mm tall. Leaf
blades to 40 mm long; to 2 mm
wide. Spikelets 4-7 mm long;

1.0- 1.5 mm wide. Vegetative
parts have glandular hairs with
swollen tips; basal sheaths dense-
ly covered with long, bulbous-
based hairs; inflorescence dense-
ly contracted, pedicels stout; spikelets narrowly lanceolate,
rachilla persistent, lemmas and/or paleas breaking up from
the base upwards; lower glume 1/2 the length of the lemma
above in the intact spikelet; lemma acute, 1 .0-1.6 mm long,
palea keels scabrid; anthers 3, 0.2 mm long; caryopsis sub-
globose.

Flowering March to May. Shallow depressions on sandy
flats. Locally common (confined to flat areas north of high
sand dune between Luderitz and the Kuiseb river). Biome:
Desert. Endemic. Similar to E. kingesii , which does not
have bulbous-based hairs on the leaf sheaths and has an ob-
long-elliptic caryopsis.

Description: De Winter 1969 (72). Voucher: De Winter
& Hardy 8050. PRECIS code 9902860-06600.

Eragrostis racemosa (Thunb.) Steud.

PI. 79.

( -E . chalcantha Trin.) 1.

Smalhartjie-eragrostis,
narrow heart love grass.

Perennial; densely tufted (lea-
ves mainly basal); to 800 mm tall.

Leaf blades 60- 1 00 mm long; 2-5
mm wide. Spikelets 3-10 mm
long; 1.5-4. 5 mm wide. Basal sheaths glabrous or thinly
silky-hairy; inflorescence open or contracted, sparsely
branched, secondary branches present or absent, primary
branches stiff, with 2-4 spikelets on short stout pedicels;
spikelets dark greenish-grey, olive or brownish-grey,
outline usually smooth, rachilla persistent, lemmas and/or
paleas breaking up from the base upwards; glumes
cartilaginous, ovate, boat-shaped; lemma with lateral
nerves indistinct; palea keels a narrow ridge, entire, scaber-
ulous; anthers 3, 1 .0-1.6 mm long; caryopsis almost square.

Flowering August to May. On shallow sandy, stony or
clayey soils. Common (widespread). Biome: Fynbos,
Savanna, and Grassland. North to Sudan, and in
Madagascar. Erosion control (useful cover on shallow soils
and in heavily grazed areas). Often confused with E.
nindensis , which has yellowish -green spikelets with a
serrate outline and the rachilla fragile and breaking up from
the apex downwards. Similar to E. sclerantha subsp.
sclerantha , which has the basal sheaths with dense woolly
hairs.

ter in Chippindall 1955 (165), Clayton et al. 1970-1982
(230). Illustration: De Winter in Chippindall 1955 (fig.
135). Voucher: Smook 4792, Kluge 1098. PRECIS code
9902860-06700.

159

Eragrostis remotiflora De Winter

Weak perennial, or annual;
tufted (erect or with some culms
geniculate); to 600 mm tall. Leaf
blades to 250 mm long; to 2.5 mm
wide. Spikelets to 5 mm long; to
1 mm wide. Inflorescence open,
branches spreading, pedicels
slender; spikelets oblong to lin-
ear, lemmas on the same side of
the rachilla either not or just overlapping the one above, ra-
chilla persistent, lemmas and/or paleas breaking up from the
base upwards; lower glume less than 1/3 the length of the
lemma above in the intact spikelet, weakly keeled; lemma
1.0-1. 8 mm long; palea keels scabrid to glabrous; anthers
3, 0.2-0. 3 mm long; caryopsis oblong.

Flowering January to April. Wet and damp areas in pans,
vleis and river floodplains, also disturbed areas in semi-arid
regions. Locally common. Biome: Savanna. Endemic. Pas-
ture (grazed). Similar to E. pilosa and E. aethiopica , which
have the lemmas on the same side of the rachilla overlap-
ping the lemma above.

Description: De Winter 1961 Bothalia 7 (477).
Illustration: De Winter 1961 (478). Voucher: Acocks
14016, Smook & Gibbs Russell 2371. PRECIS code
9902860-06800.

Eragrostis rigidior Pilg.

Curly leaf, krulblaar.

Perennial; loosely tufted
(erect to geniculate); to 1000 mm
tall. Leaf blades to 200 mm long;
to 5 mm wide. Spikelets 3. 5-7.0
mm long; 1.0-1. 5 mm wide.

Unbranched culm nodes gla-
brous; basal sheaths papery,
nerves rounded and well apart, glabrous to slightly hairy at
the very base only; leaf blades narrowing abruptly into a
long tapering apex, curly when dry; inflorescence open,
branches spreading, lowest branches usually whorled,
spikelets usually contracted along the branches; spikelets
linear to oblong, rachilla fragile in the upper portion, lem-
mas and/or paleas breaking up from the base upwards;
lower glume 4/5 to longer than the lemma directly above
and upper glume 1/2-3/4 the length of the lemma directly
above in the intact spikelet; palea margins wide apart except
at the base and occasionally touching at the apex, keels a
thin line, smooth to scaberulous; anthers 3, 0.8-1. 2 mm
long.

Flowering September to May. Sand, loam, humus loam
or calcrete soils in open patches, disturbed areas and old
cultivation sites. Locally common. Biome: Savanna. East
Africa. Pasture (valuable fodder in dry areas). Intermediates
with E. barbinodis, which has hairy nodes, have been
recorded. Resembles E. lehmanniana , which has leaf blades
that gradually narrow to the apex and do not curl when dry
and the inflorescence with the lowest branches never
whorled.

Description: Chippindall & Crook 1976 ( 170), De Win-
ter in Chippindall 1955 (149), Clayton et al. 1970-1982
(242). Illustration: De Winter in Chippindall 1955 (fig.
115). Voucher: Smook 5300. PRECIS code

9902860-06900.

Eragrostis rogersii C.E. Hubb.

Annual; tufted (geniculate to
rooting at the nodes); to 400 mm
tall. Leaf blades to 90 mm long;
to 4 mm wide. Spikelets 6-12 mm
long; 2. 5-4. 2 mm wide. Basal
sheaths glabrous or with bulbous-
based hairs along the margins or
scattered near the leaf blades; in-
florescence with branches slight-

Fig. 92.

Fig. 92. Eragrostis rigidior

160

ly spreading, spikelets few and distant; spikelets with the
rachilla fragile, breaking up from the apex downwards; up-
per glume 2/3 — 3/4 the length of the lemma above in the
intact spikelet; lemma acute (in profile), a median mucro
present or absent; palea keels entire, scaberulous; anthers
3, 1.1 mm long.

Flowering March. Disturbed sandy soils. Infrequent.
Biome: Savanna. Zimbabwe, Zambia. Resembles E. dinteri ,
which has acuminate and usually awned lemmas. Either
rare or poorly collected in FSA area, more common in
Zimbabwe.

Description: Hubbard 1934 Kew Bull. (115). Voucher:
Ellis 2752. PRECIS code 9902860-07000.

Eragrostis rotifer Rendle

(=£. margaritacea Stapf) 1 .

Perennial; tufted (erect, occas-
ionally decumbent); to 1500 mm
tall. Leaf blades to 300 mm long;
to 4 mm wide. Spikelets 4-10 mm
long; to 1 mm wide. Basal sheaths
densely hairy at the base; inflo-
rescence open, branches spread-
ing, lowest branches whorled, branches and pedicels dense-
ly covered with prickles giving a greenish-white
appearance, spikelets contracted to the branches; spikelets
linear to oblong, rachilla persistent, lemmas and/or paleas
breaking up from the base upwards; glumes 1/2-2/3 the
length of the lemmas directly above in the intact spikelet;
lemma greyish-green usually flushed purple in the upper
portion above which is a yellow portion, and the apex has
a white membranous margin; palea margins touching to
overlapping at the apex, keels a obscure line, smooth or sca-
berulous; anthers 3, 0.5-0. 8 mm long; caryopsis ellipsoid.

Flowering November to July. Mainly sandy soils in
moist areas like vleis, pan edges and river beds, also dis-
turbed areas. Locally common (often pure stands in damp
places). Biome: Savanna and Nama-Karoo. Northwards to
Tanzania. Pasture (remaining green for long time, valuable
fodder in dry areas). Similar to E. heteromera , which has
the basal sheaths glabrous or obscurely hairy at the very
base.

Description: Chippindall & Crook 1976 (171), De Win-
ter in Chippindall 1955 (156), Clayton et al. 1970-1982
(216). Illustration: Muller 1984 (fig. 82). Voucher: De
Winter 2397. PRECIS code 9902860-07100.

Eragrostis sabinae Launert

Perennial; stoloniferous and
tufted (densely; leaves short,
mainly basal); 90-150 mm tall.

Leaf blades to 250 mm long; to
1.5 mm wide. Spikelets 4-7 mm
long; 0. 8-1.0 mm wide. Basal
sheaths densely hairy; culm nodes
usually with long spreading white
hairs; leaf blades only slightly
tapering to the apex; inflorescence 25-80 mm long, open,
moderately branched, spikelets in loose clusters at the ends
of the branches or spreading; spikelets with rachilla persis-
tent in lower portion and fragile in upper part, lemmas
and/or paleas breaking up from the base upwards; glumes
translucent, smooth or scaberulous along the keel, 1/2-2/3
the length of the lemmas directly above in the intact spike-
let; lemma green to grey-green, usually whitish at the apex;
palea margins meeting along the entire length or only touch-
ing at the apex, keels a thin line; anthers 3, 0. 4-0.8 mm
long.

Flowering February to May. Brackish or saline soils
around vleis, pans and springs. Locally common. Biome:
Savanna. Endemic. Resembles some forms of E. laevissima ,
which has glandular dots on the lateral nerves of the lem-
mas.

Description: Launert 1970 (225). Voucher: Mueller
1474, Smook 5128. PRECIS code 9902860-07200.

Eragrostis sabulosa (Steud.) Schweick.

Perennial; tufted and rhizo-
matous (rhizome long and
creeping); 60-150 mm tall. Leaf
blades 1 0 — 40 mm long; to 3 mm
wide. Spikelets to 7 mm long;

1 .5-2.0 mm wide. Culm internod-
es short and exceeded by the leaf
sheaths; leaves mainly cauline;
inflorescence usually less than 40
mm long, very dense with branches appressed to the main
axis and spikelets densely crowded and appressed to the
branches; spikelets dark olive-grey, rachilla tardily break-
ing up between the florets; upper glume acute to obtuse and
jagged; lowest lemma 1.6-2. 5 mm long, lateral veins usual-
ly indistinct; palea keels with cilia less than 0.1 mm long;
anthers 3, 1.2 mm long; caryopsis ellipsoid.

Flowering March, and October to November. Sandy
soils especially beach sand. Locally common. Biome: Fyn-
bos. Endemic. Similar to E. sarmentosa , which has the low-
est lemma 1.5 mm long and anthers 0.2-0. 3 mm long.
Resembles Sporobolus virginicus, which has only one floret
per spikelet, in habit and habitat.

Description: De Winter in Chippindall 1955 (161).
Illustration: De Winter in Chippindall 1955 (fig. 129).
Voucher: Crook 1040. PRECIS code 9902860-07300.

Eragrostis sarmentosa (Thunb.) Trin.

Mat-forming perennial; short
rhizomatous and tufted (culms
occasionally geniculate, decum-
bent and rooting at the nodes); to
400 mm tall. Leaf blades to 100
mm long; to 4.5 mm wide. Spike-
lets 3-7 mm long; 1.5-1. 7 mm
wide. Leaves mainly cauline; in-
florescence narrow and contract-
ed, branches appressed to the main axis, pedicels thick,
spikelet groups often distant on the main axis, spikelets ap-
pressed to the branches; spikelets greyish-green to purple,
rachilla persistent, the upper part often becoming fragile,
lemmas and/or paleas breaking up from the base upwards;
upper glume acute; lowest lemma 1 .5 mm long, lateral veins
distant, usually not reaching the margin, never excurrent in-
to a mucro; palea keels scaberulous; anthers 3, 1 .2 mm long;
caryopsis ellipsoid to ovoid.

Flowering July to May. Moist, sandy areas such as the
edges of riverine vegetation, along floodplains and dams,
also in disturbed overgrazed areas. Locally common.
Biome: Fynbos, Savanna, and Succulent Karoo. Zambia,
Zimbabwe. Similar to E. sabulosa , which has the lowest
lemma 1.6-2. 5 mm long and anthers 0.6-1. 3 mm long.

Description: Stapf 1898-1900 (618), De Winter in Chip-
pindall 1955 (162). Illustration: De Winter in Chippindall
1955 (fig. 130). Voucher: De Winter & Vahlmeijer 8584,
Smith 773. PRECIS code 9902860-07400.

Eragrostis sclerantha Nees subsp. sclerantha

Perennial; densely tufted; to
700 mm tall. Leaf blades to 200
mm long: to 6.5 mm wide. Spike-
lets 2-5 mm long; 1. 5-3.0 mm
wide. Basal sheaths densely
woolly-hairy, sometimes this on-
ly visible on the inner sheaths; in-
florescence branches spreading,
with many spikelets, either ap-
pressed to the branches or spreading; spikelets dark olive-
green, rachilla persistent, lemmas and/or paleas breaking up
from the base upwards; lemma acute, lateral nerves
conspicuous; palea keels entire, scabrid; anthers 3, 0. 7-1.0
mm long; caryopsis ellipsoid.

Flowering January to May. Sandy soils and sandy loams
between rocks (often quartzite). Infrequent (but
widespread). Biome: Savanna and Grassland. Zimbabwe,

161

Angola. Similar to£. racemosa , which has the basal sheaths
glabrous or hairy, but not densely woolly-hairy, and subsp.
villosipes, which has a contracted inflorescence with the
branches appressed to the main axis. Resembles E. desolata
Launert found in Zimbabwe, which has the basal sheaths
glabrous or hairy, not woolly-hairy, leaf blades 0.5-2. 5 mm
wide and anthers 1.2-1. 3 mm long.

Description: Chippindall & Crook 1976 (172), Stapf
1898-1900 (615), De Winter in Chippindall 1955 (116),
Clayton et al. 1970-1982 (229). Illustration: De Winter in
Chippindall 1955 (fig. 136). Voucher: Cohen 862, Burtt
Davy 9240. PRECIS code 9902860-07500.

Eragrostis sclerantha Nees subsp. villosipes (Jedw.)

Launert

(=E. sclerantha Nees var.
villosipes (Jedw.) De Winter) 7.

Perennial; densely tufted
(erect); to 600 mm tall. Leaf blad-
es to 200 mm long; 2-6 mm wide.

Spikelets 2-8 mm long; 1. 5-2.0
mm wide. Leaves forming a
dense basal tuft; basal sheaths with dense, yellowish, wool-
ly hairs; inflorescence narrow, sparsely branched, branches
appressed to the main axis, spikelets appressed, solitary or
in groups of 2-3, pedicels short; spikelets olive-green, ra-
chilla persistent, lemmas and/or paleas breaking up from the
base upwards; glumes cartilaginous, upper glume ovate,
boat-shaped; lemma with lateral nerves indistinct and not
reaching the margin; palea keels a narrow line, scaberulous;
anthers 3, 0.7-1. 0 mm long; caryopsis ellipsoid.

Llowering Lebruary. Wooded grassland. Infrequent.
Biome: Savanna. Tropical Africa. Said to occur in
Botswana, and there is a doubtful fragment from Namibia.
In the Transvaal only a single specimen has been collected
in 1912. Differs from subsp. sclerantha , which has the in-
florescence spreading.

Description: Launert 1961 Bot. Soc. Brot. 35,2 (19),
Chippindall & Crook 1976 (172), De Winter in Chippindall
1955 (66), Clayton et al. 1970-1982 (229). Voucher: Bell
PRE 5985. PRECIS code 9902860-07600.

Eragrostis scopelophila Pilg.

Bergpluimgras.

Wiry, much branched shrub or
dwarf shrub; tufted (forming
dense bushes); to 1000 mm tall.

Leaf blades to 250 mm long; to 5
mm wide. Spikelets 3-10 mm
long; 2. 2-3. 3 mm wide. Basal
sheaths glabrous; leaves mainly
cauline; inflorescence open, pedicels (excluding terminal
ones) shorter than or to as long as the spikelets; spikelets
green to dark greenish grey, the opposite rows of florets
hardly overlapping, rachilla visible and subpersistent with
the upper portion fragile; lemma acute to obtuse, lateral
nerves distinct; palea margins not touching along the entire
length, keels entire, narrow, rounded, scaberulous; anthers
3, 1.3-1. 5 mm long; caryopsis broadly oblong.

Llowering December to April (and August and October).
Mountainous areas, often associated with dolomite. Locally
common. Biome: Savanna and Nama-Karoo. Endemic.
Lairly palatable pasture (stays green in winter).

Description: De Winter in Chippindall 1955 (160).
Illustration: Muller 1984 (fig. 83). Voucher: De Winter
2342. PRECIS code 9902860-07700.

Eragrostis stapfii De Winter

Perennial; densely tufted
(erect); to 500-900 mm tall. Leaf
blades to 200 mm long; to 2 mm
wide. Spikelets to 4 mm long;

0. 5-1.0 mm wide. Basal sheaths
glabrous at the base; leaves form-
ing a dense basal tuft, usually
tightly rolled and curling; inflo-
rescence open, delicate, lowest
branches whorled; spikelets with 2-3(-5) florets, rachilla
subpersistent upper part fragile, lemmas and/or paleas
breaking up from the base upwards; lower glume to 3/4 the
length of the lemma directly above and upper glume
1 /3 — 4/5 as long as the lemma above in the intact spikelet;
lemma pale yellowish grey-green, not strongly keeled, more
rounded at the back with the keel obscure at the base; palea
margins almost touching along the entire length, overlap-
ping at the apex, keels an obscure line, smooth or
scaberulous; anthers 3, 0.8 mm long.

Llowering November to April. Shallow sand or coarse
sandy soils, sometimes in wet disturbed areas. Locally com-
mon. Biome: Savanna. Southern tropical Africa. Resembles
E. micrantha, which is a weak perennial with lemmas
strongly keeled, and£. habrantha, which has broadly ovate
spikelets.

Description: De Winter in Chippindall 1955 (152).
Illustration: De Winter in Chippindall 1955 (fig. 119).
Voucher: Giess 14219, Pole Evans 3141. PRECIS code
9902860-07900.

Eragrostis stenothyrsa Pilg.

Perennial; tufted; to 500 mm
tall. Leaf blades to 100 mm long;
to 2 mm wide. Spikelets 4—8 mm
long; 0.8-1 .5 mm wide. Leaves in
a dense basal tuft; inflorescence
contracted, branches closely ap-
pressed to the main axis; spikelets
lanceolate to narrowly ovate, yel-
lowish, often flushed purple, ra-
chilla persistent, lemmas and/or paleas breaking up from the
base upwards; lemmas on the same side of the rachilla over-
lapping the lemma above by 2/3 or more; glumes
chartaceous; lemma with with lateral nerves not reaching
the margin; palea keels a narrow line less than 0. 1 mm wide,
scaberulous; anthers 3, 1.3 mm long.

Llowering March to April. Moist areas around pans.
Rare. Biome: Savanna. Endemic.

Description: De Winter in Chippindall 1955 (141).
Voucher: Giess, 14851. Volk
9902860-08000.

Eragrostis superba Peyr.

Weeluiseragrostis, sawtooth
love grass.

Perennial; densely tufted (of-
ten geniculate); to 1000 mm tall.

Leaf blades to 400 mm long; 3-12
mm wide. Spikelets 6-16 mm
long; 3-10 mm wide. Spikelets
strongly flattened with the sides
appearing jagged, disarticulating below the glumes at
maturity and falling as an entire unit; lemma narrowly ovate
in profile, keel winged and scaberulous; palea keels broadly
winged, entire, minutely ciliolate, hardly projecting lateral-

12473. PRECIS code

PI. 80.

162

ly from the lemma; anthers 3, 1.5-2. 5 mm long.

Flowering August to May. Sandy and stony soils in dis-
turbed places or drainage areas. Widely common. Biome:
Fynbos, Savanna, and Grassland. Northwards through east
Africa to Sudan. Hay and readily grazed, fairly palatable
and drought resistant pasture (cultivated in USA), or
erosion control (reseeding denuded areas). This species and
E. pilgeriana, which is annual, are the only Eragrostis
species in the FSA area with spikelets that disarticulate
below the glumes at maturity and fall as entire units.

Description: Chippindall & Crook 1976 (163), Stapf
1898-1900 (622), De Winter in Chippindall 1955 (171).
Illustration: Muller 1984, De Winter in Chippindall 1955
(fig. 141). Voucher: Wederman & Oberdieck 2773; Kinges
1426. PRECIS code 9902860-08100.

Eragrostis tef (Zucc.) Trotter

(=E. abyssinica (Jacq.)

Link) 9.

Teff.

Annual; loosely tufted (erect);
to 600 mm tall. Leaf blades to 300
mm long; to 4 mm wide. Spike-
lets 5. 5-9.0 mm long; 1. 5-2.0
mm wide. Inflorescence open or contracted, branches usual-
ly more than 40 mm long, flexible and slender, pedicels
slender; spikelets with rachilla persistent and the lemmas
and paleas remaining intact at maturity; upper glume
1/2-2/3 the length of the lemma above in the intact spikelet;
lemmas 2.0-2. 7 mm long; palea keels scaberulous; anthers
3, 0.3-0. 5 mm long; caryopsis oblong.

Flowering November to May (and July and September).
An escape from cultivation which grows in weedy places,
along roadsides and where it has been naturalized. Infre-
quent to locally common. Naturalized from northern Africa
and Ethiopia. Biome: Fynbos, Savanna, Grassland, and
Nama-Karoo. Northwards to east Africa. Introduced to
most tropical countries where it can become adventitious.
Food and drink (a staple cereal crop in Ethiopia), or pasture
(planted as hay), or erosion control (rehabilitation of road
reserves).

Description: Chippindall & Crook 1976 (165), De Win-
ter in Chippindall 1955 (154), Clayton et al. 1970-1982
(213). Voucher: Smook 5423. PRECIS code

9902860-08200.

Eragrostis tenella (L.) Roem. & Schult.

Annual; tufted (erect and geni-
culate); to 500 mm tall. Leaf blad-
es 60-90 mm long; to 3 mm wide.

Spikelets 1.5-2. 5 mm long. Inflo-
rescence open, with branches
spreading and the spikelets dis-
tant, eglandular, or with non-
sticky glands; spikelets with ra-
chilla fragile, breaking up from
the apex downwards; lemma keel smooth or scabrid; palea
keels with hairs 0. 3-0.4 mm long and exserted from the
lemma; anthers variable, 2-3, (on the same inflorescence),
0.15-0.20 mm long.

Flowering January to April. Bare, moist, sandy soils in
disturbed places like pathsides and cultivated lands. Locally
common. Biome: Savanna. Throughout Tropics. Weed.
Distiguished from E. viscosa, which has sticky glandular
areas on the inflorescence, but the boundary is not sharp and
intermediates are found. Similar to E. ciliaris, in which the
lemma has long stiff hairs on the keel, and E. arenicola,
which has the inflorescence contracted with the spikelets

densely appressed to the branches.

Description: Hitchcock & Chase 1950 (168), Clayton et
al. 1970-1982 (206). Voucher: Smook 4142. PRECIS code
9902860-08300.

Eragrostis tenuifolia (A. Rich.) Steud.

Weak perennial, or annual;
loosely tufted; to 400 mm tall.

Leaf blades 40-300 mm long; 1-3
mm wide. Spikelets 4-16 mm
long; 1-3 mm wide. Inflorescence
open, pedicels slender; spikelet
outline coarsely serrate, the lem-
mas on the same side of the ra-
chilla distinctly overlapping the
lemma above, rachilla persistent, lemmas and/or paleas
breaking up from the base upwards; upper glume barely
reaching to just covering the base of the lemma directly
above, lowest lemma 1 .2-2.5 mm long; palea keels scaberu-
lous; anthers 3, 0.4— 0.5 mm long; caryopsis oblong, strong-
ly laterally flattened.

Flowering January (and a single specimen from the Cape
flowering in April). On sandy soils, gravels, clays and
loams, usually areas of high moisture, also in disturbed
areas. Locally common. Biome: Savanna and Grassland.
Throughout tropical Africa, Madagascar, India, New
Guinea, Australia and South America. Weed. Allied to £.
plana , which has an identical caryopsis but is a strong
perennial with glands on the lateral nerves of the lemmas.
Either poorly collected or only starting to invade in
southern Africa.

Description: Chippindall & Crook 1976 (166), Clayton
et al. 1970-1982 (238). Voucher: Acocks 23487, Smook
1857. PRECIS code 9902860-08400.

Eragrostis trichophora Coss. & Dur.

( -E . atherstonei Stapf) 7;

(=£. henrardii Jansen) 7.

Blousaadgras.

Slender, wiry perennial; sto-
loniferous and tufted (erect to
geniculate, rooting at the nodes);
to 600 mm tall. Leaf blades to 150
mm long; to 3 mm wide. Spikelets 3. 5-5.0 mm long;
1.0-1. 2 mm wide. Culms usually branched above; basal
sheaths glabrous or hairy, papery and the nerves rounded
and well apart at the base, below the collar of the leaf
sheaths or around the culm nodes round glandular dots, usu-
ally flushed with purple are often present; inflorescence
open, branches purplish-yellow with large prickles, spread-
ing, whorled at the base with long hairs in the axils; spike-
lets oblong to narrowly oblong, with 3-5 florets, rachilla
subpersistent, fragile in the upper part, lemmas and/or pa-
leas breaking up from the base upwards; glumes 4/5 as long
to longer than the lemmas directly above them in the intact
spikelet, lower glume wide and covering most of the lemma
above; lemma pale greenish-grey to dark grey, the apex
usually membranous and whitish; palea margins apart for
most of its length, nearly touching to touching at the apex,
keels a narrow line, smooth to scaberulous; anthers 3, 0.8
mm long.

Flowering November to May. On sand and loam, on
shallow soils and dolomite in moist places, road verges and
other disturbed or overgrazed areas. Common. Biome:
Savanna and Nama-Karoo. Southern tropical and north
Africa. Pasture (grazed by goats). Similar to E.
cylindriflora, which is annual, and E. lehmanniana, which
has the lowest inflorescence branches 1-2 and not whorled.

163

Description: Chippindall & Crook 1976 (174), De Win-
ter in Chippindall 1955 (149). Illustration: Muller 1984 (fig.
85). Voucher: Smook 6243, De Winter & Giess 6823.
PRECIS code 9902860-08500.

Eragrostis truncata Hack.

Mat-forming perennial; rhizo-
matous (rhizome short and bran-
ched), or tufted (forming raised
cushions and characteristic
rings); to 400 mm tall. Leaf blad-
es (lower) 10-40 mm long (upper
blades to 100 mm long); 1.0-1. 5
mm wide. Spikelets 5-7 mm
long; 2. 5-3.0 mm wide. Basal
sheaths with dense, long woolly hairs; spikelets completely
pallid to flushed with dark purple, spikelet rachilla very fra-
gile, breaking up from the apex downwards, the florets be-
ing shed with the rachilla internode; lowest lemma truncate;
palea keels scaberulous; anthers 3, 0.8-1. 2 mm long.

Flowering October to May. Mainly in limestone soils,
especially in and around pans. Locally common. Biome: Sa-
vanna and Nama-Karoo. Endemic. Palatable pasture (graz-
ed by game). Barely distinguishable from E. bergiana , and
an indepth study is needed in these two taxa.

Description: Stapf 1898-1900 (624), De Winter in Chip-
pindall 1955 (176). Illustration: De Winter in Chippindall
1955 (fig. 148). Voucher: Ellis 2630, Smook 3487. PRECIS
code 9902860-08600.

Eragrostis virescens Presl

Chilean love grass.

Annual; tufted (erect or geni-
culate); to 700 mm tall. Leaf blad-
es to 250 mm long; 3. 5-7.0 mm
wide. Spikelets 3. 0-4. 5 mm long;

1.0- 1. 2 mm wide. Basal leaf
sheaths glabrous; leaf blade mar-
gins eglandular, scabrid and the
midvein eglandular; inflorescence open, side branches
spreading, pedicels long and slender, spikelets tending to
be condensed to the branches; spikelets oblong, rachilla
persistent, lemmas and/or paleas breaking up fom the base
upwards; lower glume 1/3-1/2 the length of the lemma
above in the intact spikelet; lemma 1. 3-1.7 mm long; palea
keels scabrid; anthers 3, 0.2-0. 3 mm long; caryopsis ob-
long.

Flowering December to March. Sandy soils in cultivated
and disturbed areas. Locally common. Naturalized from
South America. Biome: Savanna, Grassland, Nama-Karoo.
South and North America. Weed (in gardens and lands).

Description: Hitchcock & Chase 1950 (151), De Winter
in Chippindall 1955 (154). Voucher: Smook 3835, Smook
2839. PRECIS code 9902860-08800.

Eragrostis viscosa (Retz.) Trin.

Sticky love grass.

Annual; tufted (erect and geni-
culate); to 500 mm tall. Leaf blad-
es 40-100 mm long; 2-5 mm
wide. Spikelets 2-3 mm long;

1.0- 1. 5 mm wide. Inflorescence
with sticky glands (noticable due
to particles adhering to them),
open, with the branches spreading and the spikelets distant;
spikelets with the rachilla fragile, breaking up from the apex
downwards; lemma keel glabrous, palea keels with hairs
0.4-0. 5 mm long and exserted from the lemma; anthers 3,
0. 1-0.3 mm long .

Flowering February to August. Disturbed and open plac-
es with sandy or shallow soils. Locally common. Biome:
Savanna. Northwards through east Africa to Nigeria; to
India, Thailand and Philippines, with a few records from
tropical America. Weed and indicator (of poor soil condi-
tion and overgrazing). Strong smelling when fresh. The
boundary between E. viscosa and E. tenella, which has the
inflorescences eglandular or with non-sticky glands, is not
always sharp and intermediates are found. Similar to E.
arenicola , which has the inflorescence contracted, and E.
ciliaris, which has the lemma with long hairs on the keel.

Description: Chippindall & Crook 1976 (150), Clayton
et al. 1970-1982 (206). Illustration: De Winter in Chippin-
dall 1955 (fig. 155). Voucher: Muller 1298. PRECIS code
9902860-08900.

Eragrostis volkensii Pilg.

Perennial; densely tufted (with
straggling matted wiry culms; of-
ten procumbent); 400-1200 mm
tall. Leaf blades 20-80 mm long;

1-5 mm wide. Spikelets 3. 5-7.0
mm long; 1. 5-4.0 mm wide. In-
florescence sparsely branched,
the primary branches stiffly
spreading, pedicels short; spike-
lets olive-green, rachilla persistent, lemmas and/or paleas
breaking up from the base upwards; glumes obtuse; lemma
broadly ovate (in profile), lateral nerves indistinct; palea
keels entire, winged, scaberulous; anthers 3, 1 mm long;
caryopsis narrowly ovate.

Flowering October to January. Damp soils in
mountainous areas. Infrequent. Biome: Grassland.

Northwards to east Africa, Cameroun and Zaire.

Description: Clayton et al. 1970-1982 (222). Voucher:
De Winter & Codd 216, Kluge 439. PRECIS code
9902860-09000.

Eragrostis walteri Pilg.

Perennial; hydrophyte (occas-
ionally), or tufted (with culms
erect, semi-decumbent or
floating); to 1140 mm tall (or
long). Leaf blades to 100 mm
long; to 4.5 mm wide. Spikelets
to 7 mm long; 1. 5-3.0 mm wide
(excluding awns). Culms either
straggling and matted or erect to
geniculate and separate and not matted; inflorescence nar-
row, sparsely branched, branches spreading from or appres-
sed to the main axis, pedicels short and the spikelets close
together; spikelets light green to purple, granular, rachilla
subpersistent, fragile in the upper portion; glumes acumin-
ate, upper glume tapering into a long, thick, awn-like,
acuminate apex; lemma lanceolate, acute to acuminate in
profile; palea margins not touching, keels flat, very broad
in the lower 2/3, narrowing sharply to the apex and
excurrent into a soft mucro; anthers 3, 0. 6-1.0 mm long;
caryopsis elliptic.

Flowering throughout the year. In damp, sandy and
brackish soils around seepage areas or stagnant pools or in
running water, especially that of springs and often associat-
ed with calcium carbonate. Locally common (only in a few
specific moist areas). Biome: Nama-Karoo and Desert.
Endemic. Ellis 1984 S. Afr. J. Bot. 3,6 (380) reports the first
record for the sub-family Chloridoideae of non-Kranz leaf
anatomy, which implies it utilizes the C3 photosynthetic
pathway.

Description: De Winter in Chippindall 1955 (176).
Voucher: Giess 8104, Mueller 1285. PRECIS code
9902860-09100.

164

Eriochloa Kunth

Aglycia Steud ,,Glandiloba (Raf.) Steud., Helopus Trin.,
Oedipachne Link.

Annual, or perennial; long-stoloniferous, or caespitose
to decumbent. Culms 200-1700 mm high; herbaceous;
branched above, or unbranched above. Leaf blades usually
flat. Ligule a fringed membrane (very reduced), or a fringe
of hairs.

Inflorescence of spike-like main branches (of simple or
compound racemes )\ open; espatheate. Spikelet-bearing
axes persistent.

Spikelets solitary, or in pairs; consistently in ‘long-and-
short’ combinations, or not in distinct ‘long-and-short’
combinations; supported on a peculiar, hardened, cupuli-
form ‘callus’ . Spikelets adaxial; compressed dorsiventrally;
falling with the glumes. Glumes two (but G1 very
modified); very unequal; awned (G2, when aristulate), or
awnless; very dissimilar (the lower reduced to a small
cupuliform strip adherent to the thickened rachilla
internode). Proximal incomplete florets 1; paleate, or
epaleate, palea when present fully developed to reduced;
male, or sterile.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes; rugose; becoming indurated, or not becoming
indurated (papery to crustaceous); hairless (glabrous or
apically puberulous); having the margins tucked in onto the
palea; with a clear germination flap; 5 nerved; entire;
mucronate to awned (the mucro or awn barbellate). Awns
apical; non-geniculate; much shorter than the body of the
lemma. Palea present; relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit small; hilum
short; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional. Biochemical type PCK (5
species); XyMS+. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. 30
species. Subtropical. Helophytic to mesophytic; in open
habitats (damp ground and weedy places); glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, and Cape Province. 5 indigenous species.

References. 1. Gibbs Russell. 1980. Bothalia 13: 2.
Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Lower glume produced above beadlike swelling at
base of spikelet; upper glume acute but not

mucronate or awned 2

Lower glume apparently absent, adnate to beadlike
swelling at base of spikelet; upper glume

acuminate, mucronate or short-awned 3

2(1). Lower glume a short, truncate, in-turned cuff at

base of spikelet

E.meyeriana subsp. meyeriana

Lower glume acute, 1/4-1/2 the length of the
spikelet . . E. meyeriana subsp. grandiglumis
3(1). Spikelets 2. 0-2. 5 mm long; inflorescence branches
few, appressed to main axis . E. parvispiculata
Spikelets (2.5-)3.0-5.0 mm long; inflorescence
branches several to many, ascending, not

appressed to main axis 4

4(3). Lower floret sterile, lacking a palea; plant annual

E. fatmensis

Lower floret staminate, with a palea almost as long

as lemma; plant annual or perennial 5

5(4). Plant perennial, robust; spikelets 3^4- mm long;

upper glume mucronate or with an awn-point to
1 mm long; upper lemma with mucro 0.3-0. 7 mm

long; Transvaal, Swaziland, Natal

E. stapfiana

Plant annual; spikelets 3.5-5.0 mm long; upper
glume with an awn-point 1-3 mm long;
Botswana E. macclounii

Fig. 93. Eriochloa meyeriana subsp. meyeriana

Eriochloa fatmensis (Hochst. & Steud.) Clayton

( -E . nubica (Steud.) Hack. &

Stapf ex Thell.) 2.

Annual; 100-1200 mm tall
(culms erect or geniculate). Leaf
blades 30-300 mm long; 2-10
mm wide. Spikelets (2.5-)3.0-5.0
mm long. Inflorescence branches
several to many, ascending; lower
glume apparently absent, upper glume with a short awn;
lower floret sterile; palea absent.

Flowering January to April. Wet places, usually on black
clay, but also weedy by ephemeral water on other soils. In-
frequent. Through tropical Africa and Arabia to India.

Description: Clayton et al. 1970—1982 (571). Voucher;
Merxmueller & Giess 1560. PRECIS code 9901020—00050.

Eriochloa macclounii Stapf

Annual; tufted; to 1200 mm
tall. Leaf blades 80-600 mm
long; 3-12 mm wide. Spikelets
3. 5-5.0 mm long. Inflorescence
branches several to many,
ascending; raceme rachises with
long hairs; lower glume
apparently absent; upper glume
with an awn-point 1-3 mm long;
lower floret staminate, with palea almost as long as lemma;
female-fertile lemma with mucro 1-2 mm long.

Flowering April. Floodplain grassland. Rare (in
southern Africa). Biome: Savanna. To Tanzania and
Mozambique. Similar to the more robust E. stapfiana.

Description: Clayton et al. 1970-1982 (570). Voucher:
P.A. Smith 4299. PRECIS code 9901020-00075.

165

Eriochloa meyeriana (Nees) Pilg. subsp. grandiglumis

(Stent & Rattray) Gibbs Russell

{=Panicum meyerianum Nees
var. grandeglume Stent &

Rattray) 1.

Perennial. Similar to the
typical subspecies but with the
lower glume acute, extending
1/4- 1/2 the length of the spikelet.

Flowering October to June.

Lowveld riverbanks and floodplains. Rare. Biome: Savan-
na. Also in Zimbabwe. Intergrades with subsp. meyeriana,
which has a shorter lower glume.

Description: Clayton et al. 1970-1982 (569). Voucher:
Codd 5421. PRECIS code 9901020-00095.

Eriochloa meyeriana (Nees) Pilg. subsp. meyeriana

Fig. 93. PI. 81.

( =E . borumensis sensu
Hack., non Stapf) 2; ( =Panicum
meyerianum Nees var.
meyerianum) 2.

Robust perennial (culms geni-
culate, rooting at nodes below);

300-1500 mm tall. Leaf blades
50-250 mm long; 3-15 mm wide.

Spikelets 2. 5-3. 5 mm long. Lower glume produced above
the beadlike swelling as a short, truncate in-turned cuff;
upper glume acute, not mucronate or awned.

Flowering October to May. Riverbanks and wet places,
on sandy or clay soils. Locally common. Biome: Savanna.
To tropical Africa. Intergrades with both subsp.
grandiglumis and E. stapfiana.

Description: Clayton et al. 1970-1982 (569). Voucher:
De Winter & Codd 544. PRECIS code 9901020-00100.

Eriochloa parvispiculata C.E. Hubb.

Perennial; tufted; 300-1200
mm tall. Leaf blades 80-300 mm
long; 3-10 mm wide. Spikelets
2. 0-2. 5 mm long. Inflorescence
branches few, appressed to main
axis; lower glume apparently
absent; upper glume with a
mucro.

Flowering January to April.

Riverbanks and floodplain pans. Rare. Biome: Savanna. To
tropical east Africa. Intergrades with E. stapfiana , which
has longer spikelets; some specimens are difficult to
distinguish from the annual E. fatmensis.

Description: Clayton et al. 1970-1982 (570). Voucher:
Liebenberg 4390. PRECIS code 9901020-00300.

Eriochloa stapfiana Clayton

(=£. borumensis sensu Stapf,
non Hack.) 2.

Robust perennial; tufted;

600-1700 mm tall. Leaf blades
40-200 mm long; 3-8 mm wide.

Spikelets 3^1 mm long. Inflores-
cence branches several to many,
ascending; raceme rachises with
short hairs; lower glume apparently absent; upper glume
mucronate or with an awn-point to 1 mm long; lower floret
staminate, with palea almost as long as lemma; female-
fertile lemma with mucro 0.3-0. 7 mm long.

Flowering October to May. Riverbanks and wet places
on heavy soils. Infrequent. Biome: Savanna. To tropical
east Africa. Intergrades with E. meyeriana subsp. meyeriana,
which has a shorter mucronate tip on the upper lemma.

Description: Clayton et al. 1970-1982 (569). Voucher:
De Winter & Codd 552. PRECIS code 9901020-00400.

Eriochrysis P. Beauv.

Plazerium Kunth.

Perennial; caespitose. Culms 400-1200 mm high; herba-
ceous; unbranched above. Leaf blades linear; usually flat,
or folded (rarely). Ligule a fringed membrane, or a fringe
of hairs. Plants bisexual, with bisexual spikelets. The
spikelets of sexually distinct forms on the same plant ; ho-
momorphic (but the pedicellate spikelets smaller).

Inflorescence of spike-like main branches (spiciform
'racemes', in a raceme or panicle, with tawny-red hairs)',
contracted (narrow); non-digitate\ espatheate; not com-
prising ‘partial inflorescences' and foliar organs. Spikelet-
bearing axes ‘racemes’; solitary; with substantial rach-des;
disarticulating at the joints.

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets female-only . Female-fertile spikelets compressed
dorsiventrally; falling with the glumes. Glumes two; more
or less equal; awnless; very dissimilar (G2 thinner, not 2-
keeled). Proximal incomplete florets /; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire to incised (truncate to serrate);
awnless. Palea absent. Lodicules 2; fleshy; glabrous.
Stamens 3 (rudimentary in the pedicellate spikelets). Ovary
glabrous. Hilum short; embryo large.

Fig. 94. Eriochrysis pallida

166

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Andropogoninae. 7
species. Tropical America and tropical Africa. Helophytic;
in open habitats (in swamps and moist places); glycophytic.
Botswana, Transvaal, Swaziland, Natal, and Cape Province.
2 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.
Species treatment by G.E. Gibbs Russell.

1(0). Inflorescence hairs longer than the spikelets; sessile

spikelets 3. 5-5.0 mm long E. pallida

Inflorescence hairs shorter than the spikelets; sessile
spikelets 5-6 mm long E. brachypogon

Eriochrysis brachypogon (Stapf) Stapf

(=£. brachypogon (Stapf) Stapf
subsp. australis J.G. Anders.) 1.

Perennial; tufted; 600-900
mm tall. Leaf blades to 300 mm
long; 1-3 mm wide. Spikelets
(sessile) 5-6 mm long (pedicel-
late shorter). Golden callus hairs
shorter than spikelets.

Flowering November to March. Vleis and riverbanks.
Rare and conservation status not known. Biome: Grassland.
Tropical Africa.

Description: Clayton et al. 1970-1982 (707).

Illustration: Clayton et al. 1970-1982 (fig. 162). Voucher:
Compton 30488. PRECIS code 9900420-00100.

Eriochrysis pallida Munro

Perennial; tufted; 400-900
mm tall. Leaf blades 100-240
mm long; 1-4 mm wide. Spike-
lets (sessile) 3. 5-5.0 mm long
(pedicellate shorter). Golden
callus hairs longer than spikelets.

Flowering July to June. Vleis
and riverbanks. Infrequent.

Biome: Savanna. Tropical Africa.

Description: Chippindall 1955 (475), Clayton et al.
1970-1982 (706). Illustration: Chippindall 1955 (fig. 391).
Voucher: Killick 251. PRECIS code 9900420-00200.

Eulalia Kunth

Puliculum Haines.

Perennial (rarely annual); caespitose, or decumbent.
Culms 100-1500 mm high; herbaceous; unbranched above.
Leaf blades linear; flat. Ligule an unfringed membrane, or
a fringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets homomorphic.

Inflorescence of spike-like main branches (very hairy or
silky, often brown or purple ); digitate or subdigitate
( usually with a short axis)', espatheate; not comprising
‘partial inflorescences’ and foliar organs. Spikelet-bearing
axes ‘racemes’ (spiciform); with very slender rachides', dis-
articulating at the joints. ‘Articles’ disarticulating
obliquely.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets hermaphrodite. Female-fertile spikelets com-
pressed dorsiventrally; falling with the glumes (the
pedicellate falling from the pedicel, the sessile falling with
the joint and pedicel). Glumes two; more or less equal;

Fig. 94. PI. 82.

Fig. 95. Eulalia villosa

167

awned (rarely, G1 is bilobed or 2-awned), or awnless; very
dissimilar (both villous, rigid to coriaceous; the lower
flattened to depressed on the back and more or less
bicarinate, the upper naviculate). Proximal incomplete
florets present or absent; when present 1; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire, or incised; awned. Awns 1; from
the sinus (usually); geniculate; much longer than the body
of the lemma. Palea present, or absent; when present very
reduced. Lodicules 2; fleshy; ciliate, or glabrous. Stamens
3 (rarely 2). Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x - 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 30 species. Tropical and
subtropical Africa, Asia, Australia. Helophytic to meso-
phytic; in open habitats (grassland, sometimes in moist
places); maritime-arenicolous ( E . ridleyi), or glycophytic.
Namibia, Botswana, Transvaal, Swaziland, Natal, and Cape
Province. 2 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Hairs on racemes white; leaf sheaths hairy; spikelets

5-7 mm long; culms erect E. villosa

Hairs on racemes golden - brown; leaf sheaths
glabrous; spikelets 3. 5-4.0 mm long; culms often
decumbent E. aurea

Eulalia aurea (Bory) Kunth

(=£. geniculata Stapf) 1.

Perennial; creeping rhizomat-
ous; to 1000 mm tall. Leaf blades
30-150 mm long; 3-6 mm wide.

Spikelets 3.5^4 mm long (sessile
and pedicellate alike). Culms
often decumbent; sheaths
glabrous; raceme hairs golden-
brown.

Flowering December to March. Riverbanks and
floodplains. Infrequent. Biome; Savanna. Tropical Africa,
Reunion and Australia.

Description: Chippindall 1955 (486), Clayton et al.
1970-1982 (713). Illustration: Chippindall 1955 (pi. 16).
Voucher: De Winter & Marais 4484. PRECIS code
9900530-00150.

Eulalia villosa (Thunb.) Nees ^ ^ p^ ^

Perennial; tufted; 300-1400
mm tall. Leaf blades 50-250 mm
long; 3-8 mm wide. Spikelets 5-7
mm long (sessile and pedicellate
alike). Culms erect; sheaths hairy;
raceme hairs white; lemma awns
15-20 mm long.

Flowering September to May.

Open grassland on hillsides. In-
frequent. Biome: Savanna and Grassland. Through eastern
tropical Africa and Madagascar to India. Resembles hairy-
leaved individuals of Ischaemum fasciculatum in the
inflorescence form and reddish colour, but in that species
the lemma awns are only about 5-10 mm long.

Description: Chippindall 1955 (485), Clayton et al.
1970-1982 (713). Illustration: Chippindall 1955 (fig. 397),
Clayton et al. 1970-1982 (fig. 165). Voucher: Devenish
1282. PRECIS code 9900530-00200.

Eustachys Desv.

Chloroides Regel, Langsdorffia Regel, Schultesia
Spreng.

Perennial; caespitose. Culms 200-1000 mm high; herba-
ceous. Leaf blades linear; flat, or folded. Ligule a fringe of
hairs.

Fig. 96. Eustachys paspaloides

168

Inflorescence of spike-like main branches ( spikes or
spicate racemes)', digitate or subdigitate', espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; 1.7-5 mm long; compressed
laterally to not noticeably compressed; disarticulating
above the glumes. Glumes two; relatively large (thinly
membranous); very unequal, or more or less equal; decid-
edly shorter than the adjacent lemmas; awned (the upper
only, from below its apex)', very dissimilar (membranous,
G2 broader and awned). Incomplete florets distal to the
female-fertile florets (usually 2, greatly reduced), merely
underdeveloped; proximal incomplete florets absent.

Female-fertile florets I . Lemmas decidedly firmer than
the glumes (firmly membranous to papery, dark brown); 3
nerved; entire, or incised (notched); awnless, or mucronate.
Palea present. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small; ellipsoid; hilum short; pericarp
fused; embryo large.

Photosynthetic pathway and related features. C4;
NAD-ME (distichophylla)', XyMS+. PCR sheath outlines
uneven, or even. PCR sheath extensions absent. PCR cell
chloroplasts centrifugal/peripheral, or centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 10
species. Tropical America, West Indies, tropical and South
Africa. Mesophytic; in open habitats (savanna, on a variety
of soils); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Eustachys paspaloides (Vahl) Lanza & Mattei

(-E. mutica auctt.) 1 .

Red Rhodes grass, bruinhoen-
derspoor.

Perennial; rhizomatous and
tufted (erect or geniculately
ascending); 200-950 mm tall.

Leaf blades 20-180 mm long; 2-5
mm wide. Spikelets 1 .5-2.5 mm long. Leaf sheaths strongly
compressed, blades folded, with blunt apices; spikes
(3— )4— 1 0(— 15), 50-150 mm long; spikelets golden-brown;
lower glume ovate, boat-shaped; upper glume oblong-
elliptic with awn 0.5-1. 5 mm long; lemma 1. 5-2.4 mm
long, with or without a mucro up to 1 mm long.

Flowering October to May. Sandy and stony soils,
occasionally on clay. Common. Biome: Fynbos, Savanna,
and Grassland. Tropical Africa to Arabia. Very closely
related to Chloris, from which it is distinguished by broader
glumes, a short awn on the upper glumes and dark brown
almost awnless female-fertile lemmas.

Description: Chippindall & Crook 1976 (23), Chippin-
dall 1955 (194), Clayton et al. 1970-1982 (335).
Illustration: Chippindall 1955 (fig. 170), Clayton et al.
1970-1982 (fig. 95). Voucher: Smook 3124. PRECIS code
9903020-00200.

Festuca L.

Amphigenes Janka, Anatherum Nabelek, Argillochloa
Weber ,Bucetum Parnell , Drymochloa Holub , Drymonaetes
Fourr., Festucaria Fabric., Gnomonia Lunell, Gramen
Krause, Hellerochloa Rauschert, Hesperochloa (Piper)
Rydberg, Leiopoa Ohwi, Lojaconoa Gand., Nabelekia
Roshev., Pseudobromus K. Schum., Wasatchia M.E. Jones.

Perennial', long-rhizomatous, or long-stoloniferous, or
caespitose, or decumbent. Culms 20-2000 mm high; herba-
ceous; unbranched above. Sheath margins joined, or free.

Leaf blades linear to linear-lanceolate; narrow, flat, or
folded, or rolled (convolute or involute). Ligule an
unfringed membrane (sometimes ciliolate).

Inflorescence paniculate', open, or contracted (rarely);
espatheate. Spikelet-bearing axes persistent.

Spikelets not secund', 3-20 mm long; compressed
laterally; disarticulating above the glumes. Hairy callus
absent. Glumes two; very unequal ; markedly shorter than
the spikelets; awnless; similar (usually narrow to ovate-
lanceolate). Lower glume I nerved. Incomplete florets distal
to the female-fertile florets, merely underdeveloped;
proximal incomplete florets absent.

Female-fertile florets 2-14 (rarely 1). Lemmas similar
in texture to the glumes; non-carinate', 3-7 nerved; entire,
or incised; awnless, or mucronate, or awned. Awns when
present 1; from the sinus, or apical; non-geniculate; much
shorter than the body of the lemma (usually), or about as
long as the body of the lemma (sometimes — rarely
somewhat longer). Palea present; relatively long. Lodicules
2; membranous', ciliate, or glabrous. Stamens 3. Ovary
glabrous, or hairy. Fruit small, or medium sized, or large;
fusiform, or ellipsoid; hilum long-linear (usually about as
long as the grain, but sometimes elliptical and only half as
long)', embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. 360 species (or
more). Worldwide temperate & mountains. Helophytic
(rarely), or mesophytic (mostly), or xerophytic (rarely);
maritime-arenicolous, or halophytic, or glycophytic.
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. Indigenous species ( 8) , naturalized
species (1).

Intergeneric hybrids with Vulpia ( X Festulpia Melderis
ex Stace & Cotton), with Lolium ( X Festulolium Aschers.
& Graebn.) and supposedly with Bromus (X Bromofestuca
Prodan — Bull. Grad. Bot. Univ. Club 16, 93 (1936)).

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton.
1985. Kew Bull. 40: 727. 3. Linder. 1986. Bothalia 16: 61.
4. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Spikelets 1 -flowered with a prominent rachilla
extension; lemma extending into an awn 1 0-20 mm

long F. africana

Spikelets 3-10-flowered, lacking a rachilla extension;
lemmas acute or with an awn to 6 mm long ... 2
2(1). Panicle almost candelabrum-shaped, lowest branches
about as long as the central axis; branches rigid,
straight and bare, bearing a few spikelets near the

tip F. longipes

Panicle open or contracted, lowest branches less than
half the length of the central axis; branches
flexuous, either bare in the lower third or bearing

spikelets from the base 3

3(2). Old leaf sheaths persistent, breaking into fibres and
forming a dense protective layer around young

shoots 4

Old leaf sheaths not persistent or breaking into fibres

6

4(3). Leaf sheaths covered with velvety hairs at the base;
base of culms bulbous or thickened; panicle dense,

almost spike-like, branches rigid F. scabra

Leaf sheaths glabrous; base of culms not bulbous or
thickened; panicle open, branches flexuous ... 5
5(4). Leaf blades filiform, often inrolled, 0.2-2. 0 mm wide;

ligule shorter than 1 mm; basal fibres fine

F. caprina

Leaf blades expanded or inrolled, 3-10 mm wide;

ligule 3-12 mm long; basal fibres coarse

F. costata

6(3). Leaf auricles very well developed, to 5 mm wide . .

F. elatior

Leaf auricles absent or poorly developed, less than 1

mm wide 7

169

7(6). Lemma awned, awn 1-4 mm long; panicle branches
flexuous, bearing fewer than 5 spikelets near the

tip; leaves cauline F. dracomontana

Lemma awnless or minutely awned; panicle branches
fairly rigid, either bearing spikelets from the base
or clustered at the tips; leaves basal or cauline . 8
8(7). Leaf blades inrolled, mostly basal; panicle pyramidal
and open; branches with spikelets in the upper half

F. killickii

Leaf blades expanded, mostly cauline; panicle narrow
and contracted; branches widely spaced, spikelets
solitary on long pedicels along the central axis, with
short branches bearing up to 3 spikelets scattered
in between F. vulpioides

Fig. 97. Festuca costata

Festuca africana (Hack.) Clayton

(-Pseudobromus africanus
(Hack.) Stapf) 2;

{=P seudobromus silvaticus K.

Schum.) 2.

Perennial; rhizomatous; 500-
1 200(— 1 600) mm tall. Leaf blades
250-500 mm long; 8-15 mm
wide. Spikelets 7-9 mm long.

Leaves cauline; ligule to 8 mm long; panicle scanty,
branches delicate and flexuous with spikelets solitary at the
tips; spikelets 1 -flowered with a prominent rachilla
extension and a rudimentary floret; glumes unequal; lemma
extending into a long straight awn 10-20 mm long.

Flowering January to April (occasionally in other
months). Open patches in forests, usually in the shade. In-
frequent. Biome: Forest. Northwards to Kenya. Easily
confused with another forest species, Stipa dregeana, which
has glumes equal and longer than the lemmas and lemmas
hairy with a bent and twisted awn. Distinguished from other
South African Festuca species, which have spikelets 3-9-
flowered and usually lack a rachilla extension.

Description: Linder (15), Stapf 1898-1900 (763), Chip-
pindall 1955 (61). Illustration: Chippindall 1955 (fig. 34).
Voucher: Schweickerdt 1563. PRECIS code 9904170-
00075.

Festuca caprina Nees

Bokbaardgras.

Perennial; rhizomatous and
tufted (densely); 250-1000 mm
tall. Leaf blades 40-250 mm
long; filiform, to 1.5 mm wide.

Spikelets 7-15 mm long. Old leaf
sheaths persisting as fine fibres;
ligule shorter than 1 mm; leaves
usually not more than half the culm length; panicle open,
lax, 50-200 mm long; spikelets 4-6(-9)-flowered; lemmas
awned, awn 1-4 mm long.

Flowering September to March. Moist or wet areas near
vleis in high altitude mountain grassveld. Common to local-
ly dominant. Biome: Grassland and Afromontane.

Northwards to Tanzania. Closely related to F. scabra ,
which has flat leaf blades, an almost spikelike panicle and
swollen culm bases, and to/7, costata , which has expanded
or inrolled leaf blades, a longer ligule and coarse sheath
fibres. The different varieties previously recognized are not
upheld because of the variability in the species.

Description: Linder (13), Stapf 1898-1900 (719), Chip-
pindall 1955 (55), Clayton et al. 1970-1982 (62).
Illustration: Chippindall 1955 (fig. 25). Voucher: Hoener
2114. PRECIS code 9904170-00200.

Festuca costata Nees

Tussock fescue, polswenk-
gras.

Tough, erect perennial; rhizo-
matous (rhizome short and stout
or long and deeply burrowed);

600-1700 mm tall. Leaf blades
700-1000 mm long; 3. 5-6.0 mm
wide. Spikelets 10-20 mm long.

Leaf sheaths persistent, breaking into coarse fibres that
form a protective layer around the young shoots; ligules
3-12 mm long; panicle open, 120-240 mm long; spikelets
3-7-flowered; lemma acute or minutely awned.

Flowering September to January. Moist places in high
altitude mountain grassland, encouraged by frequent
burning. Common to locally dominant. Biome:Afromontane
Grassland. Endemic. Closely related to F. caprina , which

Fig. 97. PI. 85.

170

has filiform leaf blades and fine sheath fibres, and to F.
scabra, which has bulbous culm bases, velvet-hairy leaf
sheaths and a dense, almost spikelike, panicle. The different
varieties previously recognized are not upheld because of
the variability in the species.

Description: Chippindall & Crook (210), Linder (11),
Stapf 1898-1900 (721), Chippindall 1955 (56), Clayton et
al. 1970-1982 (59). Illustration: Chippindall 1955 (fig. 28).
Voucher: Mohle 47. PRECIS code 9904170-00500.

Festuca dracomontana Linder

Perennial; rhizomatous; 500-
800 mm tall. Leaf blades 80-200
mm long; 2-8 mm wide. Spike-
lets 10-12 mm long. Leaves cau-
line, blades expanded; ligules
shorter than 1 mm; panicle open,
to 250 mm long, branches flexu-
ous, bare for the longest part, with

1-5 spikelets near the tips;
spikelets 3-7-flowered; lemmas awned, awn 2-4 mm long.

Flowering October. Sour grassveld in high mountains.
Rare. Biome: Grassland. Endemic.

Description: Linder 1986 Bothalia 16,1 (59). Voucher:
Du Toil 2714. PRECIS code 9904170-00725.

Festuca longipes Stapf

Perennial; rhizomatous (rhi-
zome long and slender); 300-750
mm tall. Leaf blades 100-300
mm long; 2-6 mm wide. Spike-
lets 8-12 mm long. Ligules 2-3
mm long; panicle almost
candelabrum-shaped, 150-330
mm long, branches rigid,
flattened on the inner side, borne
in remote pairs, unbranched and bare for most of their
length, bearing 1-6 spikelets at the tips, lower branches
almost as long as the central axis; spikelets closely 3-6-
flowered; glumes and lemmas awnless.

Flowering November to April. Moderate to steep grassy
slopes and sandstone ridges, often on the edges of forests
in partial shade, at altitudes higher than 1 500 m. Infrequent.
Biome; Grassland. Endemic. The shape of the panicle
distinguishes this species from other South African Festuca
species, which have open or contracted panicles with
branches flexuous and less than half the length of the
central axis.

Description: Linder (10), Stapf 1898-1900 (721), Chip-
pindall 1955 (58). Voucher: Mullins PRE 56847. PRECIS
code 9904170-00900.

Festuca elatior L.

Festuca scabra Vahl

( -F . arundinacea Schreb.) 3.

Meadow fescue, English blue-
grass.

Perennial; rhizomatous and
tufted; 800-2000 mm tall. Leaf
blades 100-600 mm long; 3-12
mm wide. Spikelets 10-18 mm
long. Leaf sheaths not breaking into fibres, sheaths
glabrous; leaf auricles well developed, to 5 mm wide;
ligules to 2 mm long; panicle lanceolate to ovate, 100-500
mm long, nodding; spikelets 3-10-flowered; lemmas acute
or awned, awn to 4 mm long.

Flowering September to April. Disturbed places near
streams, reservoirs and on roadsides. Infrequent. Natural-
ized from Europe and temperate Asia. Biome: Fynbos to
Grassland. Widely introduced to tropical countries. Winter
pasture (and in experimental cultivation). The unusually
large leaf auricles are unique for the southern African
Festuca species.

Description: Bor 1985 (1730), Hitchcock & Chase 1950
(168), Chippindall 1955 (56), Clayton et al. 1970-1982
(58). Illustration: Chippindall 1955 (fig. 27), Hitchcock &
Chase 1950 (fig. 78). Voucher: Smook 5977. PRECIS code
9904170-00750.

Festuca killickii K. -O'Byrne

Erect, coarse perennial; rhizo-
matous and tufted; 500-950 mm
tall. Leaf blades erect, 200-600
mm long; inrolled, 3-6 mm wide.

Spikelets 6-9 mm long. Leaves
and sheaths glabrous; ligules

2-3(-4) mm long; panicle
pyramidal, to 200 mm long,

20-60 mm wide, branches bare
for lower half; spikelets shortly pedicelled, 4-6-flowered;
lemmas acute or with awn shorter than 0.5 mm.

Flowering December to February. Subalpine grassveld
at high altitudes up to 3000 m, on Cave sandstone and
basalt, amongst boulders, along streams and on ledges. Lo-
cally common to locally dominant. Biome: Grassland.
Endemic.

Description: Kennedy-O’Byrne 1963 Kew Bull. 16,1
(461), Linder (13). Illustration: Kennedy-O’Byme 1963
(fig. 1). Voucher: Du Toit 2320. PRECIS code

9904170-00800.

Munnik fescue, Munnik
swenkgras.

Perennial; rhizomatous (rhi-
zome long or oblique); 300-1000
mm tall. Leaf blades 50-300 mm
long; inrolled or expanded and to
10 mm wide. Spikelets 7-15 mm
long. Plants dioecious; culm base
bulbous at maturity; leaf sheaths velvet-hairy at the base,
old sheaths splitting into fine fibres; ligules 2-7 mm long;
panicle 50-300 mm long, 10-30 mm wide, narrow,
contracted, sometimes spike-like or interrupted; spikelets

3-7-flowered; lemmas awnless or minutely awned.

Flowering September to February. Sandy soils in
undisturbed, high altitude mountain grassveld and
extending into Valley Bushveld, often in moist places and
partial shade, stimulated by fire. Common to locally domin-
ant. Biome: Fynbos, Savanna, Grassland, and Succulent
Karoo. Endemic. Well eaten natural or cultivated pasture.
Morphologically very variable. If the bulbous culm bases
and fibrous leaf sheaths are not distinct in young plants, it
can be distinguished by the velvet-hairy basal part of the
leaf sheaths. Very closely related to F . caprina and F.
costata , which have glabrous leaf sheaths and open lax pan-
icles.

Description: Linder (8), Stapf 1898-1900 (722), Chip-
pindall 1955 (56). Illustration: Chippindall 1955 (fig. 26).
Voucher: Hoener 1923. PRECIS code 9904170-01000.

Festuca vulpioides Steud.

Perennial; tufted (or
sprawling); 500-1000 mm tall.

Leaf blades 100-250 mm long;

3-7 mm wide. Spikelets 15-20
mm long. Leaf blades cauline,
expanded; ligules shorter than 1
mm; panicle narrow, to 300 mm
long; spikelets borne either
solitary on long pedicels or on
short branches with up to five widely spaced spikelets;
spikelets 5-8-flowered; lemmas acute or minutely awned.

Flowering January. Scattered tufts at high altitudes in
the Dohne Sourveld. Rare. Locally common. BiomefGrass-
land. Endemic. The only F . vulpioides specimen at PRE,
Acocks 20228, has much smaller spikelets, 8-14 mm, than
the length given for this species.

171

Description: Linder (6), Stapf 1898-1900 (720).
Voucher: Acocks 20228. PRECIS code 9904170-01 100.

Fingerhuthia Nees

Lasiotrichos Lehm.

Perennial (or rarely annual in desert areas); caespitose.
Culms 50-1170 mm high; herbaceous; unbranched above.
Leaf blades long linear; flat, or folded. Ligule a fringe of
hairs. The spikelets of sexually distinct forms on the same
plant (the lowest spikelets sometimes barren), or all alike
in sexuality.

Inflorescence a single raceme, or paniculate (to 120 mm
long)\ densely contracted; espatheate. Spikelet-bearing axes
persistent.

Female-fertile spikelets solitary; 4-7 mm long; com-
pressed laterally (strongly); falling with the glumes
( disarticulating from persistent pedicels)-, not disarticu-
lating between the florets. Glumes two; more or less equal;
much exceeding the spikelets; awned, or awnless (shortly
awned or mucronate); similar (narrow, folded, thin). Incom-
plete florets distal to the female-fertile florets, merely un-
derdeveloped (male or rudimentary); incomplete florets
absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (rather firm);
without a germination flap; 5 nerved, or 7 nerved (rarely 3
nerved); entire; mucronate (median nerve excurrent). Palea
present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit ellipsoid.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 2
species. Southern Africa, Afghanistan and Arabia. Helo-
phytic, or mesophytic, or xerophytic; in open habitats;
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 2 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

1 (0). Glumes with keels densely hairy; lemma tips rounded;
plant base not robust; rhizomes slender; basal leaf

sheaths dull; growing in well-drained soils

F. africana

Glumes with keels sparsely hairy; lemma tips
acuminate; plant base robust; rhizomes very well
developed; basal leaf sheaths glossy; growing in

waterlogged and poorly-drained soils

F. sesleriiformis

Fingerhuthia africana Lehm.

Vingerhoedgras, thimble
grass.

Perennial (sometimes annual);
rhizomatous; 100-910 mm tall.

Leaf blades 25-400 mm long; 2-4
mm wide. Spikelets 4. 0-5. 5 mm
long. Panicle spike-like, 15-50
mm long; glumes with stiff dense
hairs on the keels; lemma apex rounded.

Flowering September to May. Well-drained sandy or
gravelly soils, often on limestone outcrops. Common.
Biome: Savanna, Grassland, Nama-Karoo, Succulent
Karoo, and Desert. Disjunct distribution between the FSA
area and Afghanistan and Arabia. Pasture.

Fig. 98. PI. 86.

Description: Stapf 1898-1900 (691), Chippindall 1955
(207). Illustration: Chippindall 1955 (fig. 184 (spikelet).
Voucher: Smook 2887. PRECIS code 9903710-00100.

Fingerhuthia sesleriiformis Nees

Vingerhoedgras, thimble
grass.

Perennial; rhizomatous and
tufted (forming large tussocks);

300-1170 mm tall. Leaf blades
1 20-240 mm long; 3-5 mm wide.

Spikelets 5-6 mm long. Rhi-
zomes robust and well developed;
panicle spike-like, to 80 mm long; glumes sparsely hairy
on the keels; lemma apex acuminate.

Flowering November to April. Black clay in vleis or
clayey soils near rivers. Common (often in dense, pure
stands). Biome: Savanna, Grassland, and Nama-Karoo.
Endemic. Erosion control, or pasture, or domestic use -
(brooms).

Description: Stapf 1898-1900 (692), Chippindall 1955
(207). Illustration: Chippindall 1955 (fig. 183). Voucher:
Smook 5886. PRECIS code 9903710-00200.

Fig. 98. Fingerhuthia africana

172

Gastridium P. Beauv.

Annual ; caespitose (or solitary culms). Culms 100-600
mm high; herbaceous. Leaf blades linear; flat. Ligule an
unfringed membrane.

Inflorescence paniculate', contracted; espatheate.
Spikelet-bearing axes persistent.

Spikelets 3-6.5 mm long; compressed laterally; disartic-
ulating above the glumes. Glumes two (swollen, globular
and more or less cartilaginous at the base, membranous
above); relatively large; more or less equal (the lower
somewhat longer); long relative to the adjacent lemmas;
conspicuously ventricose ; awnless; similar. All florets
female-fertile, proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the

Fig. 99. Gastridium phleoides

glumes; 5 nerved; incised (more or less dentate); mucronate
to awned (the midvein usually excurrent). Awns when
present 1; dorsal; geniculate; much shorter than the body
of the lemma, to much longer than the body of the lemma.
Palea present; relatively long. Lodicules 2; membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit small; hilum
short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 2 species.
Canaries, western Europe, Mediterranean. Mesophytic to
xerophytic; in open habitats (grassy places and arable land).
Cape Province. 1 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past.
Species treatment by G.E. Gibbs Russell.

Gastridium phleoides (Nees & Meyen) C.E. Hubb.

Fig. 99. PI. 87.

Annual; 100-600 mm tall.

Leaf blades to 15 mm long; 1-4
mm wide. Spikelets 5-7 mm long.

Glumes swollen at base around
floret; panicle narrow and
spike-like.

Flowering October to Decem-
ber. Open veld. Rare. Naturalized
from the Mediterranean. Biome:

Fynbos. Widely introduced, originally from north Africa
and the Mediterranean region. Only known from Cape
Town, Stellenbosch and Wellington.

Description: Chippindall 1955 (96), Clayton et al.
1970-1982 (100). Illustration: Chippindall 1955 (fig. 67),
Clayton et al. 1970-1982 (fig. 34). Voucher: Loxton 237.
PRECIS code 9902480-00100.

Hackelochloa Kuntze

Sometimes included in Rytilix.

Annual', caespitose. Culms (50-)300-1000 mm high;
herbaceous; branched above. Leaf blades linear-lanceolate',
flat. Ligule a fringed membrane. Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant; overtly heteromorphic (the pedicellate
spikelets narrowly ovate and winged, herbaceous).

Inflorescence paniculate (the numerous ‘racemes’
solitary in their spathes, usually in fascicles)', spatheate', a
complex of ‘partial inflorescences’ and intervening foliar
organs; spikelet-bearing axes spikelike; with substantial
rachides; disarticulating at the joints (the sessile spikelets
falling with the joint and the pedicellate spikelet). ‘Articles’
with a basal callus-knob.

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; these pedicellate/sessile. Pedicels of the
‘pedicellate’ spikelets discernible, but fused with the rachis.
The sessile spikelets hermaphrodite. The ‘pedicellate’
spikelets hermaphrodite (the glumes herbaceous, similar;
lower lemma present or absent), or male-only, or sterile.
Female-fertile spikelets 1-3 mm long; compressed dorsi-
ventrally (to globose); falling with the glumes. Glumes two;
relatively large; difference in glume length not great, but
G2 hooded by Gl; awnless; very dissimilar (lower
cartilaginous, globose, reticulate-pitted; upper thinner and
embedded in the axis). Proximal incomplete florets 1 ;
epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline, flimsy); entire; awnless. Palea present, or
absent; when present relatively long, or conspicuous but
relatively short, or very reduced. Lodicules fleshy. Stamens
3. Ovary glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 7 (?). Panicoideae; Andropogonodae; Andro-
pogoneae; Rottboelliinae. 2 species. Tropics, southern
China and southern U.S.A. Helophytic to mesophytic; in
open habitats (grassland and disturbed ground);
glycophytic. Transvaal. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA. 3. Veldkamp. 1986. Blumea 31:
281.

Species could be transferred to Mnesithea Kunth.

Species treatment by G.E. Gibbs Russell.

173

Fig. 100. Hackelochloa granularis

Hackelochloa granularis (L.) Kuntze

Fig. 100. PI. 88.

Lizardtail grass.

Annual; 50-1000 mm tall.

Leaf blades 20-150 mm long;

6-12 mm wide. Spikelets (ses-
sile)!.0-1. 5 mm long (pedicellate
slightly longer). Sessile spikelet
globose, lower glume pitted and
tuberculate.

Flowering March to April. Disturbed places. Rare (in
South Africa). Throughout tropics. Weed (ruderal in the
tropics).

Description: Chippindall 1955 (523), Clayton et al.
1970-1982 (849). Illustration: Chippindall 1955 (fig. 418),
Clayton et al. 1970-1982 (fig. 200). Voucher: Parker
8 — 4— 1 959. PRECIS code 9900220-00100.

Hainardia Greuter

Monerma auctt., non (Willd.) Coss & Dur.

Annual ; caespitose. Culms 50-400 mm high; herba-
ceous; branched above. Sheath margins free. Leaf blades
linear; flat, or rolled (convolute). Ligule an unfringed
membrane.

Inflorescence a single spike (with a hard, cylindrical,
articulated rachis, the spikelets embedded in alternate
notches)', espatheate. Spikelet-bearing axes disarticulating;
disarticulating at the joints.

Spikelets solitary; distichous; 4-8 mm long; compressed
dor siventr ally, falling with the glumes. Glumes one per
spikelet (the G2); relatively large (firm); long relative to the
adjacent lemmas; awnless. All florets female-fertile;
proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes (membranous); 3 nerved; entire; awnless. Palea
present; relatively long. Lodicules 2; membranous;
glabrous. Stamens 1-3. Ovary glabrous. Fruit small; hilum
short; embryo small.

Fig. 101. Hainardia cylindrica

174

Cytology, classification, distribution. Chromosome base
number, x = 13. Pooideae; Poodae; Poeae. 1 species.
Mediterranean to Iraq. Mesophytic; in open habitats
(meadows, etc., often coastal). Cape Province. 1 naturalized
species.

References. 1. Greuter. 1967. Boissiera 13: 178. 2.
Clayton & Renvoize. 1986. Gen. Gram.

Species treatment by G.E. Gibbs Russell.

Hainardia cylindrica (Willd.) Greuter

Fig. 101. PI. 89.

( =Monerma cylindrica auctt.,
non (Willd.) Coss & Dur.) 1,2.

Erect annual; 50-350 mm tall.

Leaf blades to 7 mm long; to 2.5
mm wide. Spikelets 5-8 mm long.

Spikes cylindrical, solitary;
spikelets sunk in the rhachis, with
one glume and one floret.

Flowering November to December. Weedy in moist
places. Rare. Naturalized from southern Europe. Biome:
Fynbos. Mediterranean. Weed. Similar to Parapholis
incurva, which has two glumes, Lepturus repens, which is
a stoloniferous perennial, and Lolium rigidum , which has
several florets per spikelet.

Description: Chippindall 1955 (73). Illustration: Chip-
pindall 1955 (fig. 45). Voucher: Adamson 3309. PRECIS
code 9904423-00100.

Harpochloa Kunth

Perennial; densely caespitose. Culms 300-900 mm high;
herbaceous; unbranched above. Leaf blades stiffly linear;
flat, or rolled. Ligule a fringed membrane.

Inflorescence a single spike (very rarely two); usually
non-digitate; espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; alternately biseriate (along the midrib
of the rachis); 6-7 mm long ; compressed laterally (darkly
pigmented ); disarticulating above the glumes; not disarticu-
lating between the florets. Glumes two (dark grey-green);
very unequal (G2 much larger); about equalling the
spikelets (i.e. the upper glumes); awnless; very dissimilar
(G 1 smaller, 1 -keeled, thinner. G2 2-keeled, firm). Incom-
plete florets distal to the female-fertile florets (the second
and third male, the fourth if present sterile, these all
enclosed in the lemma of the lower male floret and not
exceeding the LI), merely underdeveloped, awnless;
proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes (firmly membranous); without a germination flap;
3 nerved; entire; awnless. Palea present (hairy near tip); rel-
atively long. Lodicules fleshy (winged); glabrous. Stamens
3 (in hermaphrodite and male florets). Ovary glabrous. Fruit
small (3 mm); ellipsoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath extensions absent. PCR cell
chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 1
species. Southern Africa. Mesophytic; in open habitats
(grassland); glycophytic. Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. 1 indige-
nous species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

i

Harpochloa falx (L. f.) Kuntze

Fig. 102. PI. 90.

Ruspergras, caterpillar grass.

Perennial; distinctly rhizomat-
ous and tufted (densely); 400-900
mm tall. Leaf blades 100-250
mm long; often inrolled, 2— 4(— 6)
mm wide. Spikelets 6-9 mm long.

Solitary one-sided, ‘toothbrush’
spikes, up to 80 mm long and 10
mm wide, sickle-shaped at maturity; spikelets in two rows,
3^1-flowered.

Flowering September to April. Stony well-drained to
compacted soils on moist slopes. Common (often in large
dense stands). Biome: Fynbos, Savanna, and Grassland.
Pasture (highly palatable).

175

Description: Chippindall 1955 (191). Illustration: Chip-
pindall 1955 (fig. 166). Voucher: Van Wyk & Theron 4706.
PRECIS code 9902980-00100.

Helictotrichon Schult.

Avenochloa Holub, Avenula (Dumort.) Dumort.,
Danthorhiza Ten ,,Heuffelia Schur.

Perennial, caespitose. Culms 150-1500 mm high; her-
baceous; unbranched above. Leaf blades linear; flat, or
folded, or rolled (convolute). Ligule an unfringed
membrane ( sometimes puberulent).

Inflorescence paniculate ; open ; espatheate. Spikelet-
bearing axes persistent.

Spikelets 8-25 mm long; compressed laterally ; disartic-
ulating above the glumes. Callus pointed, hairy . Glumes
two; very unequal, or more or less equal; decidedly shorter
than the adjacent lemmas; awnless; similar (persistent,
hyaline to scarious or firm & herbaceous). Incomplete
florets distal to the female-fertile florets, merely underde-
veloped, owned; proximal incomplete florets absent.

Female-fertile florets 2-7 . Lemmas decidedly firmer
than the glumes; non-carinate (dorsally rounded)', 5-7
nerved; incised (usually bidentate); awned. Awns 1;
median; dorsal; geniculate; much longer than the body of
the lemma (always?). Palea present; relatively long. Lodi-
cules 2; membranous; glabrous. Stamens 3. Ovary hairy.
Fruit medium sized; hilum long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 7. Pooideae; Poodae; Aveneae. About 90
species. Europe, Africa, Southeast Asia, North & South
America. Mesophytic to xerophytic, or helophytic (rarely);
mostly in open habitats (dry hillsides, meadows, margins
of woods). Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 13 indigenous species.

References. 1. Schweickerdt. 1937. Bothalia. 3,2. 2.
Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

1(0). Rachilla internodes glabrous H. leoninum

Rachilla intemodes sparsely or densely hairy .... 2
2(1). Rachilla intemodes 2. 5^4. 5 mm long; spikelets

usually loosely flowered 3

Rachilla internodes to 2.3 mm long; spikelets usually
closely flowered (if loosely flowered then leaves

setaceous) 6

3(2). Leaves setaceous; panicle open; spikelets fewer than

20 H. sp. (=Ellis 4663)

Leaves flat; panicle lax or contracted; spikelets more

than 20 4

4(3). Panicles 150-300 mm long, linear; upper glume
shorter than half the spikelet length; spikelets

15-30 mm long H. longum

Panicles 60-120 mm long, oblong or ovate; upper
glume longer than half the spikelet length, spikelets

10-17 mm long 5

5(4). Lemmas coarsely granular and scabrid below the
insertion of the awn; plants about 300 mm tall . .

H. namaquense

Lemmas smooth below the insertion of the awn; plants

about 800 mm tall H. barbatum

6(2). Lemmas scabrid or scaberulous 7

Lemmas glabrous 9

7(6). Glumes almost equal and as long as the spikelet,

broadly lanceolate, almost enclosing the spikelet,
usually scabrous especially on the nerves; leaves

usually woolly-hairy H. galpinii

Glumes unequal to subequal, upper glume 2/3 the
spikelet length, narrowly lanceolate, glabrous;
leaves usually glabrous 8

8(7). Lemmas about 7 mm long; rachilla internodes 1.5 mm

long H. hirtulum

Lemmas about 10 mm long; rachilla internodes 2.0

mm long H. capense

9(6). Lemmas shorter than 7 mm, nerves usually raised .

H. natalense

Lemmas longer than 7 mm, nerves usually

inconspicuous 10

10(9). Spikelets 12-15 mm long; lemma lobes above the
insertion of the awn 6-8 mm long; panicle
usually very dense and contracted, spikelets

yellowish H. dodii

Spikelets 8-12 mm long; lemma lobes above the
insertion of the awn 3-5 mm long; panicle lax,
open or slightly contracted; spikelets green, often

variegated with purple 11

11(10). Leaves setaceous, 200^400 mm long; rachilla hairs
3^1 mm long; spikelets loosely flowered, with
the rachilla usually exposed between the florets

H. longifolium

Leaves flat, 60-150 mm long; rachilla hairs 2 mm
long; spikelets closely flowered, with the rachilla
usually not showing H. turgidulum

Helictotrichon barbatum (Nees) Schweick.

Perennial; densely tufted;

600-800 mm tall. Leaf blades
150-320 mm long; 1.5-3. 5 mm
wide. Spikelets 14-17 mm long.

Leaves flat; panicle 80-100 mm
long, oblong, lax; upper glume
longer than 1/2 the spikelet;
rachilla internodes bearded,

3. 0-3. 5 mm long; lemmas smooth
below the insertion of the awn.

Flowering November. Lower mountain slopes. Very
rare. Biome: Succulent Karoo. Endemic. Known from two
localities only: Kamiesberg and Hantamberge in the
western Cape.

Description: Schweickerdt 1937 (190), Chippindall
1955 (78). Illustration: Chippindall 1955 (fig 48(8)).
Voucher: Acocks 18632. PRECIS code 9901970-00100.

Helictotrichon capense Schweick.

Perennial; tufted; to 1000 mm
tall. Leaf blades to 250 mm long;

1-2 mm wide. Spikelets about 15
mm long. Leaf blades filiform,
expanded; panicle to 200 mm
long; upper glume 2/3 the spikelet
length; rachilla internodes 2 mm
long; lemma scabrous, about 10
mm long.

Flowering November, December, and May. Sandy soils,
occasionally in disturbed places. Infrequent. Biome: Fyn-
bos and Savanna. Endemic. A poorly defined species,
which is usually larger than H. hirtulum in all dimensions.

Description: Schweickerdt 1937 (193), Chippindall
1955 (78). Illustration: Chippindall 1955 (fig. 48(3) & 49).
Voucher: Sim 2803. PRECIS code 9901970-00200.

Helictotrichon dodii (Stapf) Schweick.

Perennial; tufted; 500-1000
(-1250) mm tall. Leaf blades
300-500 mm long; 3-5 mm wide.
Spikelets 12-15 mm long. Leaf
blades flat; panicle 120-300 mm
long, narrow, contracted, usually
dense; glumes lanceolate, upper
glume 2/3 the spikelet length;
rachilla internodes 2 mm long.

176

with hairs 3.0-3. 5 mm long; lemmas glabrous, linear in
outline, lobes delicately awned, 6-8 mm long.

Flowering sporadic, but mainly October to December.
On coastal sandflats, disturbed places and vlei margins. In-
frequent. Biome: Fynbos and Grassland. Endemic. The very
dense, contracted panicle and long lemma lobes (which give
a delicate, slender appearance), make this a very well
defined species. Related to H. turgidulum, which has
shorter lemma lobes, to H . natalense , which has shorter

spikelets, and to H. longifolium, which has setaceous leaf
blades.

Description: Schweickerdt 1937 (197), Stapf 1898-1900
(475), Chippindall 1955 (79). Illustration: Chippindall 1955
(fig. 48(12)). Voucher: Pole Evans 518. PRECIS code
9901970-00300.

Helictotrichon galpinii Schweick.

Perennial; tufted; to 600 mm
tall. Leaf blades 120-160 mm
long; 2. 5-3.0 mm wide. Spikelets
8-10 mm long. Leaves expanded,
hairy; panicle 100-160 mm long,
contracted; glumes almost equal,
broadly lanceolate, as long as the
spikelet; rachilla intemodes 1.25
mm long, hairy; lemmas covered
with papillae and prickles.

Flowering January to March. On humic soils in wet
places. Infrequent. Biome: Grassland. Endemic. Readily
distinguished by the broad glumes that are as long as the
spikelets and the scabrid lemmas.

Description: Schweickerdt 1937 (192), Chippindall
1955 (78). Illustration: Chippindall 1955 (fig. 48(10)).
Voucher: P.C.V. du Toit 2311. PRECIS code

9901970-00400.

Helictotrichon hirtulum (Steud.) Schweick.

Fig. 103.

Slender, weak perennial;
loosely tufted; to 1000 mm tall.

Leaf blades to 250 mm long;
filiform, to 2 mm wide. Spikelets
8-1 1 mm long. Panicle lax,
almost spike-like, to 200 mm
long; glumes unequal, upper to
2/3 the spikelet length; rachilla
internodes 1 .5-2.0 mm long, with
hairs to 2 mm long; lemma scabrid below the insertion of
the awn.

Flowering November to March. On mountain slopes,
often in shady, wet places and also in disturbed areas. Infre-
quent to locally common. Biome: Fynbos, Savanna,
Grassland and Nama-Karoo. Endemic. Very similar to H.
capense , which normally is a larger, more robust plant.

Description: Schweickerdt 1937 (193), Chippindall
1955 (78). Illustration: Chippindall 1955 (fig. 48(2)).
Voucher: Sim 2803. PRECIS code 990 1970-00500.

Helictotrichon leoninum (Steud.) Schweick.

Perennial; loosely to fairly
densely tufted; 1 50 — 450( — 650)
mm tall. Leaf blades 40-100
(-180) mm long; filiform, to 4
mm wide. Spikelets 12-14 mm
long. Leaf blades flat; panicle
60-100 mm long, linear, contrac-
ted; upper glume about 1/2 the
spikelet length; rachilla inter-
nodes glabrous, 1. 5-2.0 mm long; lemmas very densely
papillose.

Flowering August to November. On mountain slopes,
along mountain roads and in humic seepage areas. Infre-
quent. Biome: Fynbos. Endemic. This is the only southern
African Helictotrichon species with glabrous rachilla inter-
nodes, but one should be careful that the hairs on the
densely tufted callus are not mistaken for rachilla hairs.

Description: Schweickerdt 1937 (191), Chippindall
1955 (78). Illustration: Chippindall 1955 (fig'. 48(6)).
Voucher: P.C.V. du Toit 1419. PRECIS code 9901970-
00600.

177

Helictotrichon longifolium (Nees) Schweick.

Perennial; tufted; 300-900
mm tall. Leaf blades 200^100
mm long; setaceous, to 1.5 mm
wide. Spikelets 8— 1 0(— 1 2) mm
long. Panicle to 200 mm long,
usually open; upper glume about
3/4 the spikelet length; rachilla
internodes about 2 mm long, with
hairs 3-4 mm long; lemma
smooth or finely papillate, lobes above the insertion of the
awn shorter than 7 mm.

Flowering December to April. On moist and rocky
mountain slopes. Locally common. Biome: Grassland and
Nama-Karoo. Endemic. Very closely related to H. dodii ,
which has flat leaf blades and longer spikelets, to H.
turgidulum , which has shorter rachilla hairs and flat leaf
blades, and to H. natalense , which has shorter spikelets.

Description: Schweickerdt 1937 (195), Stapf 1898-1900
(All), Chippindall 1955 (79). Voucher: Smook 1375.
PRECIS code 9901970-00700.

Helictotrichon longum (Stapf) Schweick.

Perennial; long rhizomatous
and tufted (with new shoots
spreading for a short distance
underground before emergence);

600—1 100(— 1600) mm tall. Leaf
blades 1 50— 300(— 400) mm long;

2.5-10mm wide. Spikelets 15-30
mm long. Leaves flat; panicle
150-300 mm long, linear, usually
contracted; upper glume shorter than 1/2 the spikelet;
rachilla internodes 2. 5-3.0 mm long, bearded; lemmas
minutely granular below the insertion of the awn.

Flowering September to November. Sandy flats in
coastal fynbos, occasionally in moist areas. Infrequent.
Biome: Fynbos. Endemic. A very distinct tall plant with
flat, wide leaf blades and a long panicle with notably long
spikelets.

Description: Schweickerdt 1937 (189), Stapf 1898-1900
(473), Chippindall 1955 (77). Illustration: Chippindall 1955
(fig. 48(5)). Voucher: Acocks 19730. PRECIS code
9901970-00800.

Helictotrichon namaquense Schweick.

Perennial; densely tufted;

250^150 mm tall. Leaf blades
50-150 mm long; 2 -A mm wide.

Spikelets 10-17 mm long. Leaves
flat; panicle 60-120 mm long,
ovate, contracted, lower branches
sometimes spreading; upper
glume longer than 1/2 the spike-
let; rachilla internodes 2. 5-3.0
mm long, bearded; lemmas coarsely granular, scabrid
below the insertion of the awn.

Sandy flats in Renosterbosveld. Rare. Locally common.
Biome: Nama-Karoo. Endemic. Recorded only from the
Sutherland district.

Description: Schweickerdt 1937 (189), Stapf 1898-1900
(473), Chippindall 1955 (78). Illustration: Chippindall 1955
(fig. 48(4)). Voucher: Acocks 17178. PRECIS code
9901970-00900.

Helictotrichon natalense (StapD Schweick.

Perennial; tufted; 400-800
(-1000) mm tall. Leaf blades
100-250 mm long; 3-5 mm wide.

Spikelets 7-9 mm long. Leaf
blades flat; panicle to 250 mm
long, usually open, with branches
spreading; upper glume 2/3 the
spikelet length; rachilla inter-
nodes about 1.5 mm long, hairy;

lemmas glabrous, usually smooth, nerves usually raised,
column of awn a loose spiral.

Flowering November to January. On rocky hillsides and
in wet places such as streamsides. Infrequent to locally
common. Biome: Savanna and Grassland. Endemic.
Related to H . longifolium , which has setaceous leaves and
longer spikelets, and to H. turgidulum and H. dodii, which
have longer spikelets.

Description: Schweickerdt 1937 (194), Stapf 1898-1900
(477), Chippindall 1955 (79). Illustration: Chippindall 1955
(fig. 48(1)). Voucher: De Wet 1722. PRECIS code
9901970-01000.

Helictotrichon quinquesetum (Steud.) Schweick.

Perennial; tufted; 500-750 mm tall. Leaf blades to 250
mm long; to 4 mm wide. Spikelets 12-18 mm long. Leaves
expanded or folded; panicle 120-180 mm long, contracted,
almost spike-like; rachilla internodes 3. 5-4. 5 mm long,
with hairs to 4 mm long; lemma with prominently raised
nerves, finely granular.

Slopes of Table Mountain. Extremely rare. Biome: Fyn-
bos. Endemic. Represented in most herbaria only by
duplicates of the type specimen. Ecklon 929. None at PRE.

Description: Schweickerdt 1937 (188), Stapf 1898-1900
(474), Chippindall 1955 (77). Illustration: Chippindall 1955
(fig. 48(9)). PRECIS code 9901970-01 100.

Helictotrichon turgidulum (StapD Schweick.

PI. 91.

Perennial; tufted; 300-1000
mm tall. Leaf blades
60— 1 50(— 250 ) mm long; 1.5-6. 0
mm wide. Spikelets 10-12 mm
long. Leaf blades flat; panicle
70-300 mm long, open or
contracted, often interrupted;
glumes broadly lanceolate, upper
glume 2/3 the spikelet length;
rachilla internodes 2 mm long, with hairs to 2 mm long;
lemma glabrous, smooth or papillate, very often variegated
with purple, lobes, above the insertion of the awn, 3-5 mm
long.

Flowering October to April. Usually in wet places on
mountain slopes and in vleis, occasionally at roadsides.
Common. Biome: Fynbos, Savanna, and Grassland.
Endemic. Vegetatively a very variable species and by far
the most widespread of all the South African Helictotrichon
species. Related to H. dodii, which has longer lemma lobes
and broader glumes, to H. longifolium, which has setaceous
leaves, and to H. natalense, which has smaller spikelets.

Description: Schweickerdt 1937 (196), Stapf 1898-1900
(474), Chippindall 1955 (79). Illustration: Chippindall 1955
(fig. 48(7) & 50). Voucher: Smook 2560. PRECIS code
9901970-01200.

Helictotrichon sp. (=Ellis 4663)

Perennial; densely tufted;

300-600 mm tall. Leaf blades
100-200 mm long; setaceous, to
1.5 mm wide. Spikelets 13-18
mm long. Panicle open, branches
spreading, bare for most of their
length; spikelets fewer than 20;
upper glume about 2/3 the spike-
let length; rachilla internodes
2. 8-3. 3 mm long, densely hairy; lemmas glabrous.

Flowering October. In shallow humic soils between
limestone outcrops. Rare. Biome: Fynbos. This species was
collected by Ellis on two occasions at De Hoop. It has
setaceous leaves, an open, lax panicle with very few spike-
lets and long rachilla internodes. This combination of
characters are not matched in any other southern African
Helictotrichon species.

Voucher: Ellis 4663. PRECIS code 9901970-99999.

178

Hemarthria R.Br.

Lodicularia P. Beauv.

Perennial; long-stoloniferous, or caespitose, or
decumbent. Culms 300-1500 mm high; herbaceous;
branched above. Leaf blades linear, or linear-lanceolate
(usually); flat. Ligule a fringed membrane. Plants bisexual,
with bisexual spikelets. The spikelets overtly heteromorphic
( the sunken, ‘sessile’ spikelets with dissimilar glumes, the
non-sunken ‘pedicellate’ spikelets with similar glumes).

Inflorescence of spike-like main branches, or paniculate
(of ‘spikes' arising one or more from the sheaths of each
of the upper leaves)’, spatheate (usually)’, a complex of
‘partial inflorescences’ and intervening foliar organs.

Spikelet-bearing axes spike-like (often curved ); solitary, or
clustered (fascicled); with substantial rachides; tardily dis-
articulating (the rachis initially tough); ultimately disarticu-
lating at the joints. ‘Articles’ with a basal callus-knob
(rarely), or without a basal callus-knob (usually).

Spikelets in pairs (each pair comprising a sessile spikelet
and the ‘pedicellate’ member of the ‘pair’ below); consis-
tently in ‘long-and-short’ combinations (the fused pedicels
discernible). Pedicels of the ‘pedicellate’ spikelets
discernible, but fused with the rachis. The sessile spikelets
hermaphrodite. The ‘pedicellate’ spikelets hermaphrodite.
Female-fertile spikelets 3-7 mm long; compressed dorsi-
ventrally; falling with the glumes. Glumes present; two;
more or less equal; awnless (but G2 sometimes long-
acuminate); very dissimilar (in the embedded spikelet, the
outer tough, the inner membranous), or similar (both tough
in the ‘pedicellate’ spikelets). Proximal incomplete florets
1; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present; conspicu-

ous but relatively short. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 9, or 10. Panicoideae; Andropogonodae;
Andropogoneae; Rottboelliinae. 12 species. Tropical
Africa, Madagascar, eastern Asia, Indomalayan region,
Australia. Hydrophytic to helophytic; in open habitats (in
water or in wet places); glycophytic. Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Hemarthria altissima (Poir.) Stapf & C.E. Hubb.

Batavian quick grass, red
swamp grass, perdegras, rooi-
kweek.

Perennial; rhizomatous and
stoloniferous; 300- 1500 mm tall.

Leaf blades 50-150 mm long; to
6 mm wide. Spikelets (sessile and
pedicellate) 5-7 mm long. Plants
rust-red; racemes very narrowly cylindrical, culm-like,
spikelets sunken.

Flowering October to June. Wet places. Sometimes lo-
cally dominant (in vleis and river margins). Biome: Fynbos,
Savanna, Grassland, Nama-Karoo, and Succulent Karoo.
Southern tropical Africa and Madagascar, Mediterranean
region, southeast Asia, introduced to America. Pasture and
weed (ruderal).

Description: Chippindall 1955 (519), Clayton et al.
1970-1982 (851). Illustration: Chippindall 1955 (fig. 414).
Voucher: De Winter 4221 . PRECIS code 9900210-00100.

Heteropogon Pers.

Spirotheros Raf.

Annual, or perennial; caespitose. Culms 200-1000 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear; flat. Ligule a fringed membrane. Plants
bisexual, with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant (heterogamous in upper
parts of inflorescence only)’, overtly heteromorphic; in both
homogamous and heterogamous combinations ( lower pairs
homogamous and homomorphic , male or sterile).

Inflorescence a single raceme, or paniculate (of single
‘racemes' , sometimes in false panicles)’, spatheate, or
espatheate; not comprising ‘partial inflorescences’ and
foliar organs. Spikelet-bearing axes ‘racemes' (of several
to many joints)’, solitary, disarticulating at the joints
(between the heterogamous upper spikelet pairs).

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile (but the pedicel reduced
to a short stump, the spikelet being supported on a long,
slender callus). Pedicels free of the rachis. The sessile
spikelets hermaphrodite (in upper regions of spike-like
panicles only), or female-only. The pedicellate spikelets
male-only, or sterile; awnless, dorsally flattened, rather
asymmetric. G1 herbaceous, many nerved, winged above.
Female-fertile spikelets not noticeably compressed (rarely),
or compressed dorsiventrally; falling with the glumes.
Glumes two; more or less equal; awnless; very dissimilar
(the upper with deep longitudinal grooves). Proximal in-
complete florets 7; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline, but stipitate-cartilaginous beneath the
awn); entire; awned. Awns 1; median; apical; geniculate;
much longer than the body of the lemma. Palea present, or

Fig. 104. PI. 92.

179

absent; when present very reduced. Lodicules when present
2; fleshy; glabrous. Stamens 0-3. Ovary glabrous. Hilum
short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10 and 11. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 7 species. Tropical.

Mesophytic to xerophytic; in open habitats (dry places,
often on poor soils); glycophytic. Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 2 indigenous species.

References. 1. Chippindall 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

1(0). Perennial; leaf blades usually folded, 3-8 mm wide;
glands absent; pedicelled spikelets 8-13 mm long

H. contortus

Annual; leaf blades flat, to 12 mm wide; dark
crateriform glands present on spathes, peduncles
and lower glumes of pedicelled spikelets;

pedicelled spikelets 16-20 mm long

H. melanocarpus

Heteropogon contortus (L.) Roem. & Schult.

Tanglehead,pylgras,assegaai-
gras.

Perennial; rhizomatous; 200-
1000 mm tall. Leaf blades 30-
300 mm long; 3-8 mm wide.

Spikelets (sessile) 5.5-7 mm long
(pedicellate 8-13 mm long and
glandless). Leaves usually folded,
the tips rounded and often hooded; inflorescence a single
spike with velvety awns from the upper half.

Flowering October to June. Hillsides and rocky places
on well-drained soils. Common. Biome: Fynbos, Savanna,
Grassland, and Nama-Karoo. Tropical and warm regions.
The large, awned, single-raceme inflorescence resembles
Urelytrum agropyroides and Trachypogon spicatus , but in
both these species the inflorescence has awns throughout
its length. H. contortus often occurs with Themeda triandra
and Schizachyrium sanguineum and may resemble them
vegetatively. However, Themeda has tapering leaf tips and
the ligule is usually notched and S. sanguineum has a
strongly curved ligule and the plant is red or purple tinged.

Description: Chippindall 1955 (492), Clayton et al.
1970-1982 (827). Illustration: Chippindall 1955 (fig. 400),
Clayton et al. 1970-1982 (fig. 191), Hitchcock & Chase
1950 (fig. 1 182). Voucher: Giess, Volk & Bleissner 6429.
PRECIS code 9900800-00100.

Heteropogon melanocarpus (Ell.) Benth.

Sweet tanglehead, eenjarige
assegaaigras.

Robust annual; tufted; 500-
2000 mm tall. Leaf blades to 500
mm long; to 12 mm wide. Spike-
lets (sessile) 10-11 mm long
(pedicellate with lower glume
16-20 mm long, with a row
of depressed glands in the middle). With stilt roots; leaf
blades flat.

Flowering January to May. Roadsides and rocky places,
often on turf soil. Conservation status not known. Locally
common. Biome: Savanna. Tropical Africa to India,
tropical America.

Description: Chippindall 1955 (494), Clayton et al.
1970-1982 (827). Illustration: Hitchcock & Chase 1950
(fig. 1 183). Voucher: De Winter & Marais 4601. PRECIS
code 9900800-00200.

Fig. 105. PI. 93.

180

Holcus L.

Arthrochloa R. Br., Ginannia Bub., Homalachna
Kuntze, Nothoholcus Nash, Notholcus Hitchc., Sorghum
Adans.

Annual (rarely), or perennial; long-rhizomatous to long-
stoloniferous, or caespitose. Culms 80-1500 mm high; her-
baceous; unbranched above. Leaf blades linear to lirear-
lanceolate; flat; without readily visible transverse veins.
Ligule an unfringed membrane to a fringed membrane .

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 3-8 mm long; compressed laterally; falling
with the glumes. Glumes two; more or less equal; about
equalling the spikelets; awned (rarely), or awnless; similar
(membranous). Upper glume 3 nerved. Incomplete florets
distal to the female-fertile florets (spikelets 2-flowered, the
lower hermaphrodite, the upper usually male-only), clearly
specialised and modified in form; awned (with a short
dorsal awn); proximal incomplete florets absent.

Fig. 106. Holcus lanatus

Female-fertile florets l(-2). Lemmas decidedly firmer
than the glumes ( leathery ); 3-5 nerved; entire, or incised;
awnless, or awned. Awns when present 7; dorsal; genicu-
late. Palea present; relatively long. Lodicules 2;
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short, or long-linear (rarely); embryo small.

Cytology, classification, distribution. Chromosome base
number, x =4 and 7. Pooideae; Poodae; Aveneae. 6 species.
8 Canary Is., North Africa, Europe to Asia Minor &
Caucasus; 1 South Africa. Mesophytic; in shade and in open
habitats (grassland, open woodland, disturbed ground);
glycophytic. Transvaal, Natal, and Cape Province. Indige-
nous species (1), naturalized species (1).

References. 1. Tutin. 1980. FL. Europ. 5: 230.
Species treatment by T.M. Sokutu.

1(0). Perennial; leaves villous; upper glume awn to 1 mm

long; upper floret usually male H. lanatus

Annual; leaves puberulous; upper glume with an awn
2-6 mm long; upper floret sterile .... H. setiger

Holcus lanatus L.

Velvet grass, Yorkshire fog,
soft grass.

Perennial; loosely tufted; 300-
1000 mm tall. Leaf blades 1.5-
8.0 mm wide. Spikelets 3-4 mm
long. Leaves conspicuously
woolly, greyish to blue-green;
lower glume awn never ex-
ceeding 0.5 mm; upper glume awn to 1 mm long; upper
floret male.

Flowering November to January. Vleis, damp sheltered
places, on sandy to nutrient rich soils. Common (in its
habitats). Naturalized from Europe. Biome: Fynbos, Savan-
na, and Forest. Europe and the Mediterranean. Occasionally
cultivated pasture and weed. A distinct species which
cannot be confused because of its relatively short-awned
glumes, although variable in the glume and lemma awn
length. In the past the name 77. mollis was missapplied to
some specimens of this taxon.

Description: Hubbard 1954 Grasses (237), Tutin 1980
(5: 230), Stapf 1898-1900 (465), Chippindall 1955 (87).
Illustration: Chippindall 1955 (fig. 58). Voucher: Smook
4878, Dyer 6277. PRECIS code 9901920-00100.

Fig. 106. PI. 94.

Holcus setiger Nees

Annual; culms solitary or
loosely tufted; 150-300 mm tall.

Leaf blades 25-140 mm long;

1.0-4. 5 mm wide. Spikelets 3-4
mm long. Leaves puberulous,
rarely hairy, pale to dark green;
lower glume awn to 1 mm long;
upper glume awn 2-6 mm long;
upper floret sterile.

Flowering November to January. On damp and/or
sheltered places, sandy to sandy loam soils. Locally com-
mon. Biome: Fynbos and Succulent Karoo. Endemic.
Chippindall (1955) comments that the forms from George
and Namaqualand are exceptionally slender and weak.
However, this character is not sufficient to distinguish these
forms, as it falls within the variability of the species. In the
past the name 77. mollis has been missapplied to some
specimens of this taxon.

Description: Adams. & Salter 1950 (66), Stapf
1898-1900 (464), Chippindall 1955 (87). Voucher: Acocks
22966, Taylor 3489. PRECIS code 9901920-00300.

Hordeum L.

Critesion Raf., Critho Meyer, Zeocrithon P. Beauv.,
Zeccriton Wolf.

Annual, or perennial; caespitose (or solitary culms).
Cu ms 50-1300 mm high; herbaceous; unbranched above.
Leaf blades linear; usually flat, or folded (convolute).
Ligule an unfringed membrane . The spikelets of sexually
distinct forms on the same plant (the lateral spikelets sterile
in Critesion , male in Hordeum s. str. ).

181

Inflorescence a false spike, with clusters of spikelets on
reduced axes ; contracted; espatheate. Spikelet-bearing axes
disarticulating (e.g., Critesion), or persistent (Hordeum s.
str.); when disarticulating, disarticulating at the joints.

Female-fertile spikelets in triplets (the triplets shed
together); distichous, or not two-ranked (2-6 rows); consis-
tently in ‘long-and-short’ combinations, or not in distinct
‘long-and-short’ combinations (rarely, the laterals also
sessile). Female-fertile spikelets compressed laterally to not
noticeably compressed; falling with the glumes (the triplets
falling together), or not disarticulating (in cultivated forms).
Glumes two; more or less equal; awned; similar (persistent,
narrow, awn- or bristle-like above). Spikelets with female-
fertile florets only; proximal incomplete florets absent.
Male or sterile spikelets, when present, awnless.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes (coriaceous); 5 nerved; entire, or incised
(sometimes trifid); nearly always awned. Awns 1; median;
apical; non-geniculate; much shorter than the body of the
lemma, to much longer than the body of the lemma. Palea
present; relatively long. Lodicules membranous; ciliate.
Stamens 3. Ovary hairy. Fruit small, or medium sized, or
large; hilum long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Triticodae; Triticeae. About 40
species. North temperate & South America. Mesophytic, or
xerophytic; in open habitats (open weedy places, mostly on
dry soils); maritime-arenicolous, halophytic, and
glycophytic. Transvaal, Orange Free State, Natal, Lesotho,
and Cape Province. Indigenous species (1), naturalized
species (3).

Intergeneric hybrids with Elytrigia (X Elytrohordeum
Hylander), Agropyron ( X Agrohordeum A. Camus), Secale

(X Hordale Ciferri & Giacom.), Sitanion (X Sitordeum
Bowden), Triticum (X Tritordeum Aschers. & Graebn.). See
alsoX Elyhordeum Zizan & Petrowa.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Bothmer et al. 1980. Bot. Notiser 133: 539. 3. Humphries.
1980. FI. Europ. 4. Dewey. 1984. Genomic classification
in Gustafson, Gene manipulation: 209.

Species treatment by M. Koekemoer.

1(0). Spikes narrower than 6 mm; spikelets with awns

shorter than 10 mm H. stenostachys

Spikes wider than 6 mm; spikelets with awns longer

than 10 mm 2

2( 1 ). Plants perennial; leaf sheaths lacking auricles, sheaths
fibrous with age; upper leaf blades usually rigid .

H. capense

Plants annual; leaf sheaths auricled, not fibrous with

age, upper leaf blades soft or firm 3

3(2). Glumes of lateral spikelets scabrid or smooth

H. marinum subsp. gussoneanum

Glumes of lateral spikelets long-ciliate 4

4(3). Anthers of central spikelet 0.2-0. 5 mm long;

prolongation of rachilla of lateral spikelets stout,
orange-brown ... H. murinum subsp. glaucuin
Anthers of central spikelet 0.7-1. 4 mm long;

prolongation of rachilla of lateral spikelets slender,

green 5

5(4). Central spikelet sessile or with a pedicel not more
than 0.6 mm long; lateral spikelets as long as or

shorter than the central spikelet

H. murinum subsp. murinum

Central spikelet with pedicel (0.7— )0.9— 1 .8 mm long;
lateral spikelets longer than the central spikelet .
H. murinum subsp. leporinum

Hordeum capense Thunb.

(=H. nodosum auctt., non
L.) 1.

Perennial; tufted; 200-600
mm tall. Leaf blades 60-170
(-240) mm long; 3-6 mm wide.

Leaf sheaths fibrous with age;
blades rigid, lacking auricles;
spike 8-17 mm wide (includ-
ing awns), rachis readily disarticulating at maturity; spike-
let awns to 20 mm long.

Flowering October to April. Usually in moist areas such
as streamsides, riverbanks and around dams, occasionally
in disturbed areas. Infrequent to locally common. Biome;
Fynbos, Savanna, Grassland, Nama-Karoo, and Succulent
Karoo. Endemic. Potential pasture. Similar to H. marinum
and H. murinum, which are annual and have leaf sheaths
auricled.

Description: Chippindall 1955 (72). Voucher: Loxton
239. PRECIS code 9904510-00100.

Hordeum marinum Huds. subsp. gussoneanum (Pari.)
Thell.

(=Critesion marinum (Huds.)
Loeve) 4.

Mediterranean barley.

Annual; culms solitary or
loosely tufted (often geniculate);
150^400 mm tall. Leaf blades
20-80 mm long; 2-4 mm wide.

Fig. 107. Hordeum murinum

182

Leaf sheaths auricled, not fibrous with age; blades soft;
spike 15-25 mm wide, fragile, rachis disarticulating readily
at maturity; spikelet awns to 30 mm long.

Flowering September to November. Usually on
roadsides or in moist waste and disturbed places. Locally
common. Naturalized from Europe. Biome: Fynbos and
Succulent Karoo. Europe, Mediterranean basin and in the
U.S.A. Closely related to H. murinum, which has the
glumes of the lateral spikelets long ciliate.

Description: Bor 1985 (1833), Hitchcock & Chase 1950
(266). Voucher: Vlok 1575. PRECIS code 9904510-00250.

Hordeum murinum L. subsp. glaucum (Steud.) Tzvel.

Differs from subsp. murinum
in that the anthers of the central
spikelet are 0.2-0. 5 mm long and
the prolongation of the rachilla of
the lateral spikelets stout and
orange-brown.

Flowering August to October.

On sandy soils, usually in
disturbed areas. Locally common.

Naturalized from Europe. Biome: Fynbos, Savanna, and
Succulent Karoo. Mediterranean Basin of Europe. Weed.

Description: Humphries 1980 (5:204). Voucher: Oliver
151. PRECIS code 9904510-00325.

Hordeum murinum L. subsp. ieporinum (Link)

Archangeli

Differs from subsp. murinum
in that the central spikelet has a
pedicel (0.7-)0.9-l .8 mm long
and lateral spikelets longer than
the central one.

Flowering September to Nov-
ember. Disturbed areas. Infre-
quent to locally common. Nat-
uralized from Europe. Biome:

Fynbos and Grassland. Europe. Weed.

Description: Humphries 1980 (5:204). Voucher: Van
Breda 2011. PRECIS code 9904510-00330.

Hordeum murinum L. subsp. murinum

Fig. 107. PI. 95.

( =Critesion murinum (L.)

Loeve) 4.

False barley, muiswildegars.

Annual; culms solitary or
loosely tufted; 50-500 mm tall.

Leaf blades 20-150(-250) mm
long; 2-8 mm wide. Leaf sheaths
auricled, not fibrous with age; blades soft; spike 15-25 mm
wide, rachis disarticulating readily at maturity, the central
spikelet sessile or with a pedicel shorter than 0.6 mm;
lateral spikelets shorter or as long as central spikelet; awns
to 30 mm long; anthers of central spikelet 0.7- 1 .4 mm long;
prolongation of rachilla of lateral spikelets slender and
green.

Flowering October to December. On sandy soil, usually
in disturbed areas. Locally common. Naturalized from
Europe. Biome: Fynbos, Savanna, and Succulent Karoo.
Introduced and naturalized in many countries. Weed (in
sandy waste places). The long-ciliate glumes of the lateral
spikelets distinguish this taxon from H. marinum subsp.
gussoneanum.

Description: Humphries 1980 (5:204), Hitchcock &
Chase 1950 (268), Chippindall 1955 (72). Illustration:
Chippindall 1955 (fig. 44). Voucher: Orchard 51 1 . PRECIS
code 9904510-00335.

Hordeum stenostachvs Godr.

( =Critesion stenostachys
(Godr.) Loeve) 4; (=//.
compressum Griseb.) 2.

Perennial; tufted; (280-)350-
1 200(-1500) mm tall. Leaf blades
25— 90(— 1 02) mm long; 1. 5-5.0
(-7.0) mm wide. Leaf sheaths
lacking auricles; spike 3-5 mm
wide (including awns), often
partly enclosed in uppermost leaf sheath; rachis readily dis-
articulating at maturity; spikelet awns 2. 5-8.0 mm long.

Flowering October to March. In moist or seasonally
moist areas, vleis and also in disturbed places. Locally com-
mon. Probably naturalized from south America. Biome:
Grassland and Nama-Karoo. Tropical South America.
Pasture (eaten by stock). Easily distinguished from the
other Hordeum species in southern Africa by the narrow,
slender spikes and short awns.

Description: Von Bothmer, Jacobsen & Nicora 1980
Bot. Notiser 133 (546). Voucher: Comins 845. PRECIS
code 9904510-00385.

Hyparrhenia Fourn.

Annual (rarely), or perennial (usually large); caespitose.
Culms 300-3000(^1000) mm high; herbaceous. Leaf blades
linear; usually flat, or folded (sometimes). Ligule an
unfringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant (the lower spikelet pairs homogamous, the
upper pairs heterogamous); overtly heteromorphic
(imperfect spikelets sometimes with awned glumes, the L2
awnless).

Inflorescence paniculate (leafy)', with capillary
branchlets (i.e., the articles of the racemes, and the
peduncles); spatheate ; a complex of ‘partial inflorescences’
and intervening foliar organs. Spikelet-bearing axes
‘racemes’; paired (with a common peduncle, the upper
raceme base usually much shorter than 9 mm — by contrast
with Exotheca); with very slender rachides; disarticulating
at the joints.

Spikelets in pairs (with terminal triplets); consistently in
‘long-and-short’ combinations; these pedicellate/sessile.
Pedicels free of the rachis. The sessile spikelets hermaphro-
dite (in the upper pairs only). The pedicellate spikelets
male-only, or sterile;, usually longer than the sessile, G1
often mucronate or aristate. L2 awnless, sometimes
suppressed. Female-fertile spikelets 3.5-10 mm long; not
noticeably compressed to compressed dorsiventrally;
falling with the glumes. Glumes two; more or less equal;
awnless; very dissimilar (lower rounded or dorsally
flattened; upper narrower, shallowly naviculate). Lower
glume not two-keeled (striate or grooved). Proximal incom-
plete florets 1; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline, but hardening and stipitate to the awn);
incised; awned. Awns 1; median; from the sinus (flanked
by tiny teeth); geniculate; much longer than the body of the
lemma. Palea present, or absent; when present relatively
long, or conspicuous but relatively short, or very reduced.
Lodicules 2; fleshy, or membranous; glabrous. Stamens 3.
Ovary glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, jc = 1 0 and 15. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. About 55 species.
Mediterranean, Africa, Arabia, America. Mesophytic, or

183

xerophytic; in open habitats (savanna); glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. 20 indige-
nous species.

References. 1. Clayton. 1969. Kew Bull. Add. Ser. II.
2. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

Fig. 108. Hyparrhenia hirta

1(0). Upper raceme base with a scarious appendage 3-4

mm long at apex, just below spikelets 2

Upper raceme base lacking an appendage (but
sometimes with a scarious rim or short tooth to 0.5

mm long) 3

2( 1 ). Lower glume of sessile spikelet glabrous or with a few

hairs at tip H. newtonii var. newtonii

Lower glume of sessile spikelet with long hairs . . .

H. newtonii var. macra

3(1). Raceme bases terete, markedly unequal, the upper at
least 3 times longer than the lower, usually not
deflexed (but often deflexed in H. quarrel ) .... 4
Raceme bases flattened, subequal, often deflexed at

maturity, usually less than 2 mm long 14

4(3). Spikelets with reddish brown or yellowish hairs . . 5

Spikelets with white hairs, or glabrous 8

5(4). Sessile spikelets 5-7 mm long, callus 0. 8-2.0 mm

long 6

Sessile spikelets 3-5 mm long, callus 0.2-0. 8 mm

long 7

6(5). Basal leaf sheaths with spreading white hairs; upper

raceme base 2-3 mm long H. nyassae

Basal leaf sheaths glabrous or rarely with a few hairs;

upper raceme base 3. 5-7.0 mm long

H. poecilotricha

7(5). Panicle lax or contracted; spatheoles linear, 40-50
mm long; awns 9-14 per raceme pair . . . H. rufa
Panicle copiously branched; spatheoles narrowly
lanceolate, 20-40 mm long; awns 6-10 per raceme

pair H. dichroa

8(4). Upper raceme with 0 or 1 pairs of homogamous

spikelets at base 9

Upper raceme with 2 pairs of homogamous spikelets

at base 12

9(8). Spikelets glabrous or hispidulous 10

Spikelets pubescent to villous 11

10(9). Culms slender; callus cuneate, 0.8-1. 5 mm long;

upper raceme base (2— )2.5— 3.5 mm long; awns

4-5 per raceme pair H. gazensis

Culms robust; callus linear, slender, 1-2 mm long;
upper raceme base 1.5-2. 5 mm long; awns 2-6

per raceme pair H. finitima

11(9). Racemes never deflexed; awns 8-14 per raceme

pair H. hirta

Racemes, or some of them, deflexed; awns 6-10 per

raceme pair H. quarrei

12(8). Awns 4-7 per raceme pair, 25-40 mm long, with
hairs 0. 1-0.6 mm long; racemes 1 5-25 mm long;
callus 1.0-1. 8 mm long; pedicellate spikelets
awnless or with an awn-point to 2 mm long . .

H. anamesa

Awns 2-4 per raceme pair, 30-55 mm long, with
hairs 0.7-1. 2 mm long; racemes 10-12 mm long;
callus 1. 8-3.0 mm long; pedicellate spikelets

with an awn 1-5 mm long 13

13(12). Spikelets glabrous; awns 2( — 4) per raceme pair .

H. filipendula var. filipendula

Spikelets white-villous; awns (2— )4 per raceme pair

H. filipendula var. pilosa

14(3). Awns 10-15 per raceme pair H. dregeana

Awns fewer than 9 per raceme pair 15

15(14). Pedicellate spikelets glabrous or nearly so between
nerves and margins (or shortly pilose in H.

schimperi ) 16

Pedicellate spikelets villous 19

16(15). Awns 3-5 per raceme pair; peduncles 9 mm long

or less 17

Awns (4-)6-8 per raceme pair; peduncles more

than 9 mm long 18

17(16). Awns to 1 6(— 20) mm long; spatheoles 8-18 mm

long; callus square H. cymbaria

Awns 18-30 mm long; spatheoles 14-24 mm long;

callus cuneate H. variabilis

18(16). Awns 7-17 mm long; plantsslender and rambling;

callus oblong or square; pedicellate spikelets

glabrous H. pilgeriana

Awns 20-35 mm long; plants robust, erect; callus
cuneate to acute; pedicellate spikelets glabrous or

sparsely pilose H. schimperi

19(15). Basal sheaths hairy; plants densely tufted

H. tamba

Basal sheaths without hairs; plants more loosely

tufted 20

184

20(19). Awns 7-13 mm long; spatheoles 12-23 mm long;

peduncles 3-13 mm long; callus rounded; culms

robust, with stilt roots H. umbrosa

Awns more than 15 mm long; spatheoles 20^40 mm
long; peduncles 10-30 mm long; callus cuneate;
culms robust or slender, with or without stilt

roots 21

21(20). Awns 22^10 mm long; pedicellate spikelets usually
with an awn 2-6 mm long; culms very robust,
exposed at the base, with well-developed stilt
roots, lowest internodes narrower than those

above H. rudis

Awns 15-25 mm long; pedicellate spikelets with a
short awn-point 1-3 mm long; culms robust or
slender, clad in old leaf sheaths at base, without

well-developed stilt roots, lowest intemode

similar in width to upper ones 22

22(21). Culms robust, sometimes with small stilt roots;

plants 1000-2000(-3000) m tall ... H. tamba

Culms slender, without stilt roots, arising in clumps
from a short rhizome; plants 300-1300 mm tall

H. collina

Hyparrhenia anamesa Clayton

Perennial; rhizomatous and
tufted (densely); 600-1200 mm
tall. Leaf blades to 400 mm long
(but often shorter); 4 mm wide.

Spikelets (sessile) 5. 0-6. 5 mm
long (white-villous, callus 1.0-
1.8 mm long). Racemes 15-25
mm long, with 2 homogamous
pairs at base of upper raceme,
raceme pairs with 4-7 awns 25-40 mm long having hairs
to 0.6 mm long; raceme bases terete, unequal; pedicellate
spikelets awnless or with an awn-point to 2 mm long.

Flowering October to May. Dry soils, open places. Com-
mon. Biome: Fynbos, Savanna, and Grassland. Eastern
Africa. This recently-described species is intermediate
between H. hirta, which has longer racemes, more awns and
0 or 1 homogamous pairs at the upper raceme base, and H .
filipendula, which has shorter racemes with fewer awns.
Some specimens formerly assigned to H. hirta are now
segregated in this species, but its validity in the field is not
yet assessed.

Description: Clayton et al. 1970-1982 (800), Clayton
1969 (85). Illustration: Clayton 1969, (fig. 21) Clayton et
al. 1970-1982 (fig. 184). Voucher: Rodin 3821. PRECIS
code 9900730-00100.

Hyparrhenia collina (Pilg.) Stapf

Elephant grass, olifantsgras.

Slender perennial; rhizomat-
ous and tufted (loosely);

300-1300 mm tall. Leaf blades to
300 mm long; 2-5 mm wide.

Spikelets (sessile) 4. 5-5.0 mm
long (usually dark purple with
white hairs, callus cuneate).

Spatheoles 20^40 mm long; peduncles 10-25 mm long;
raceme pairs with 4-6 awns 15-25 mm long; raceme bases
subequal, flattened; pedicellate spikelets villous.

Flowering April to May. Damp soils and dry savanna.
Infrequent. Eastern Africa to Sudan. Imperfectly separated
from the closely related H. rudis, H. dregeana and H.
tamba, but it may be distinguished by its slender culms.

Description: Clayton et al. 1970-1982 (811), Clayton
1969 (130). Voucher: Du Toil 2412. PRECIS code
9900730-00200.

Hyparrhenia cvmbaria (L.) Stapf

Boat thatching grass, bootjie-
tamboekiegras.

Robustperennial; rhizomatous
and tufted (coarsely); 2000^4000
mm tall. Leaf blades to 450 mm
long; 6-20 mm wide. Spikelets
(sessile) 3.8 — 4.5 mm long

(glabrescent to shortly pubescent,
often purplish, callus square, 0.2-0. 3 mm long). With stilt
roots; spatheoles ovate, bright reddish-brown, 8-18 mm
long; peduncles 3-8 mm long; raceme pairs with 3— 5(— 6)
awns to 16 mm long; raceme bases subequal, flattened; ped-
icellate spikelets glabrous to puberulous, margins ciliate.

Flowering November to June. Forest margins, open
hillsides. Common. Biome: Savanna and Grassland.
Tropical Africa, Madagascar and Comoro Islands. Closely
related to H. umbrosa and grades into H. variabilis, but
distinguished by its small ovate spatheoles and short square
callus.

Description: Clayton et al. 1970-1982 (804), Clayton
1969 (110). Illustration: Chippindall 1955 (pi. 16,11).
Voucher: Scheepers 190. PRECIS code 9900730-00300.

Hyparrhenia dichroa (Steud.) Stapf

Perennial (culms stout); rhizo-
matous and tufted; to 3000 mm
tall. Leaf blades to 600 mm long;
to 8 mm wide. Spikelets (sessile)

4-5 mm long (hairs pale
brownish, scanty, callus 0.4— 0.8
mm long). Panicle copiously
branched; spatheoles narrowly
lanceolate, 20^40 mm long;
raceme pairs with 6-10 awns ( 1 0 — )20— 3G mm long, racemes
often clasped at the base by spatheoles at maturity; raceme
bases terete, unequal.

Flowering March to June. Moist places, weedy places,
roadsides. Locally common. Biome: Savanna. To Sudan
and Zaire. Intergrades with its close relative H. rufa, which
has exserted racemes and more awns, and similar to H.
gazensis, which has slender culms and fewer awns, and H.
finitima, which has a thin pungent callus.

Description: Clayton et al. 1970-1982 (796), Clayton
1969 (68). Voucher: Strey & Schlieben 8597. PRECIS code
9900730-00400.

Hyparrhenia dregeana (Nees) Stapf

(-H. aucta (Stapf) Stapf ex
Stent) 1; (=H. pilosissima
(Hack.) J.G. Anders.) 1.

Harige bloutamboekiegras,
hairy blue thatching grass.

Robust perennial; rhizomatous
and tufted (densely); 1500-2000
mm tall. Leaf blades to 600 mm long; 3-8 mm wide. Spike-
lets (sessile) 4-5 mm long (densely long-hairy to shortly
hairy, rarely glabrous, callus cuneate, 1 mm long). Culms
4-9 mm across; basal sheaths hairy; spatheoles 20-25 mm
long: peduncles 15-50 mm long; raceme pairs with 10-25
awns 8-20 mm long; raceme bases subequal, flattened,
short-appendaged; pedicellate spikelets villous to
hispidulous, rarely glabrous.

Flowering November to May. Stony hillsides,
streamsides, dry soils around vleis. Common. Biome: Sa-
vanna and Grassland. Eastern Africa. Related to H. collina,
H. tamba , and H. rudis, from which it may be distinguished
by its densely tufted habit and very many short awns.

185

Description: Clayton et al. 1970-1982 (809), Clayton
1969 (124). Voucher: Liebenberg 6820. PRECIS code
9900730-00500.

Description: Clayton et al. 1970-82 (797), Clayton 1969
(72). Voucher: Van Vuuren 1685. PRECIS code 9900730-
00800.

Hyparrhenia filipendula (Hochst.) Stapf var. filipendula Hyparrhenia gazensis (Rendle) Stapf

Fine thatching grass, fyntam-
boekiegras.

Delicate and graceful perenni-
al; rhizomatous and tufted;

600-2000 mm tall. Leaf blades to
300 mm long; to 4 mm wide.

Spikelets (sessile) 5.5-7 mm long
(glabrous, callus 1. 8-3.0 mm
long). Racemes 10-12 mm long; raceme pairs with 2(^4)
awns 30-55 mm long, with hairs to 1.2 mm long; raceme
bases terete, unequal, with 2 pairs of homogamous spikelets
at base of upper raceme; pedicellate spikelets with an awn
1-5 mm long.

Flowering November to April. Woodlands, higher
rainfall areas, open veld. Common. Biome: Savanna.
Tropical Africa, Madagascar, Ceylon to Australia. Domes-
tic use (thatching). Intergrades with var. pilosa , which in
turn intergrades with H. hirta and H. anamesa , but
recognized by its graceful appearance, with many slender
branches, drooping peduncles and small, few-awned
racemes.

Description: Clayton et al. 1970-1982 (803), Clayton
1969 (95). Illustration: Chippindall 1955 (fig. 408B).
Voucher: Codd 6880. PRECIS code 9900730-00600.

Hyparrhenia filipendula (Hochst.) Stapf var. pilosa

(Hochst.) Stapf

Fyntamboekiegras, fine
thatching grass.

Perennial; rhizomatous and
tufted; 600-2000 mm tall. Leaf
blades to 300 mm long; to 4 mm
wide. Spikelets (sessile) 5. 5-7.0
mm long (white-villous, callus
1. 8-3.0 mm long). Racemes
10-12 mm long; raceme pairs with (2— )4 awns 30-55 mm
long, with hairs to 1.2 mm long; raceme bases terete,
unequal, with 2 pairs of homogamous spikelets at base of
upper raceme; pedicellate spikelets with an awn 1-5 mm
long.

Flowering December to April. Open veld and disturbed
places in higher rainfall areas. Common. Biome: Savanna
and Grassland. Tropical Africa, southern Asia to Australia.
Domestic use (thatching). Forms a bridge between var.
filipendula and H. hirta and H. anamesa, and most recently
not treated as separate from the typical variety (Clayton &
Renvoize 1982).

Description: Clayton 1969 (97), Voucher: De Winter
2863. PRECIS code 9900730-00700.

Hyparrhenia finitima (Hochst.) Anderss. ex Stapf

Robust perennial; rhizomat-
ous; 1000-2000 mm tall. Leaf
blades to 600 mm long; to 8 mm
wide. Spikelets (sessile) 5. 5-6.0
mm long (yellowish, glabrous to
shortly white hairy, callus linear,
pungent, 1-2 mm long). Raceme
pairs with 2-6 awns 25-40 mm
long; raceme bases terete,
unequal, upper raceme base 1 .5-2.5 mm long, homogamous
spikelets 0-1 pair at base of upper racemes.

Flowering December to March. Rocky places, disturbed
places. Infrequent. Biome: Savanna. Tropical Africa.
Closely related to H. gazensis, but distinguished by its
robust culms and thin callus.

Polgras.

Perennial; rhizomatous and
tufted (loosely); 500-1800 mm
tall. Leaf blades 80-200 mm
long; 2-5 mm wide. Spikelets
(sessile) 4. 0-5. 5 mm long (white-
hispidulous, callus cuneate,

0. 8-1.5 mm long). Culms

slender; raceme pairs with 4-5 awns 20-30 mm long;
raceme bases terete, unequal, upper raceme base
(2.0— )2.5— 3 .5 mm long, homogamous spikelets 0-1 pair at
base of upper racemes.

Flowering November to May. Ruderal on poor soils,
roadsides. Locally common. Biome: Savanna. Southern
tropical Africa. Related to H. finitima and H. dichroa, but
distinguished by its combination of slender culms, cuneate
callus and few awns.

Description: Clayton et al. 1970-1982 (797), Clayton
1969 (71). Voucher: De Winter & Codd 145. PRECIS code
9900730-00900.

Hyparrhenia hirta (L.) Stapf

Common thatching grass, dek-
tamboekiegras.

Perennial; rhizomatous and
tufted (wiry); 300-800 mm tall.

Leaf blades 20-150 mm long;

1 — 2( — 4) mm wide. Spikelets (ses-
sile) 4. 0-6. 5 mm long (yellowish
green to violet, white-villous,
callus acute). Culms slender; panicle scanty, of 2-10
raceme pairs, the pairs with 0-1 homogamous pairs at base
of upper racemes and 8-14 awns 10-35 mm long with hairs
to 0.3 mm long; raceme bases terete, unequal; racemes
never deflexed, 20-40 mm long.

Flowering September to June. Stony soils. Dominant.
Biome: Fynbos, Savanna, Grassland, and Nama-Karoo.
Throughout Africa to the Mediterranean and Pakistan. Do-
mestic use (thatching), or indicator (climax). The most
widespread of all the hyparrhenias, H. hirta is linked
through H. quarrei to H . nyassae, through H. anamesa to
H . filipendula and also to H.dregeana and H . finitima. H.
hirta may be recognized by its hard basal tussock, harsh
narrow leaves and scanty panicle of white villous racemes
which do not deflex.

Description: Clayton et al. 1970-1982 (798), Clayton
1969 (75). Illustration: Chippindall 1955 (fig. 408 A).
Voucher: De Winter 2579. PRECIS code 9900730-01000.

Hyparrhenia newtonii (Hack.) Stapf var. macra Stapf

Perennial; rhizomatous and
tufted (densely); 600-1200 mm
tall. Leaf blades to 300 mm long;
to 3 mm wide. Spikelets (sessile)

6-10 mm long (lower glume
hairy, callus acute to pungent,

1 .5—2.0 mm long). Basal sheaths
tomentose or glabrous, raceme
pairs with 2-4 awns 25-55 mm
long; raceme bases covered with stiff hairs, upper raceme
base with a linear scarious appendage 3-4 mm long.

Flowering December to April. Stony hillsides. Infre-
quent. Biome: Savanna and Grassland. Southern tropical
Africa. This variety is doubtfully distinct, distinguished
from the typical var. newtonii only by the hairy lower glume
of the sessile spikelet.

Fig. 108. PI. 96.

186

Description: Clayton et al. 1970-1982 (816), Clayton
1969 (150). Voucher: Louw 2720. PRECIS code

9900730-01300.

Hyparrhenia newtonii (Hack.) Stapf var. newtonii

Bearded thatching grass.

Perennial; rhizomatous and
tufted; 300-1000 mm tall. Leaf
blades to 300 mm long; to 3 mm
wide. Spikelets (sessile) 6-10
mm long (lower glume glabrous
or with a few hairs at tip, callus
acute to pungent, 1. 5-2.0 mm
long). Basal sheaths tomentose or glabrous, raceme pairs
with 2-4 awns 25-55 mm long; raceme bases covered with
stiff hairs, upper raceme base with a linear scarious
appendage 3-4 mm long.

Flowering December to March. Stony hillsides.. Infre-
quent. Biome: Savanna and Grassland. Western tropical
Africa, Madagascar, southeast Asia, Indonesia.

Description: Clayton et al. 1970-1982 (816), Clayton
1969 (149). Illustration: Clayton et al. 1970-1982 (fig.
186). Voucher: Story 1645. PRECIS code 9900730-01350.

Hyparrhenia poecilotricha (Hack.) Stapf

(=//. buchanani (Stapf) Stapf
ex Stent) 1.

Perennial; rhizomatous and
tufted; 600-1300 mm tall. Leaf
blades to 300 mm long; to 3 mm
wide. Spikelets(sessile) 5. 5-7.0
mm long (with yellow or reddish-
brown hairs; callus acute to
pungent, 1-2 mm long). Racemes pairs with 4-7 awns
25-40 mm long; raceme bases terete, unequal, upper
raceme base 3. 5-7.0 mm long, with 2 pairs of homogamous
spikelets.

Flowering December to April. Bushveld. Locally com-
mon. Biome: Savanna and Grassland. Eastern tropical
Africa. A variable species that connects//. rufa,H. nyassae,
H .filipendula and the tropical species H.familiaris (Steud.)
Stapf, probably through introgressive hybridization. It may
be recognized by its tendency to a long upper raceme base.

Description: Clayton et al. 1970-1982 (796), Clayton
1969 (69). Voucher: Giess, Volk & Bleissner 6452.
PRECIS code 9900730-01600.

Hyparrhenia nyassae (Rendle) Stapf

Hyparrhenia quarrei Robyns

Bronsaartamboekiegras,
bronze awned thatching grass.

Perennial; rhizomatous and
tufted; 600-1300 mm tall. Leaf
blades to 450 mm long; 2-5 mm
wide. Spikelets (sessile) 5-6 mm
long (yellowish-green to violet,
with golden-yellow hairs, the
callus linear or narrowly cuneate, 0.8-1. 2 mm long). Basal
sheaths with dense spreading white hairs; raceme pairs with
6-14 awns 20^10 mm long; raceme bases terete, unequal,
upper raceme base 2-3 mm long.

Flowering November to March. Moist places in open
veld. Locally common. Biome: Savanna and Grassland.
Tropical Africa and southeast Asia. Related to H. rufa ,
which has no hairs on the basal sheaths, a shorter callus and
less hairy racemes. H. nyassae also intergrades with H.
quarrei, which has white raceme hairs.

Description: Clayton et al. 1970-1982 (793), Clayton
1969 (53). Voucher: De Winter & Codd 432. PRECIS code
9900730-01400.

Hyparrhenia pilgeriana C.E. Hubb.

Slender perennial; rhizomat-
ous and tufted (but lax and
rambling); 300-600 mm tall. Leaf
blades 50-1 10 mm long; 2-4 mm
wide. Spikelets (sessile) 4 mm
long (glabrous or with very short
white hairs, callus oblong or
square). Peduncles 9-30 mm
long, racemes exserted from
spatheoles; raceme pairs with 6-7 awns 7-17 mm long;
raceme bases subequal or the upper somewhat longer (to 1 .5
mm), flattened, shortly appendaged; pedicellate spikelets
glabrous.

Flowering February to March. Seasonal swamps, old
fallow land. Infrequent. Biome: Grassland. Eastern Africa.
Resembles H. cymbaria, which is a robust plant with
racemes enveloped by the short spatheoles, and H. gazensis,
which has longer awns.

Description: Clayton et al. 1970-1982 (807), Clayton
1969 (115). Illustration: Clayton 1969 (fig. 27). Voucher:
McClean 101. PRECIS code 9900730-01500..

Perennial; short rhizomatous
and tufted; 1000-2000 mm tall.

Leaf blades to 400 mm long; to 5
mm wide. Spikelets (sessile)

4. 5-5. 5 mm long (white pubes-
cent to villous, callus slender,

0.7-1. 2 mm long). Raceme pairs
with 6-10 awns 18-36 mm long;
raceme bases unequal, terete,
with 0 or 1 homogamous pairs at base of upper racemes;
racemes deflexed at maturity.

Flowering January to June. Forest margins. Common.
Biome: Savanna and Grassland. Tropical Africa. This
species links H. hirta, which does not have deflexed
racemes, and H. nyassae, which has yellow raceme hairs,
and may be a product of introgression between these
species.

Description: Clayton et al. 1970-1982 (799), Clayton
1969 (82). Voucher: Pole Evans 3699. PRECIS code
9900730-01700.

Hyparrhenia rudis Stapf

Robustperennial; rhizomatous
and tufted (coarsely); 2000-3000
mm tall. Leaf blades 300-600
mm long; 3-18 mm wide. Spike-
lets (sessile) 5-6 mm long (pale
or reddish brown, with silky
white hairs, callus cuneate).

Culms to 8 mm thick, exposed at
base, with stilt roots; spatheoles
25-40 mm long; peduncles 10-20 mm long; raceme pairs
with 4-7 awns 22-40 mm long; raceme bases subequal,
flattened, short-appendaged; pedicellate spikelets villous,
with an awn 2-6 mm long.

Flowering February to May. Moist soils. Locally com-
mon. Biome: Savanna and Grassland. Central Africa,
Madagascar. Closely related to H. dregeana, H. tamba and
H. collina , from which it is distinguished by its long awns
and loosely tufted culms that increase in diameter above the
lowest internodes, and to H. schimperi, which has glabrous
or sparsely hairy pedicellate spikelets.

Description: Clayton et al. 1970-1982 (811), Clayton
1969 (128). Voucher: Scheepers 242. PRECIS code
9900730-01800.

187

Hyparrhenia rufa (Nees) Stapf var. rufa

often violet-tinged, usually glossy, glabrous or with scanty
reddish-brown hairs, callus rounded or wedge-shaped,
0.2-0. 8 mm long). Panicle lax or contracted; spatheoles
linear, 40-50 mm long; raceme pairs with 7-14 awns 20-30
mm long; raceme bases terete, unequal.

Flowering December to June. Disturbed moist places
and roadsides. Common. Biome: Savanna. Tropical Africa,
introduced to America. Domestic use (thatching), or pasture
(when young). A widespread, common and very variable
species, best recognized by the glossy lower glume. It is
closely related to H. dichroa, which has fewer awns and
often has spatheoles clasping the raceme bases, and H.
poecilotricha , which has longer sessile spikelets.

Description: Clayton et al. 1970-1982 (794), Clayton
1969 (62). Illustration: Hitchcock & Chase 1950 (fig.
1667). Voucher: Scheepers 215. PRECIS code

9900730-01900.

Hyparrhenia schimperi (A. Rich.) Stapf

Robust, erect perennial;
shortly rhizomatous and tufted
(coarsely); 2000-4000 mm tall.

Leaf blades to 600 mm long; to 20
mm wide. Spikelets (sessile) 4-5
mm long (sparsely hairy to nearly
glabrous, callus cuneate to acute).

With stilt roots; culms to 8 mm
thick; peduncles 10-15 mm long;
raceme pairs with 6-8 awns 20-35 mm long; raceme bases
subequal, flattened and short-appendaged; pedicellate
spikelets glabrous to sparsely pilose.

Flowering December to May. Open moist places.
Locally common. Biome: Fynbos, Savanna, and Grassland.
Eastern Africa and Madagascar. Grades into H. variabilis
which has fewer awns, and closely related to H. rudis,
which has long hairs on the pedicellate spikelets.

Description: Clayton et al. 1970-1982 (808), Clayton
1969 (118). Voucher: Codd 208. PRECIS code

9900730-02000.

Hyparrhenia tamba (Steud.) Stapf

Fig. 109.

{-H. glauca Stent) 1 .

Bloutamboekiegras, blue
thatching grass.

Stout, robust perennial; rhizo-
matous and tufted (densely);

1000-3000 mm tall. Leaf blades
to 800 mm long; 3-7 mm wide.

Spikelets (sessile) 5 mm long (becoming dark purplish-grey,
with long white hairs, callus cuneate). Sometimes with
small stilt roots; culms to 4 mm thick; basal sheaths hairy;
spatheoles 26-40 mm long; peduncles 20-30 mm long;
raceme pairs with 5-8 awns 1 6—25 mm long; raceme bases
subequal, flattened, short-appendaged; pedicellate spikelets
villous.

Flowering December to June. Streamsides and
roadsides. Common. Biome: Savanna and Grassland.
Tropical Africa. Domestic use (thatching). Closely related
to H. collina and H. dregeana, and possibly imperfectly
separated from the latter, but distinguished by its
combination of appendaged raceme base and few awns.

Description: Clayton et al. 1970-1982 (810), Clayton
1969 (126). Illustration: Chippindall 1955 (PI. 24), Flower.
PI. Afr. (47: 1842). Voucher: Killick 2359. PRECIS code
9900730-02100.

Hyparrhenia umbrosa (Hochst.) Anderss. ex Clayton

Robust perennial; rhizomat-
ous; 1300-2000 mm tall. Leaf
blades to 600 mm long; to 12 mm
wide. Spikelets (sessile) 4 mm
long (with long white hairs, callus
0.4 mm long, oblong with
rounded tip). With stilt roots;
culms rambling and slender
below, increasing to 6 mm across
above; spatheoles 12-23 mm long; peduncles 3-13 mm
long; raceme pairs with 4-6 awns 7-13 mm long; raceme
bases flattened, subequal; pedicellate spikelets villous.

FloweringMay, June, and July. Roadsides, old lands. In-
frequent. Biome: Savanna and Grassland. Tropical Africa.
Closely related to H. cymbaria and H rudis, and possibly
not a distinct species, but distinguishable by the rambling
culm bases and oblong callus.

Description: Clayton et al. 1970-1982 (810), Clayton
1969 (127). Voucher: Pole Evans 3775. PRECIS code
9900730-02200.

Hyparrhenia variabilis Stapf

Robust perennial; rhizomat-
ous; 1500-3000 mm tall. Leaf
blades to 450 mm long; to 15 mm
wide. Spikelets (sessile)4-5 mm
long (nearly glabrous to sparsely
and shortly white-hairy, callus
cuneate, 0.5-1. 0 mm long). With
stilt roots; culms 3. 0-5. 5 mm
across; spatheoles 14-24 mm
long; peduncles 3-9 mm long; raceme pairs with 3-5 awns
18-30 mm long; raceme bases subequal, flattened and
short-appendaged; pedicellate spikelets glabrous, except
margins ciliate.

Flowering January to May. Forest margins. Locally
common. Biome: Savanna. Eastern Africa, Madagascar,
Comoro Islands, Java. Grades into H. cymbaria, which has
short ovate spatheoles and a square callus, and H.
schimperi, which has more awns.

Description: Clayton et al. 1970-1982 (805), Clayton
1969 (113). Illustration: Clayton et al. 1970-1982 (fig.
185). Voucher: Galpin 8887. PRECIS code

9900730-02300.

Hyperthelia Clayton

Sometimes included in Hyparrhenia ; including H.
dissoluta.

Annual, or perennial; caespitose. Culms 1000-7500 mm
high', herbaceous; branched above (to form compound
inflorescences). Culm internodes hollow. Leaf blades
linear; flat, or rolled (on drying). Ligule an unfringed
membrane (usually), or a fringed membrane (rarely).
Plants bisexual, with bisexual spikelets. The spikelets of
sexually distinct forms on the same plant (hermaphrodite
and male)', overtly heteromorphic.

Inflorescence of spike-like main branches, or paniculate
(a large, leafy false panicle)', spatheate; a complex of
‘partial inflorescences' and intervening foliar organs.
Spikelet-bearing axes very much reduced, or ‘racemes’
(rarely); paired', with very slender rachides; disarticulating
at the joints. ‘Articles’ appendaged (raceme-base with a
long scarious appendage at the tip, which opposes the basal
homogamous spikelets in the bud).

Spikelets in pairs, or in triplets (sometimes having one
female-fertile spikelet with a pair of pedicellate male
spikelets, the triplet disarticulating in its entirety); consis-
tently in ‘long-and-short’ combinations; these pedicellate/

sessile. Pedicels free of the rachis. The sessile spikelets her-
maphrodite (in the heterogamous combinations), or male-
only (in the homogamous combinations). The pedicellate
spikelets male-only. The homogamous and pedicellate
spikelets male, linear-lanceolate, with two hyaline lemmas.
Female-fertile spikelets 8-35 mm long; compressed dorsi-
ventrally; falling with the glumes. Glumes two; more or less
equal; awned (G2 sometimes aristate), or awnless; very dis-
similar. Proximal incomplete florets 1\ epaleate; sterile.

189

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline at margins and tips); incised; awned. Awns
1 ; median; from the sinus; geniculate; much longer than the
body of the lemma. Palea present, or absent; when present
conspicuous but relatively short, or very reduced. Lodicules
2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
narrowly ellipsoid; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 6 species. Tropical and southern
Africa. Mesophytic; in open habitats (grasslands and
savanna); glycophytic. Namibia, Botswana, Transvaal,
Swaziland, and Natal. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russel! & M.
Koekemoer.

Hyperthelia dissoluta (Nees ex Steud.) Clayton

(=Hyparrhenia dissoluta
(Steud.) C.E. Hubb.) 1.

Geeltamboekiegras, yellow
thatching grass.

Robust perennial; tufted;

1000-3000 mm tall. Leaf blades
to 300 mm long; to 12 mm wide.

Spikelets (sessile) 6. 5-7. 5 mm long (pedicellate to 14 mm
long). Plant yellow and green; culms and awns yellow,
leaves and spikelets green; spikelets glabrous, lower glume
of sessile spikelets narrowly grooved, awns 50-100 mm
long.

Flowering throughtout the year (mostly in autumn).
Roadsides and disturbed places. Common. Biome: Savanna.
Tropical Africa, Madagascar. Domestic use (thatching).

Description: Chippindall 1955 (512), Clayton et al.
1970-1982 (786). Illustration: Chippindall 1955 (fig. 410),
Clayton et al. 1970-1982 (fig. 183). Voucher; Compton
27058. PRECIS code 9900731-00100.

Imperata Cirillo

Syllepis Fourn.

Perennial; long-rhizomatous. Culms 100-1500 mm
high; herbaceous; unbranched above. Ligule a fringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets all alike in sexuality ; homomorphic.

Inflorescence paniculate (spiciform or loosely
contracted, the branches with numerous short ‘racemes' ,
with dense silky white hairs)-, contracted ; espatheate; not
comprising ‘partial inflorescences’ and foliar organs.
Spikelet-bearing axes short ‘racemes’; with very slender
rachides; persistent.

Spikelets in pairs-, consistently in ‘long-and-short’ com-
binations; unequally pedicellate in each combination.
Pedicels free of the rachis. The short-pedicellate spikelets
hermaphrodite. The long-pedicellate spikelets hermaphro-
dite. Female-fertile spikelets compressed dorsiventrally;
falling with the glumes (falling entire from their pedicels).
Glumes two; more or less equal ; awnless; similar
(membranous, with long silvery hairs especially towards
the base). Proximal incomplete florets /; paleate (rarely),
or epaleate; male (rarely), or sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire, or incised (denticulate); awnless.
Palea present; relatively long, or conspicuous but relatively
short, or very reduced (broad). Stamens 1, or 2 (rarely 3?).
Ovary glabrous. Fruit small; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 8 species. Tropical and
subtropical. Helophytic, or mesophytic, or xerophytic; in

open habitats (often in damp or weedy places); maritime-
arenicolous (some forms of/, cylindrica), or glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. 1 indige-
nous species.

Intergeneric hybrids procured with Saccharum.

Fig. 111. Imperata cylindrica

Fig. 1 10. PI. 97.

190

References. 1. Launert. 1970. FSWA. 2. Clayton &
Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Imperata cylindrica (L.) Raeusc

(=/. cylindrica (L.)

Raeuschel var. africana
(Anderss.) C.E. Hubb.) 1, 2; (= 1 .
cylindrica (L.) Raeuschel var.
major (Nees) C.E. Hubb.) 1, 2.

Sygras, cottonwool grass,
donsgras, silverspike.

Perennial; strongly rhizomatous; 100-1200 mm tall.
Leaf blades to 1500 mm long; 2-12 mm wide. Spikelets all
alike 3-6 mm long. Leaves broad in the middle, narrowed
at tip and base, reddish in winter; panicle dense, silky,
white, cylindrical.

Flowering August to June. Riverbanks, vleis and
seasonally wet places. Common. Biome: Fynbos, Savanna,
and Grassland. Old World tropics. Weed (because the tough
rhizomes make it difficult to eradicate).

Description: Chippindall 1955 (476), Clayton et al.
1970-1982 (700). Illustration: Chippindall 1955 (fig. 392),
Clayton et al. 1970-1982 (fig. 159). Voucher; Gibbs
Russell 2197. PRECIS code 9900370-00050.

Fig. 111. PI. 98.

Ischaemum L.

Argopogon Mimeur, Collardoa Cav., Ischaemopogon
Griseb., Meoschium P. Beauv.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 100-3500 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear (usually), or linear-lanceolate to
lanceolate; flat. Ligule an unfringed membrane. Plants
bisexual, with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant, or all alike in sexuality;
overtly heteromorphic (the pedicellate spikelet sometimes
much smaller, often asymmetric), or homomorphic.

Inflorescence of spike -like main branches (terminal or
axillary ); digitate or subdigitate (usually)', spatheate
(uppermost leaf reduced to a spatheate sheath), or espathe-
ate; not comprising ‘partial inflorescences’ and foliar
organs. Spikelet-bearing axes ‘racemes’; paired, or
clustered; with substantial rachides (these stout,
triangular)', disarticulating at the joints.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; in pedicellate/sessile combinations, or unequally
pedicellate in each combination. Pedicels free of the rachis.
The ‘shorter’ spikelets hermaphrodite. The ‘longer’
spikelets hermaphrodite, or male-only (rarely), or sterile
(rarely). Female-fertile spikelets compressed dorsiven-
trally; falling with the glumes. Glumes two; more or less
equal; awned, or awnless; very dissimilar (lower
coriaceous, usu. 2-keeled; upper 1 -keeled above, sometimes
awned). Upper glume 5-11 nerved. Proximal incomplete
florets 1 ; paleate, palea fully developed; male.

Female-fertile florets 1. Lemmas less firm than the
glumes (firmly membranous); incised', mucronate, or awned
(usually). Awns when present 1; from the sinus (or
mucronate); geniculate; much shorter than the body of the
lemma, to much longer than the body of the lemma. Palea
present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Hilum' short; embryo large.

Cytology, classification, distribution. Chromosome base
number,* = 10. Panicoideae; Andropogonodae; Andropo-

goneae; Andropogoninae. 60 species. Tropical and
subtropical. Helophytic (mostly), or mesophytic, or xero-
phytic; in shade, or in open habitats (damp or shady places);
maritime-arenicolous (e.g. /. muticum, I. triticeum), or
glycophytic. Namibia, Botswana, Transvaal, Swaziland,
Natal, and Cape Province. 2 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf blades green, usually reddish-tinged, usually
wider than 5 mm, narrowing shortly to a sharp
point; lower glume of sessile spikelets convex or

flat, keels usually winged on upper half

I. fasciculatum

Leaf blades glaucous, usually narrower than 5 mm,
drawn out into a long fine tip; lower glume of

sessile spikelets concave, keels not winged

I. afrum

191

PI. 99.

Ischaemum afrum (J.F. Gmel.) Dandy

(=/. brachyatherum (Hochst.)
Hack.) 1; (=/. glaucostachyum
Stapf) 1.

Turfgras, tweevingergras.

Perennial; rhizomatous; to
1200 mm tall. Leaf blades
100-500 mm long; 2— 5(— 1 1) mm

wide. Spikelets (sessile) 5-8 mm long (pedicellate smaller).
Leaves glaucous, tips tapering to a long, fine point; lower
glume of sessile spikelets concave, keels not winged.

Flowering October to April. Black turf soil, usually near
water. Common and locally dominant (sometimes). Biome:
Savanna and Grassland. Throughout tropical Africa to
India. Weed (ruderal).

Description: Chippindall 1955 (487), Clayton et al.
1970-1982 (747). Illustration: Clayton et al. 1970-1982
(fig. 175). Voucher: Giess, Volk & Bleissner 6436. PRECIS
code 9900100-00100.

similar in form to the laterals; apical; non-geniculate; when
present much shorter than the body of the lemma to about
as long as the body of the lemma. Palea present; relatively

Fig. 112.

Ischaemum fasciculatum Brongn.

(=/. arcuatum (Nees)

Stapf) 1.

Rooivleigras.

Perennial; rhizomatous; 300-
900 mm tall. Leaf blades 50-250
mm long; (4-)5-16 mm wide.

Spikelets (sessile) 5-6 mm
long. Leaves green, becoming reddish-brown, tips
narrowing shortly to a sharp point; lower glume of sessile
spikelets convex or flat, keels usually winged on upper half;
awns 5-10 mm long.

Flowering October to May. Wet places, vleis and
riverbanks. Biome: Savanna and Grassland. Throughout
tropical Africa to southeast Asia. Weed (ruderal).
Occasional individuals with hairy leaves resemble Eulalia
villosa in reddish colour and inflorescence form, but in E.
villosa the lemma awns reach about 15-20 mm long.

Description: Chippindall 1955 (487), Clayton et al.
1970-1982 (749). Illustration: Chippindall 1955 (fig. 398).
Voucher: De Winter & Wiss 4315. PRECIS code
9900100-00200.

Kaokochloa De Winter

Annual (all vegetative parts pilose)', culms geniculate or
prostrate at base, rooting at nodes. Culms 150-800 mm
high; herbaceous; branched above. Leaf blades linear-
lanceolate to lanceolate; flat, or rolled. Ligule a fringe of
hairs.

Inflorescence paniculate', open to contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 5-8 mm long; not noticeably compressed
(‘ sub globose' ); disarticulating above the glumes (or
between the glumes, the upper glume falling with the
spikelet); not disarticulating between the florets. Glumes
two; more or less equal; about equalling the spikelets;
awnless; similar. Lower glume 9 nerved, or 1 1 nerved. In-
complete florets distal to the female-fertile florets, merely
underdeveloped, awnless (with only minute vestiges of
awns); proximal incomplete florets absent.

Female-fertile florets 3-6. Lemmas without a germin-
ation flap; 9 nerved; incised (between excurrent nerves);
awned. Awns 2, or 3, or 5; median and lateral (sometimes),
or lateral only (the two marginal nerves excurrent into large
awns, the median and other nerves occasionally
contributing smaller awns). The median awn when present

Fig. 113. Kaokochloa nigrirostris

192

long (but narrower than the lemma). Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Hilum short; pericarp
fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions absent. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chloridoideae;
Pappophoreae. 1 species. Southern Africa. Xerophytic; in
open habitats (in semi-desert); glycophytic. Namibia. 1 in-
digenous species.

References. 1. De Winter. 1961. Bothalia 7: 479.

Species treatment by G.E. Gibbs Russell.

Kaokochloa nigrirostris De Winter

Fig. 113. PI. 100.

Annual; loosely tufted;

200-600 mm tall. Leaf blades
50-1 20 mm long; 5-10 mm wide.

Spikelets to 7 mm long; to 6 mm
wide. Culms decumbent; lemmas
with two lateral nerves running
out into large awns and central
nerves forming smaller awns,
lemmas curled inward at base of
awns; awns glabrous, purple-tinged.

Flowering March to June. Flats and hillsides, in sandy
or gravelly soil. Conservation status not known. Biome:
Nama-Karoo. Endemic.

Description: De Winter 1961 (480). Voucher: De Winter
& Leistner 5848. PRECIS code 990361 1-00100.

Karroochloa Conert & Tuerpe

Sometimes included in Rytidosperma , Danthonia sensu
lato.

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 40^)00 mm high; herbaceous; unbranched above.
Leaf blades linear; to 2 mm wide ; flat, or folded, or rolled:
not disarticulating. Ligule a fringe of hairs.

Inflorescence paniculate ; contracted (10-60 mm long):
more or less ovoid ; espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary; 4-6 mm long (rarely to 7 mm); com-
pressed laterally; disarticulating above the glumes. Hairy
callus present. Glumes two: more or less equal (subequal);
about equalling the spikelets; awnless; similar
(membranous, margins and apices hyaline). Incomplete
florets distal to the female-fertile florets, merely underde-
veloped, awned; proximal incomplete florets absent.

Female-fertile florets 3-7. Lemmas similar in texture to
the glumes (membranous); hairy (with fringes or tufts of
white hairs, except in K. curva , the hairs in tufts, or not in
tufts; in transverse rows, or not in transverse rows); without
a germination flap; 9 nerved; incised; awned. Awns 1 , or
i; median, or median and lateral (by small extensions from
the lobes). The median awn different in form from the
laterals (when laterals present); from the sinus: geniculate:
about as long as the body of the lemma to much longer than
the body of the lemma. Palea present; relatively long
(almost equalling the lemma); 2-nerved. Lodicules 2;
fleshy; ciliate. Stamens 3. Ovary glabrous. Fruit small
(0.8-1 mm); hilum short; pericarp fused; embryo small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x = 6. Arundinoideae; Danthonieae. 4 species.
Southern Africa. Mesophytic; in open habitats (grassland
and among rocks); glycophytic. Namibia, Orange Free
State, Natal, Lesotho, and Cape Province. 4 indigenous
species.

References. 1. Conert & Tuerpe. 1969. Senckenb. Biol.
50: 333.

1(0). Lemmas with hairs in tufts, glabrous between tufts;

leaves and sheaths sparsely hispid 2

Lemmas with a row of hairs across backs below awn
base, glabrous or pubescent below this; leaves
usually glabrous, but if pubescent then never hispid

3

2(1). Lemmas3.0-3.5 mm long, including lobes; tufts of
hairs on lemma up to 2 mm long; palea 2. 8-3. 2 mm
long, sparsely pubescent between the margins and

keels; perennial K. purpurea

Lemmas 1.8-2. 5 mm long, including lobes; tufts of
hairs not longer than 1.2 mm; palea 1. 8-2.4 mm

long, glabrous; annual K. tenella

3(1). Plants annual; lemmas sparsely pubescent or glabrous
below row of hairs across back, margins fringed

with hairs K. schismoides

Plants perennial; lemmas densely pubescent below
row of hairs across back, margins not obviously
fringed K. curva

Species treatment by N.P. Barker.

Fig. 114. Karroochloa curva

193

Karroochloa curva (Nees) Conert & Tuerpe

Fig. 1 14.

(=Danthonia curva Nees) 1.

Perennial; stoloniferous and
tufted; to 400 mm tall. Leaf
blades to 250 mm long; to 2 mm
wide. Spikelets 5-6 mm long;
about 1.5 mm wide. Leaf sheaths
usually glabrous; leaf blades flat
or folded, usually glabrous, but if
pubescent then never hispid; panicle 15-50 mm long; spike-
lets 3-6-flowered; glumes 3. 5-6.0 mm long; lemmas

2. 5- 3. 5 mm long including lemma lobes which extend into
short, soft bristles; back of lemmas with a row of hairs
across the middle below the awn base, densely pubescent
below this, margins not obviously fringed; central awn

4. 0- 5. 5 mm long; palea 2. 2-2. 5 mm long, pubescent
between the keels.

Flowering October to May. In damp or shady habitats.
Common. Biome: Fynbos, Nama-Karoo, and Grassland.
Endemic. Natural pasture. Similar to A', schismoides, which
is annual and has a fringe of hairs along the lemma mar-
gins. The flowering time is dependent on the seasonality of
the rains.

Description: Conert & Tuerpe 1969 (295), Stapf
1898-1900 (532), Chippindall 1955 (243). Illustration:
Conert & Tuerpe 1969 (fig. 2-8, spikelet parts only), Chip-
pindall 1955 (fig. 214). Voucher: Du Toit 1999. PRECIS
code 9902044-00100.

Karroochloa purpurea (L.f) Conert & Tuerpe

PI. 101.

(-Danthonia purpurea
(Thunb.) Beauv. ex Roem. &

Schult.) 1.

Perennial; shortly rhizomatous
and tufted; to 220 mm tall. Leaf
blades to 40 mm long; to 1 mm
wide. Spikelets 5-7 mm long; to
4 mm wide. Leaf sheaths sparsely
hispid; blades rolled, falcate, sparsely hispid; panicle 10-20
mm long; spikelets 3-6-flowered; glumes 4-7 mm long,
often tinged with dark purple around the keel; lemmas

3.0- 3. 5 mm long, including the truncate lemma lobes;
backs of the lemmas with numerous tufts of hairs in a row
across the middle of the back below the awn base as well
as basal tufts near the margins and central nerve; hair tufts
to 2 mm long; central awn 3-4 mm long, geniculate; palea
2. 8-3. 2 mm long, pubescent between the margin and keels.

Flowering July to May. In mountainous areas and in
short grasslands. Common (roadsides). Biome: Fynbos,
Grassland, and Nama-Karoo. Endemic. Natural pasture (for
sheep). This species is similar toK. tenella , which is annual
and has shorter tufts of hairs on the lemmas.

Description: Conert & Tuerpe 1969 (303), Stapf 1 898—
1900 (530), Chippindall 1955 (244). Illustration: Conert &
Tuerpe 1969 (fig. 24-30), Chippindall 1955 (fig. 215).
Voucher: Barker 33. PRECIS Code 9902044-00200.

Karroochloa schismoides (Stapf ex Conert) Conert &

Tuerpe

Annual; tufted; 50-150 mm
tall. Leaf blades to 60 mm long;
about 0.5 mm wide. Spikelets

4. 5- 6.0 mm long; about 1 mm
wide. Leaf sheath usually
glabrous; leaf blades linear, open
or rolled, glabrous or pubescent
but then never hispid; panicle
10-20 mm long; spikelets 3-5-flowered; glumes 3. 5-5.0
mm long; lemmas 2. 5^1.0 mm long including lemma lobes
which extend into short, soft bristles; back of lemmas with
a fringe of hairs across the middle below the awn base.

sparsely pubescent or glabrous below this, with a fringe of
hairs along each margin; central awn 3.0^4. 5 mm long;
palea 2.2-2.4 mm long, pubescent between the keels.

Flowering dependent upon rainfall, usually July to Oct-
ober. Dry mountains. Common (in the drier parts of the
north west Cape and Namibia). Biome: Nama-Karoo and
Succulent Karoo. Endemic. Similar to K. curva , which is a
perennial and which has a densely pubescent lemma back.

Description: Conert & Tuerpe 1969 (299), Launert 1970
(160:125). Illustration: Conert & Tuerpe 1969 (fig. 13-19,
spikelet parts). Voucher: Munro s.n. PRECIS code
9902044-00300.

Karroochloa tenella (Nees) Conert & Tuerpe

( =Danthonia tenella Nees) 1.

Annual; tufted; 40— 1 50(— 250)
mm tall. Leaf blades 5-15 mm
long; to 0.8 mm wide. Spikelets
4-7 mm long; to 2 mm wide. Leaf
sheaths sparsely hispid; leaf
blades rolled, falcate, sparsely
hispid; panicle 5-20 mm long;
spikelets 3-5-flowered; glumes 4-7 mm long, sometimes
tinged with purple at apex; lemmas 1.8-2. 5 mm long,
including truncate lobes; backs of the lemmas with
numerous tufts of hairs positioned in a row across the
middle of the back below the awn base as well as basal tufts
near the margins and central nerve; hair tufts 0.5-1. 2 mm
long; central awn 2. 8-4.0 mm long, geniculate; palea
1.8-2. 4 mm long, glabrous.

Flowering June to October. Sandy soils. Common (in
disturbed areas). Biome: Fynbos, Nama-Karoo, and Succu-
lent Karoo. Endemic. In the Van Rhynsdorp area plants can
reach a height of 250 mm. Very similar to K. purpurea ,
which is perennial, shortly rhizomatous and has longer tufts
of hairs on the lemmas.

Description: Conert & Tuerpe 1969 (308), Stapf
1898-1900 (531), Chippindall 1955 (244). Illustration:
Conert and Tuerpe 1969 (fig. 36—4 1 , spikelets parts only).
Voucher: Davidse 33381. PRECIS code 9902044-00400.

Koeleria Pers.

Aegialina Schult., Aegialitis Trin., Airochloa Link,
Brachystylus Dulac, Ktenosachne Steud., Leptophyllochloa
Cald., Poarion Reichenb., Wilhelmsia Koch, sometimes
includes Lophochloa Reichenb., Rostraria Trin.

Annual ( Lophochloa ), or perennial; long-rhizomatous
(rarely), or caespitose. Culms 50-1200 mm high; herba-
ceous. Leaf blades linear; flat, or folded, or rolled
(convolute). Ligule an unfringed membrane (sometimes
puberulent and ciliolate).

Inflorescence paniculate', contracted (dense, cylindrical,
ovoid, not interrupted)', espatheate. Spikelet-bearing axes
persistent.

Spikelets not secund', 4-7 mm long', compressed
laterally; disarticulating above the glumes. Glumes two;
very unequal, or more or less equal; markedly shorter than
the spikelets, or about equalling the spikelets; awnless;
similar. All florets female-fertile, or distal incomplete
florets also present; proximal incomplete florets absent.

Female-fertile florets 2^1. Lemmas similar in texture to
the glumes; carinate', 3-5 nerved; entire (usually), or
incised (e.g. in Lophochloa ); awnless, or mucronate, or
awned (but the awns relatively inconspicuous, by contrast
with Trisetum). Awns 1 (straight, subterminal,

inconspicuous in the inflorescence); median; from the
sinus, or dorsal; non-geniculate; much shorter than the body
of the lemma to about as long as the body of the lemma.

194

Fig. 115. Koeleria capensis

Palea present; relatively long; thinner than the lemma
(membranous). Lodicules 2; membranous; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short, or
long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. About 60
species. North and south temperate. Mesophytic, or xero-
phytic; mostly in open habitats (in dry grassland and rocky
places). Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. Indigenous species (1).

Intergeneric hybrids with Trisetum: X Trisetokoeleria
Tsvelev.

References. 1. Clayton. 1970. FTEA.

Species treatment by T.M. Sokutu.

Koeleria capensis (Steud.) Nees

(=K. cristata auctt., non (L.)

Pers. var. cristata) 1; (-K.
cristata var. brevifolia (Nees)

C.E. Hubb.) 1; ( =K . cristata var.
convoluta (Steud.) C.E.

Hubb.) 1.

Perennial; tufted (to densely
so); 150-800 mm tall. Leaf blades

Fig. 115. PI. 102.

40-200 mm long; 1-4 mm wide. Spikelets 3. 5-4.0 mm
long. Inflorescence spiciform, sometimes interrupted;
spikelets 2-4-flowered, paleas projecting out of the florets
and thus conspicuous in mature spikelets.

Flowering October to January. Common in montane
areas, often among rocks and steep slopes, dry to wet areas.
Locally common (in high altitudes in Natal), or locally
dominant (sometimes). Biome: Fynbos and Grassland.
Throughout Africa. Planted pasture (to an extent). I fail to
see how this species differs from the temperate European
K. cristata (L.) Pers. The comment by Clayton (1970) on
differences in old leaf sheaths does not seem to hold and
is considered subjective.

Description: Adams. & Salter 1950 (84), Stent 1924 in
Bothalia (1:301), Stapf 1898-1900 (468), Chippindall 1955
(83), Clayton et al. 1970-1982 (79). Illustration: Chippin-
dall 1955 (fig. 54). Voucher: Behr 899, Codd 3155.
PRECIS code 9903740-00050.

Lagurus L.

Avena Scop.

Annual ; caespitose. Culms 80-500(-600) mm high
(slender); herbaceous; unbranched above. Leaf blades
linear-lanceolate\ flat. Ligule an unfringed membrane, or
a fringed membrane (rarely).

Inflorescence paniculate ; contracted; more or less ovoid
( silky-white hairy & bristly ); espatheate. Spikelet-bearing
axes persistent.

Spikelets 5-10 mm long; compressed laterally; disartic-
ulating above the glumes. Glumes two; more or less equal;
awned; similar (narrowly lanceolate, membranous, hairy,
tapering into fine bristles, thinly membranous). All florets
female-fertile\ proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (membranous);
5 nerved; incised; awned. Awns 3\ median and lateral (with
two short terminal laterals in addition to the longer median).
The median awn different in form from the laterals; dorsal;
geniculate; much longer than the body of the lemma. Palea
present; relatively long (but shorter than the lemma). Lodi-
cules 2; membranous; glabrous. Stamens 3. Ovary glabrous.
Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Aveneae. 1 species.
Mediterranean. Xerophytic; in open habitats (especially
maritime sands). Transvaal and Cape Province. 1
naturalized species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Lagurus ovatus L.

Fig. 116. PI. 103.

Harestail, haasgras.

Velvety annual; to 600 mm
tall. Leaf blades to 100 mm long;
to 10 mm wide. Spikelets 7-10
mm long (excluding long awns to
25 mm long). Panicle compact,
oval, soft from profuse spreading
glume hairs and fine awns from
glumes and lemmas.

Flowering October to November (rarely later).
Disturbed places, usually sandy soil. Locally common. Nat-
uralized from the Mediterranean. Widely naturalized. Do-
mestic use (dried flower arrangements).

Description: Chippindall 1955 (96). Illustration: Chip-
pindall 1955 (fig. 68). Voucher: Jacot Guillarmod 3951.
PRECIS code 9902610-00100.

195

Fig. 116. Lagurus ovatus

Lamarckia Moench mut. Koeler

Achyrodes Boehmer, Chrysurus Pers., Pterium Desv.,
Tinaea Garzia.

Annual ; caespitose. Culms 70-2000(-3000) mm high;
herbaceous. Leaf blades flat. Ligule an unfringed
membrane. The spikelets of sexually distinct forms on the
same plant (only the terminal spikelet in each fascicle being
hermaphrodite, the other 3—4 male-only or with 3-6 empty,
awnless, truncate lemmas).

Inflorescence paniculate and a false spike, with clusters
of spikelets on reduced axes; contracted; espatheate.
Spikelet-bearing axes disarticulating ; falling entire (the
clusters of 3-5 spikelets falling whole).

Female-fertile spikelets secund; not in distinct ‘long-
and-short’ combinations; 3.5 mm long; compressed
laterally; falling with the glumes (in the clusters). The
sterile spikelets with many florets, narrow-elongated.
Glumes two; more or less equal; about equalling the
spikelets; awned to awnless (acuminate to shortly aristate);
similar (membranous, linear-lanceolate, hyaline). Incom-
plete florets distal to the female-fertile florets, awned (the
sterile rudiment with a long awn); proximal incomplete
florets absent.

Female-fertile florets 1 . Lemmas papery; 4-5 nerved;
incised; awned. Awns 1; median; dorsal; non-geniculate.
Palea present; relatively long. Lodicules 2; membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit ellipsoid; hilum
short (linear).

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. 1 species.
Mediterranean to Pakistan. Mesophytic, or xerophytic; in
open habitats (in dry places). Cape Province. 1 naturalized
species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Linder.
Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

Fig. 117. Lamarckia aurea

196

Lamarckia aurea (L.) Moench.

Fig. 117. PI. 104.

( =Cynosurus aureus L.) 1.

Annual; loosely tufted (culms
geniculate or erect); 100-200
(-300) mm tall. Leaf blades 30-
90 mm long; 3-8 mm wide.

Spikelets (sterile ones) 6-9 mm
long; to 0.8 mm wide. Panicle
20-80 mm long, 10-25 mm wide,
soft, silky; spikelets of two kinds, in fascicles of 4-5,
terminal spikelet of each fascicle female-fertile, others
sterile; female-fertile spikelet 1 -flowered, to 2 mm long,
pedicels 2-3 mm long, villous; lemma awn more than twice
the length of the body; sterile spikelets consisting of two
glumes and numerous imbricate, obtuse, awnless, empty
lemmas.

Flowering August to October. Usually on road verges
near tarmac in stony gravel or loam. Rare. Locally common.
Naturalized from the Mediterranean basin. Biome: Fynbos.
Mediterranean and Middle East, cultivated in USA. Orna-
mental and weed (insignificant).

Description: Linder (33), Stapf 1898-1900 (689), Hitch-
cock & Chase 1950 (187), Chippindall 1955 (61).
Illustration: Chippindall 1955 (fig. 33), Hitchcock & Chase
1950 (fig. 369). Voucher: Crook 2195. PRECIS code
9903720-00100.

Leersia Swartz.

Aplexia Faf., Asprella Schreb., Blepharochloa Endl.,
Ehrhartia Weber, Endodia Raf., Homalocenchrus Mieg,
Laertia Gromov, Pseudoi'yza Griff., Turraya Wall.

Perennial; long-rhizomatous, or long-stoloniferous, or
caespitose. Culms 300-1500 mm high; herbaceous. Leaf
blades linear; flat, or folded, or rolled. Ligule an unfringed
membrane. The spikelets all alike in sexuality.

Inflorescence paniculate ; open\ espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary; 3-6 mm long; compressed laterally
(strongly so); disarticulating above the glumes (or at least,
above the rim assumed to represent them). Rachilla
terminated by a female-fertile floret. Glumes absent
(apparently reduced to a narrow rim at the tip of the
pedicel ). Proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas awn less, or mucronate
(often caudate); 3-5 nerved. Palea present; relatively long
(but much narrower than the lemma); with several nerves
(3). Lodicules 2; fleshy, or membranous; glabrous. Stamens
1-6. Ovary glabrous. Fruit small; hilum long-linear;
embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells, or without arm cells; without fusoids. Midrib with one
bundle only, or vascularization complex (rarely).

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Oryzodae; Oryzeae. 18
species. Tropical and warm temperate. Helophytic; in shade
and in open habitats. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 4 indigenous species.

References. 1. Clayton. 1970. FTEA. 2. Launert. 1971.
FZ 10(1).

Species treatment by G.E. Gibbs Russell.

1(0). Spikelets more than 1 mm across; nodes hairy; plants
not restricted to the Okavango and Caprivi .... 2

Spikelets less than 1 mm across; nodes glabrous or
hairy; plants of the Okavango swamp ........ 3

2(1). Keels of lemma and palea ciliate with stiff hairs

0.2-0. 6 mm long; plants widespread

L. hexandra

Keels of lemma and palea shortly ciliolate, with hairs

less than 0.2 mm long; western Transvaal

L. denudata

3(1). Nodes glabrous; plants robust; culms spongy, 3-5 mm

across L. friesii

Nodes hairy; plants delicate; culms not spongy, 1 mm
across L. tisserantii

197

Leersia denudata Launert

Slender perennial; hydrophyte
and tufted; to 700 mm tall. Leaf
blades 30-140 mm long; 1. 5-6.0
mm wide (flat or rolled, nearly
smooth). Spikelets 3.5 — 4.5 mm
long; 1.3-1. 6 mm wide. Culm
nodes velvety; lemma and palea
with fine cilia less than 0.2 mm
long.

Flowering February. Swampy grassland, vleis, deep
parts of temporary pans. Rare. North to tropical east Africa.

Description: Clayton et al. 1970-1982 (27). Voucher:
Kinges 1620. PRECIS code 9901590-00100.

Leersia friesii Meld.

Perennial; hydrophyte and
rhizomatous (rhizome creeping);

600-700 mm tall. Leaf blades
80-200 mm long; to 6 mm wide
(flat, smooth). Spikelets 3. 0-3. 5
mm long; 0. 9-1.0 mm wide.

Culms spongy, nodes glabrous;
lemmas sometimes subcaudate.

Flowering November to May.

Swamps. Rare. Central and eastern tropical Africa.

Description: Clayton et al. 1970-1982 (27). Voucher:
P.A. Smith 1806. PRECIS code 9901590-00150.

Leersia hexandra Swartz

Fig. 118. PI. 105.

Perennial; hydrophyte and
rhizomatous (rhizome creeping);

300-1000 mm tall. Leaf blades
100-200 mm long; 4-8 mm wide
(flat, strongly scabrous). Spike-
lets 3.4 — 4.8 mm long; 1.2-1. 4
mm wide. Culm nodes hairy;
lemma and palea keels with stiff
cilia 0. 2-0.6 mm long.

Flowering July to June. Floodplains and permanently
wet places such as vleis, pans and ditches, often forming
extensive colonies. Locally common. Throughout tropics.
The leaves and culms make a characteristic rattling sound
when shaken together.

Description: Chippindall 1955 (33), Clayton et al.
1970-1982 (27). Illustration: Chippindall 1955 (pi. 2).
Voucher: V.d. Schijff 21 15. PRECIS code 9901590-00200.

Leersia tisserantii (A. Chev.) Launert

Perennial, or annual; hydro-
phyte, or rhizomatous (some-
times), or tufted (loosely to
densely); 150-600 mm tall. Leaf
blades 40-180 mm long; 1-2 mm
wide (flat or rolled, strongly
scabrous). Spikelets 4-5 mm long
(excluding caudae); 0. 8-1.0
(-1.1) mm wide. Culm nodes
usually hairy; lemmas with a short flat cauda.

Flowering January to April. In deep water at river edges,
growing in dense colonies. Rare. Central and eastern
tropical Africa. Plants in our area have comparatively short
caudae on the lemmas, but in tropical Africa these can be
up to 7.5 mm long.

Description: Clayton et al. 1970-1982 (25), Launert.
1965. Senck. Biol. 46:129. Voucher: Gibbs Russell 2840.
PRECIS code 9901590-00300.

Leptocarydion Stapf

Annual; loosely caespitose, or decumbent (rarely,
rooting at the nodes). Culms 130-1300 mm high; herba-
ceous; branched above, or unbranched above. Leaf blades
lanceolate to ovate\ broad\ flat, or rolled. Ligule a fringed
membrane.

198

Inflorescence of spike-like main branches', contracted (to
about 200 mm long, the thin spicate laterals appressed);
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; 5-1 1 mm long; compressed
laterally; disarticulating above the glumes; disarticulating
between the florets. Glumes two; relatively large; very
unequal; markedly shorter than the spikelets; awnless;
similar (reddish, subhyaline, very narrow). Incomplete
florets distal to the female-fertile florets, merely underde-
veloped, awned; proximal incomplete florets absent.

Female-fertile florets 6-12. Lemmas similar in texture
to the glumes to decidedly firmer than the glumes (thin);
without a germination flap; 3 nerved; incised; awned. Awns
1; median (the midnerve excurrent); from the sinus; non-
geniculate (very slender); about as long as the body of the
lemma to much longer than the body of the lemma. Palea
present (linear-oblong); relatively long, or conspicuous but
relatively short. Lodicules 2; fleshy; glabrous. Stamens 2,
or 3. Ovary glabrous. Fruit small (to 1 mm); linear; hilum
short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. East and southern Africa.
Mesophytic. Namibia, Botswana, Transvaal, Swaziland,
and Natal. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Leptocarydion vulpiastrum (De Not.) Stapf

Fig. 1 19. PI. 106.

Spade grass.

Annual; loosely or compactly
tufted; 400-1070 mm tall. Leaf
blades 20-120 mm long; 6-20
mm wide. Spikelets 5-1 1 mm
long. Leaf blades papery,
lanceolate-oblong, rounded at
base; panicle a narrow silky
plume, 50-150 mm long.

Flowering February to June. Usually on sandy soil in
mopane veld, riverine woodland or rocky hillsides, often in
the shade. Infrequent to locally common. Biome: Savanna.
Tropical Africa. Easily controlled weed, pasture, and orna-
mental (in grass gardens).

Description: Chippindall & Crook 1976 (177), Stapf
1898-1900 (648), Chippindall 1955 (127), Clayton et al.
1970-1982 (294). Voucher: De Winter 2905. PRECIS code
9903430-00100.

Leptochloa P. Beauv.

Anoplia Steud., Baldomiria Herter, Diachroa Nutt,
Diacisperma Kuntze, Disakisperma Steud., Ipnum Phil.,
Leptostachys Meyer, Oxydenia Nutt, Rabdochloa P. Beauv.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms woody and
persistent, or herbaceous; branched above, or unbranched
above. Leaf blades linear-lanceolate ; flat, or rolled. Ligule
an unfringed membrane to a fringe of hairs. The spikelets
all alike in sexuality.

Inflorescence of spike-like main branches (spiciform
racemes)', open; digitate or subdigitate, or non-digitate (the
racemes often whorled, rarely subdigitate); espatheate.
Spikelet-bearing axes with very slender rachides',
persistent.

Spikelets solitary; not in distinct ‘long-and-short’ com-
binations; 1-5 mm long ( rarely up to 7); compressed
laterally to not noticeably compressed; disarticulating
above the glumes; disarticulating between the florets.

Callus short', blunt. Glumes two; very unequal, or more or
less equal; markedly shorter than the spikelets; awnless. All
florets female-fertile, or distal incomplete florets also
present, these awnless; proximal incomplete florets absent.

Female-fertile florets 2-6 (usually 3-6, rarely I).
Lemmas less firm than the glumes to similar in texture to
the glumes (membranous to hyaline); 3 nerved; entire
(rarely), or incised; awnless, or mucronate, or awned. Awns
when present 1; from the sinus; non-geniculate; much
shorter than the body of the lemma. Palea present. Lodi-
cules 2; fleshy. Stamens 2-3. Ovary glabrous. Fruit small
(0.5-2 mm); hilum short; pericarp free, or loosely adherent,
or fused; embryo large.

Photosynthetic pathway and related features. C4; PCK
(L. ciliolata), or NAD-ME (L. digitata ); XyMS+. PCR
sheath outlines uneven, or even. PCR sheath extensions
present, or absent. Maximum number of extension cells
when present 1. PCR cell chloroplasts ovoid, or elongated;

Fig. 120. Leptochloa panicea

199

with well developed grana; centrifugal/peripheral ( L .
ciliolata), or centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 27
species. Tropical and subtropical. Helophytic, mesophytic,
and xerophytic; in shade and in open habitats (woodland,
savanna, dry and swampy soils); halophytic and
glycophytic. Namibia, Botswana, Transvaal, Swaziland,
and Natal. 3 indigenous species.

References. 1. Clayton et al. 1974. FTEA. 2. Phillips.
1982. Kew Bull. 37:133.

Species treatment by M. Koekemoer.

1(0). Leaf blades lanceolate-oblong to oblong, 40-120 mm
long, 6-18 mm wide; spikelets 1 -flowered; glumes

longer than the lemmas L. uniflora

Leaf blades linear; 100-500 mm long, 3-8 mm wide;
spikelets 2-6-flowered; glumes shorter than the

lemmas 2

2(1). Plants perennial, aquatic or semi-aquatic; leaf sheaths
white and glabrous; glumes unequal; lemmas with
short dense hairs on the nerves; caryopses elliptic-

oblong, longer than 0.5 mm L. chinensis

Plants annual, in bush or grassland; leaf sheaths and
blades green and papillate-pilose; glumes subequal;
lemmas with short hairs on the back; caryopses

broadly elliptic, less than 0.5 mm long

L. panicea

Leptochloa chinensis (L.) Nees

Perennial; hydrophyte, stolon-
iferous, and tufted; 440-820 mm
tall. Leaf blades 200-500 mm
long; 3. 0-7. 5 mm wide. Spikelets
2. 1-3.2 mm long. Leaf sheaths
white, papery and glabrous; pani-
cle 200-600 mm long; spikelets
2-6-flowered; glumes unequal,
shorter than the lemmas.

Flowering December to April. In or by water. Infrequent
to locally common. Biome: Savanna. Southern tropical
Africa through India to Japan and in Indonesia. Weed
(ricefields).

Description: Clayton et al. 1970-1982 (279). Voucher:
De Winter & Codd 315. PRECIS code 9903330-00100.

Leptochloa panicea (Retz.) Ohwi

Fig. 120. PI. 107.

Annual; tufted; 320-1200 mm
tall. Leaf blades to 250 mm long;
about 7 mm wide. Spikelets
1.9-2. 5 mm long. Leaf sheaths
and blades papillate pilose, blades
linear; panicle 200-300 mm long;
spikelets (2-)3(-5)-flowered.

Flowering January to May.

Clayey loam, in or near water.

Rare. Biome: Savanna. Tropical Africa and tropical Asia.

Description: Chippindall 1955 (121), Clayton et al.
1970-1982 (279). Illustration: Chippindall 1955 (fig. 93),
Clayton et al. 1970-1982 (fig. 76). Voucher: Acocks 16789.
PRECIS code 9903330-00200.

Leptochloa uniflora A. Rich.

(-Craspedorhachis uniflora
(Hochst. ex A. Rich.)

Chippind.) 1.

Slender annual; tufted; 300-
610 mm tall. Leaf blades 40-120
mm long; 6-18 mm wide. Spike-
lets 1 .9-2.8 mm long. Leaf blades
lanceolate-oblong to oblong;

panicle 1 50 — 450 mm long; spikelets 1 -flowered.

Flowering January to May. In bushveld under trees.
Rare. Locally common. Biome: Savanna. Tropical Africa,
India and Ceylon.

Description: Chippindall 1955 (205), Clayton et al.
1970-1982 (276). Illustration: Chippindall 1955 (fig. 182).
Voucher: Killick & Leistner 3347. PRECIS code

9903330-00300.

Lepturus R.Br.

Lepiurus Dum., Leptocercus Raf., Monerma P. Beauv.

Perennial ; long-stoloniferous and caespitose. Culms
100-600 mm high; herbaceous; branched above, or un-
branched above. Leaf blades linear to linear-lanceolate; flat,
or rolled (involute). Ligule an unfringed membrane .

Inflorescence a single spike ( almost cylindrical, the
joints striate ); espatheate. Spikelet-bearing axes disarticu-
lating; disarticulating at the joints.

Spikelets solitary; distichous; 3-15 mm long (-20 mm);
compressed dor six entr ally ; falling with the glumes. Glumes
one per spikelet (G 1 usually missing), or two; very unequal;
long relative to the adjacent lemmas (i.e., G2); awnless, or
awned (G2 sometimes tapered into a short awn); very dis-
similar (lower reduced to a minute triangular scale; upper
exceeding the spikelet, thickened, with rows of minute
bristles on the back). Incomplete florets distal to the female-
fertile florets; proximal incomplete florets absent.

Fig. 121. Lepturus repens

200

Female-fertile florets 1-2. Lemmas less firm than the
glumes (membranous); 3 nerved; entire; awnless. Palea
present (lanceolate); relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit ellipsoid; hilum
short; pericarp free; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven, or even. PCR sheath
extensions absent.

Cytology, classification, distribution. Chloridoideae;
Chlorideaeserzsw lato. About 8 species. Coastal east Africa,
Madagascar, to Australia, & Polynesia. Xerophytic; in open
habitats (sandy beaches and coastal hinterland); maritime-
arenicolous (usually, a good sandbinder). Natal. 1 indige-
nous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by G.E. Gibbs Russell.

Lepturus repens (G. Forst.) R. Br.

Fig. 121. PI. 108.

Perennial; stoloniferous; 1 00—

600 mm tall. Leaf blades to 150
mm long; 2-10 mm wide. Spike-
lets (8—) 1 0— 1 4(— 22) mm long. In-
florescence a fragile spike with
the spikelets sunk on opposite
sides of the corky axis; lower
glume absent, upper glume
awned; florets 1 or 2.

Flowering September to October. Sand dunes, in salt
spray zone. Infrequent. Shores of the Indian Ocean east to
Polynesia. Erosion control (sand binder). This species is
strikingly similar to Hainardia cylindrica , which is an
annual. The inflorescence is superficially similar to
Hemarthria and allied andropogonoids, in which the spike-
lets are paired.

Description: Clayton et al. 1970-1982 (391).

Illustration: Clayton et al. 1970-1982 (fig. 104). Voucher:
Venter 6274. PRECIS code 9904420-00100.

Leucophrys Rendle

Sometimes included in Brachiaria.

Perennial. Culms 70-1000 mm high; woody and
persistent (stiffly geniculate)', branched above (plants
bushy). Leaf blades linear-lanceolate; flat, or rolled. Ligule
a fringe of hairs.

Inflorescence paniculate; contracted; espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs; with pedicellate spikelets
(the spikelets substipitate, with a short stalk fitting into the
pedicel apex); consistently in Tong-and-short’ combina-
tions, or not in distinct ‘long-and-short’ combinations.
Spikelets 4-6 mm long; abaxial to adaxial (the orientation
variable); compressed dorsiventrally; falling with the
glumes. Glumes two; relatively large; very unequal, or more
or less equal; with distinct rows of hairs (the upper with a
transverse row above the middle, the lower glabrous save
at the base)', awnless (but the tips caudate, inrolled,
membranous); very dissimilar (lower obtuse or notched at
apex, pilose at base, the upper tapering, caudate, dorsally
long-villous with a transverse fringe just above the middle).
Proximal incomplete florets 1; paleate, palea fully
developed; male.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes; smooth; becoming indurated (glossy); hairless;
having the margins tucked in onto the palea; with a clear
germination flap (basal); 5 nerved; entire; awnless. Palea
present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous.

Photosynthetic pathway. C4; XyMS+. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 1 species. Tropical and southern Africa.
Helophytic, or xerophytic; in open habitats (sandy riverbeds
in semidesert); glycophytic. Namibia and Cape Province. 1
indigenous species.

References. 1 . Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA.

Species treatment by H.M. Anderson.
Leucophrys mesocoma (Nees) Rendle
Withaargras.

Perennial; tufted; to 1000 mm
tall. Leaf blades 30-120 mm
long; 3-8 mm wide. Spikelets 7
mm long; 2-3 mm wide. Culms
woody, blue-green, profusely
branched, geniculate, nodes swol-
len; panicle 130 mm long, 30

Fig. 122. PI. 109.

Fig. 122. Leucophrys mesocoma

201

mm wide; upper glume and lemma of lower floret with
dense hairs 3-5 mm long in two tufts or a continous fringe
halfway up.

Flowering February to May. Sandy riverbeds. Infre-
quent. Biome: Nama-Karoo. Angola. Natural pasture.
Species of Melinis have a similar inflorescence. This
species may be distinguished by its much branched habit.

Description: Chippindall 1955 (379). Illustration:
Muller 1984 (186), Chippindall 1955 (fig. 325). Voucher:
Smook 5261. PRECIS code 9901030-00100.

Lintonia Stapf

Joannegria Chiov., Negria Chiov.

Perennial; caespitose, or long-rhizomatous and caespi-
tose. Culms 200-900 mm high; herbaceous; branched
above. Leaf blades linear (tapered to a fine, acuminate tip);
flat. Ligule an unfringed membrane (minutely ciliolate, with
long hairs at the auricle positions).

Inflorescence of spike-like main branches ; open; digitate
or subdigitate, or non-digitate (L. brizoides ); espatheate
(but often enveloped below by the sheath of the uppermost
culm leaf). The racemes without spikelets towards the base.
Spikelet-bearing axes persistent.

Spikelets solitary; biseriate, or not two-ranked; 4-1 1 mm
long (cuneate or elliptic, plump); compressed laterally; dis-

articulating above the glumes; not disarticulating between
the florets ( the rachilla tough). Glumes two; very unequal
(G 1 shorter); markedly shorter than the spikelets; awnless
(sub-mucronate); similar (persistent, hyaline-membranous).
Incomplete florets distal to the female-fertile florets, merely
underdeveloped, shortly awned; proximal incomplete
florets absent.

Female-fertile florets 2-4 (with several sterile ones
above). Lemmas decidedly firmer than the glumes (tough
and cartilaginous, at least in part); without a germination
flap; 5-9 nerved; incised (shortly so); awned. Awns 1;
median; dorsal; non-geniculate (curved); much shorter than
the body of the lemma to about as long as the body of the
lemma. Palea present; relatively long (about 3/4 the length
of the lemma). Lodicules 2; fleshy (cylindrical rather than
cuneate); glabrous. Stamens 3. Ovary glabrous. Fruit small
(1.3-2. 2 mm); ellipsoid; hilum short (elliptical); pericarp
free; embryo large (about half the length of the fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath
extensions absent. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Tropical east Africa.
Helophytic, or mesophytic; in shade, or in open habitats
(savanna, heavy soils in seasonally wet places);
glycophytic. Botswana, Transvaal, Swaziland, and Natal. 1
indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Lintonia nutans Stapf

Perennial; shortly stolonifer-
ous and tufted; 400-900 mm tall.

Leaf blades 30-150 mm long; 3-5
mm wide. Spikelets 6-10 mm
long; 4-8 mm wide. Leaf blades
usually glabrous; inflorescence of
2-4 digitate or subdigitate
racemes; spikelets 4-10-
flowered, wedge-shaped; lemma
shortly bilobed, with an awn 1-11 mm long between the
lobes.

Flowering December to March. On black clayey soil in
vleis or along pan edges. Conservation status not known.
Infrequent. Biome: Savanna. East Africa.

Description: Chippindall 1955 (117), Clayton et al.
1970-1982 (302). Voucher: Du Toit 169. PRECIS code
9902020-00100.

Lolium L.

Arthrochortus Lowe, Craepalia Schrank, Crypturus
Link.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 100-1300 mm
high; herbaceous; unbranched above. Leaves auriculate.
Sheath margins free. Leaf blades linear; flat, or folded, or
rolled. Ligule an unfringed membrane.

• Inflorescence a single spike ; espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary; conspicuously distichous; 7-26 mm
long; compressed laterally; disarticulating above the
glumes. Glumes one per spikelet (except that the terminal
spikelet has two)', decidedly shorter than the adjacent
lemmas, or long relative to the adjacent lemmas; awnless.
Incomplete florets distal to the female-fertile florets, merely
underdeveloped; proximal incomplete florets absent.

Female-fertile florets 2-22. Lemmas less firm than the
glumes to decidedly firmer than the glumes (membranous

202

to papery, sometimes turgid or hardening in fruit); 5-7
nerved; entire, or incised; awnless, or awned. Awns when
present 1; from the sinus, or dorsal; non-geniculate; much
shorter than the body of the lemma. Palea present; relatively
long (usually ciliate). Lodicules 2; membranous; glabrous.
Stamens 3. Ovary glabrous. Fruit small, or medium sized,
or large; hilum long-linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 7. Pooideae; Poodae; Poeae. 8 species.
Temperate Eurasia, north Africa. Mesophytic; in open
habitats. Namibia, Transvaal, Orange Free State, Natal,
Lesotho, and Cape Province. 3 naturalized species.

Intergeneric hybrids with Festuca — X Festulolium
Aschers. & Graebn. (several species of each genus
involved).

References. 1. Chippindall. 1955. Gr. & Past. 2.
Humphries. 1980. FI. Europ. 3. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Glumes appressed to the rachis, slightly gaping at
maturity, concealing or partly concealing the
spikelets in the concavities of the rachis; spikelets
1-3 mm wide L. rigidum

Glumes ascending or spreading, not concealing the

spikelets; spikelets 3-1 1 mm wide 2

2(1). Lemmas elliptical to ovate, very turgid at maturity
especially towards the base; mature caryopses to 3

times as long as wide L. temulentum

Lemmas oblong to oblong-lanceolate, not turgid at
maturity; mature caryopses more than 3 times as

long as wide 3

3(2). Lemmas awned, awn up to 15 mm long; sterile leafy
shoots absent; leaves convolute when young ....

. L. multiflorum / L. multiflorum x L. perenne
Lemmas awnless; plants with sterile leafy shoots;

leaves flat or folded when young 4

4(3). Plants perennial; leaf blades 2-4 mm long

L. perenne

Plants annual or biennial; leaf blades to 10 mm wide
L. multiflorum x L. perenne

Lolium multiflorum Lam.

PI. 111.

Italian rye grass.

Shortlived perennial, or an-
nual; loosely tufted; 200-800
(-1300) mm tall. Leaf blades
1 10-220 mm long; 3-8 mm wide.

Spikelets 8-20 mm long; 2-10
mm wide. Glumes ascending or
spreading, not concealing the
spikelets, 1 /4 — 1 /2(— 3/4) the spikelet length; lemmas acute,
not turgid at maturity, awn to 1 5 mm long; mature caryopsis
more than three times as long as wide.

Flowering October to April. On roadsides and other
disturbed areas. Common. Naturalized from Europe and the
Mediterranean area. Biome: Fynbos, Savanna, Grassland,
and Nama-Karoo. Introduced worldwide. Irrigated annual
winter pasture, or poisonous (when infected by fungi), or
weed (an escape from cultivation). Subjectively separated
from L. multiflorum x L. perenne by numerous overlapping
characters.

Description: Humphries 1980 (5: 154), Stapf 1898-1900
(739), Hitchcock & Chase 1950 (272), Chippindall 1955
(59). Illustration: Chippindall 1955 (fig. 29), Hitchcock &
Chase 1950 (fig. 534). Voucher: Smook 1705. PRECIS
code 9904330-00200.

Lolium multiflorum x L. perenne

Shortlived perennial, or an-
nual; loosely tufted; 150-1000
mm tall. Leaf blades to 300 mm
long; to 10 mm wide. Spikelets
6-25 mm long; 3-10 mm wide.

Glumes ascending or spreading,
not concealing the spikelets, 1/4
to as long as the spikelets;
lemmas not turgid at maturity,
more or less acute, awns absent or up to 10 mm long; mature
caryopsis more than three times as long as wide.

Flowering October to December. In cultivated or fallow
lands, on roadsides and in moist disturbed places. Locally
common. Naturalized from Europe. Biome: Fynbos and
Grassland. Europe, widely cultivated. Cultivated pasture
and weed (an escape from cultivation). In Europe, L.
multiflorum and L. perenne are two quite distinct species,
but they hybridize freely to form a complete range of
intermediates in southern Africa. This hybrid is subjectively
separated from L. multiflorum by numerous overlapping
characters.

Description: Linder (21), Humphries 1980 (5:154).
Voucher: Theron 927. PRECIS code 9904330-00250.

203

Lolium perenne L.

Perennial rye grass.

Perennial; tufted (with numer-
ous culms and sterile leafy
shoots); 250-500(-900) mm tall.

Leaf blades 50-140(-300) mm
long; 2-4 mm wide. Spikelets
10-15 mm long; 3-10 mm wide.

Glumes ascending or spreading,
not concealing the spikelets, 1/2-3/4 the spikelet length;
lemmas acute, not turgid at maturity, awnless; mature
caryopsis more than three times as long as wide.

Flowering March, April, November, and December. On
roadsides, in moist disturbed areas and cultivated and
fallow lands. Locally common. Naturalized from Europe.
Biome; Fynbos, Savanna, Grassland, and Nama-Karoo.
Introduced worldwide. Irrigated perennial winter pasture,
or poisonous (when infected by fungi), or weed (an escape
from cultivation). Hybridizes freely with other Lolium
species, as well as with species of Festuca.

Description: Humphries 1980 (5:154), Hitchcock &
Chase 1950 (269), Chippindall 1955 (58). Voucher:
Burrows 2216. PRECIS code 9904330-00300.

Lolium rigidum Gaudin

(=L. loliaceum (Bory &

Chaup.) Hand.-Mazz) 3.

Annual; tufted (with numer-
ous flowering culms or solitary
culms in young plants; culms
branched near the base and some-
times rooting from the lower
nodes); 100-300(-500) mm tall.

Leaf blades 100-170 mm long; 5-8 mm wide. Spikelets
7-20 mm long; 1-3 mm wide. Spikelets 2-flowered; glumes
appressed to the rachis, slightly gaping at maturity,
concealing or partly concealing the spikelets in the
concavities of the rachis, 3/4 to slightly longer than the
spikelets; lemmas obtuse to acute, not turgid at maturity,
awn absent or to 10 mm long; mature caryopsis more than
three times as long as wide.

Flowering September to January. In disturbed and
weedy places on sandy to clayey soils, preferably where it
is moist, occasionally in water and on stream edges. Locally
common. Naturalized from the Mediterranean. Biome: Fyn-
bos, Savanna, Grassland, Nama-Karoo, and Desert.
Introduced worldwide in temperate regions. Weed. Similar
in size and habit to Hainardia cylindrica, which has one-
flowered spikelets.

Description: Humphries 1980 (5: 154), Stapf 1898-1900
(740), Chippindall 1955 (59). Voucher: Smook 3649.
PRECIS code 9904330-00350.

Lolium temulentum L.

Fig. 124.

Drabok, darnel.

Robust annual; culms solitary
or tufted; 400-900 mm tall. Leaf
blades 150-300 mm long; 3-7
mm wide. Spikelets 8-28 mm
long; 3-8 mm wide. Glumes
ascending or spreading, not
concealing the spikelets, 3/4 to
1 1/2 times the spikelet length; lemmas elliptical to ovate,
very turgid at maturity, awn absent or to 20 mm long;
mature caryopsis 2-3 times as long as wide.

Flowering September to February. Usually in cultivated
or fallow lands, gardens or other disturbed areas, often
associated with wheat. Locally common. Naturalized from
the Mediterranean. Biome: Fynbos, Savanna, Grassland,
and Nama-Karoo. Naturalized in most warm temperate

countries. Weed. The fungus associated with this grass
contains an alkaloid poisonous to livestock.

Description: Humphries 1980 (5: 154), Stapf 1898-1900
(738), Hitchcock & Chase 1950 (272), Chippindall 1955
(59). Illustration: Chippindall 1955 (fig. 30), Hitchcock &
Chase 1950 (fig. 535). Voucher: Lategan s.n. PRECIS code
9904330-00400.

Lophachme Stapf

Perennial; caespitose. Culms 130-570 mm high; herba-
ceous; unbranched above (though branching below). Leaf
blades linear-lanceolate\ rolled (involute). Ligule an
unfringed membrane (in L. parva ), or a fringed membrane
(in L. digitata).

Inflorescence of spike -like main branches (a panicle of
slender spike-like racemes ); digitate or subdigitate ; espath-
eate, or spatheate (in that the spikelets sometimes subtended
by very minute, sparsely hairy, hyaline scales — vestigial
bracts or spikelets?). Spikelet-bearing axes with very
slender rachides\ persistent.

Spikelets solitary (somewhat distant); biseriate; 3.5-6
mm long; compressed laterally; disarticulating above the
glumes; disarticulating between the florets; with distinctly
elongated rachilla internodes between the florets (between
L 1 and L2 and above L2). Callus pointed. Glumes two; very
unequal (G1 shorter), or more or less equal; markedly
shorter than the spikelets to about equalling the spikelets;

Fig. 125. Lophachme digitata

204

awnless; similar (linear-lanceolate, membranous). Incom-
plete florets distal to the female-fertile florets, about 4,
clearly specialised and modified in form (reduced to awns,
forming a tuft which remains attached to the upper fertile
floret), awned; proximal incomplete florets absent.

Female-fertile florets 1, or 2 (the second floret
sometimes male-only). Lemmas less firm than the glumes
(thinly membranous); without a germination flap; 3 nerved;
incised; awned. Awns 1; median; from the sinus; non-genic-
ulate (fine, straight or recurved); about as long as the body
of the lemma to much longer than the body of the lemma.
Palea present; relatively long (reaching the bases of the
lemma lobes). Lodicules somewhat fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small (1.8 mm long);
fusiform; hilum short; pericarp loosely adherent (easily
removable after soaking); embryo large (a little more than
1/3 the length of the fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Southern tropical and
southern Africa. Helophytic, or mesophytic (open grassland
or streamsides); in shade (L. parva), or in open habitats (L.
digitata ); glycophytic. Transvaal and Natal. 1 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

Lophachme digitata Stapf

Fig. 125. PI. 1 12.

Slender perennial; rhizomat-
ous and tufted; 170-570 mm tall.

Leaf blades 30-45 mm long;

1.0-2. 5 mm wide. Spikelets 5-6
mm long. Rhizome long and
slender; basal sheaths fibrous;
spikes 2-8, to 80 mm long;
central nerve of lemma extending
to an awn longer than the floret.

Flowering February to April. Open highveld sourveld.
Infrequent to locally common. Biome: Savanna and Grass-
land. Endemic.

Description: Stapf 1898-1900 (647), Chippindall 1955
(127). Illustration: Hooker’s Icon. PI. (pi. 261 1), Chippin-
dall 1955 (fig. 101). Voucher: Codd 974. PRECIS code
9903520-00100.

Lophochloa Reichenb.

Acrospelion Schult., Parvotrisetum Chrtek, Rupestrina
Prov., Sennenia Sennen, Trisetaria Forssk., Trisetarium
Poir. , sometimes includes Rostraria Trin.

Annual ( Trisetaria ), or perennial; long-rhizomatous, or
caespitose. Culms 40-1500 mm high; herbaceous. Leaf
blades linear ; flat, or rolled (convolute). Ligule an
unfringed membrane ( sometimes puberulent or ciliolate).
The spikelets all alike in sexuality.

Inflorescence paniculate ; open, or contracted (loose, or
if dense then interrupted, neither cylindrical nor ovoid);
espatheate. Spikelet-bearing axes persistent.

Spikelets not secund\ 2.4-9 mm long', compressed
laterally; disarticulating above the glumes, or falling with
the glumes, or not disarticulating. Hairy callus present
(usually). Glumes two; very unequal, or more or less equal;
markedly shorter than the spikelets, or about equalling the
spikelets; awnless; similar. All florets female-fertile, or
distal incomplete florets also present; proximal incomplete
florets absent.

Female-fertile florets 1 (rarely), or 2-5, or 6-12 (rarely).
Lemmas similar in texture to the glumes ; carinate ; 3-7

nerved; incised', awned (usually conspicuously, contrast
Koeleria, rarely awnless). Awns when present 1, or 3;
median, or median and lateral (via setae from the lobes).
The median awn different in form from the laterals (when
laterals present); usually dorsal (or ‘subterminal’); usually
twisted; non-geniculate, or geniculate; much shorter than
the body of the lemma, to much longer than the body of the
lemma (conspicuous if inflorescence compact). Palea
present; relatively long. Lodicules 2; membranous; ciliate,
or glabrous. Stamens 3. Ovary usually glabrous. Fruit small,
or medium sized; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 6 and 7. Pooideae: Poodae; Aveneae. About
85 species. North & south temperate. Mesophytic, or xero-
phytic; mostly in open habitats (meadows, mountain slopes,
upland grasslands, weedy places); glycophytic. Namibia
and Cape Province. 2 naturalized species.

Integeneric hybrids with Koeleria (X Trisetokoeleria
Tsvelev), Sphenopholis.

Fig. 126. Lophochloa pumila

205

References. 1. Jonsell. 1980. FI. Europ. 5. 2. PRE
Herbarium practice, following Smook & Gibbs Russell.

Species treatment by T.M. Sokutu.

1(0). Spikelets 3^4-flowered; upper glume densely

pubescent, lower one less so L. pumila

Spikelets 3-6-flowered; glumes glabrous to pubescent
or lower glume pubescent and upper glume
glabrous L. cristata

Lophochloa cristata (L.) Hyl.

(=Koeleria phleoides (Vill.)

Pers.) 1.

Annual; loosely tufted; 50-
400 mm tall. Leaf blades 40—120
mm long; to 2.5 mm wide. Spike-
lets 3-5 mm long. Inflorescence
spiciform; spikelets 3-6-flower-
ed; glumes glabrous to pub-
escent, or lower glume pubescent and upper glume
glabrous; lemma awn 1-3 mm long.

Flowering October to December. Dry exposed areas or
sometimes also in moist or rocky areas. Infrequent.
Naturalized from Europe and the Mediterranean area.
Biome: Fynbos. North Africa and Europe to India. Weed,
Can be confused with L. pumila , especially when the
lowermost lemma is pubescent and the rest of the spikelet
is less hairy or glabrous.

Description: Adams. & Salt. 1950 (84), Jonsell 1980
(5:220), Stapf 1898-1900 (470), Chippindall 1955 (84).
Voucher: Cleghorn 3144. PRECIS code 9903741-00100.

Spikelets solitary, or in pairs; consistently in ‘long-and-
short’ combinations, or not in distinct ‘long-and-short’
combinations. Spikelets 6-25 mm long; compressed
laterally to not noticeably compressed; disarticulating
above the glumes. Glumes two; relatively large; very
unequal; awned, or awnless (G2 may be setaceous-
acuminate); similar. Lower glume 3 nerved ; shorter than the
female-fertile lemma. Proximal incomplete florets 1\
paleate, or epaleate (L. togoensis), palea when present fully
developed (membranous, two keeled); male, or sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes; not becoming
indurated (more or less leathery); hairy to hairless (pilose
to glabrescent, the hairs not in tufts; not in transverse rows);
the margins tucked in onto the palea; without a germination
flap; 5-9 nerved; incised (usually shortly so, rarely entire);
awned. Awns 1; median; from the sinus; geniculate; much
longer than the body of the lemma. Palea present (linear);
relatively long. Lodicules 2; fleshy; glabrous. Stamens 2
(rarely 3). Ovary glabrous. Stigmas brown. Hilum long-
linear; embryo large.

Photosynthetic pathway. C4. Organization of PCR tissue
when unconventional Arundinella type. XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x - 6 and 12. Panicoideae; Panicodae; Arun-
dinelleae. About 26 species. Tropical and southern Africa,
Madagascar, with 1 in South America. Helophytic, or meso-
phytic, or xerophytic; in open habitats (in savanna
woodland, often on poor shallow soils); glycophytic.
Namibia, Transvaal, Orange Free State, Swaziland, Natal,
and Cape Province. 6 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1972. FTEA.

Species treatment by H.M. Anderson.

Lophochloa pumila (Desf.) Bor

Fig. 126. PI. 113.

(=Trisetaria pumila (Desf.)

Maire) 2; (-Trisetum pumilum
(Desf.) Kunth) 1.

Annual; tufted; 45-400 mm
tall. Leaf blades 35-65 mm long;
to 2 mm wide. Spikelets 2. 5-4.0
mm long. Inflorescence spici-
form; spikelets 2-4-flowered;
lower glume glabrous to puberulous, ciliate on the keel;
upper glume densely pubescent; lemma awn 1 .5 — 4.0 mm
long.

Flowering September to January. Dry and/or rocky
areas, sometimes beneath bushes. Infrequent. Naturalized
from Europe. Biome: Fynbos and Succulent Karoo. Spain.
Weed. Can easily be confused with L. cristata, which has
less hairy glumes and sometimes a shorter lemma awn.

Description: Jonsell 1980 (5:220), Stapf 1898-1900
(471), Chippindall 1955 (84). Illustration: Chippindall
1955. Voucher: Acocks 15020, Smook 3652. PRECIS code
9903741-00200.

Loudetia Steud.

Annual (rarely), or perennial; caespitose. Culms
(250-)400-5000 mm high; herbaceous (usually erect,
slender or robust); branched above, or unbranched above.
Leaf blades linear (often rigid)-, flat, or rolled (convolute).
Ligule a fringed membrane (narrow), or a fringe of hairs.
Plants with hermaphrodite florets.

Inflorescence paniculate-, open, or contracted (rarely
more or less spiciform); espatheate. Spikelet-bearing axes
persistent.

1(0). Spikelets in triads, 20-30 mm long . . L. pedicel lata
Spikelets in pairs or solitary, less than 15 mm long

2

2(1). Inflorescence dense and spike-like . . L. densispica

Inflorescence an open panicle 3

3(2). Glumes obtuse or truncate 4

Glumes acute or minutely awned 5

4(3). Callus of female-fertile (upper) floret two-toothed;

leaves may be loosely hairy L. simplex

Callus of female-fertile (upper) floret pointed,
truncate or rounded; leaves densely covered with

velvety white hairs L. lanata

5(3). Culms with leaves few at base, branched after first

node; spikelets 6-8 mm long L. filifolia

Culms very leafy at base, unbranched after first node;
spikelets 8-12 mm long L. flavida

Loudetia densispica (Rendle) C.E. Hubb.

Perennial; tufted; to 900 mm
tall. Leaf blades 100-200 mm
long; to 3 mm wide. Spikelets
10-15 mm long. Panicle 40-80
mm long, spikelike and dense;
lower glume obtuse, with 2 rows
of tubercles; upper glume and
lower lemma usually glabrous;
callus of female-fertile (upper)
floret 2-toothed.

Flowering January. Open grassland. Locally common
(Komati River). Biome: Savanna. Angola, Lower Guinea.

Description: Chippindall 1955 (283). Illustration: Chip-
pindall 1955 (fig. 254). Voucher: Acocks 13308. PRECIS
code 9901751-00100.

206

Loudetia filifolia Schweick.

Slender, wiry perennial; tuft-
ed; to 600 mm tall. Leaf blades to
1 00 mm long; filiform or to 2 mm
wide. Spikelets 6-8 mm long.

Culms thin, branched, after the
first node; glumes acute or shortly
awned; callus of female-fertile
(upper) floret truncate or
rounded.

Flowering November to June. Rock crevices on cliffs
and mountain slopes. Infrequent. Biome: Savanna. Can be
distinguished from L. flavida, which has larger spikelets
(8-12 mm long) and wider leaves (2-4 mm long).

Description; Chippindall 1955 (283). Voucher: Van
Rooyen 3333. PRECIS code 9901751-00200.

Loudetia flavida (Stapf) C.E. Hubb.

Fig. 127.

Pointed russet grass.

Perennial; tufted; 800-1500
mm tall. Leaf blades 150^100
mm long; 2—4 mm wide. Spike-
lets 8-12 mm long. Leaves tend
to be widely spreading; glumes a-
cute or shortly awned, usually
glabrous and rarely tubercled;
lower lemma acute; female-fertile (upper) lemma lobes 1-2
mm long and acute, central awn 30-40 mm long, callus
pointed or truncate; stamens 3.

Flowering November to March. Shallow rocky soils,
also vlei margins. Common. Biome: Savanna and Grass-
land. East tropical Africa.

Description: Chippindall & Crook 1976 (93), Chippin-
dall 1955 (282). Illustration: Chippindall 1955 (fig. 253).
Voucher: Smook 912. PRECIS code 9901751-00300.

Loudetia lanata (Stent & Rattray) C.E. Hubb.

Woolly russet grass.

Perennial; tufted; 500 - 900
mm tall. Leaf blades 200 mm
long; 4 mm wide. Spikelets 8-12
mm long. Lower leaf sheaths
woolly; leaves thick with velvety
white hairs; panicle open, branch-
es purple or tinged; glumes ob-
tuse; callus of female-fertile (upper) floret pointed, awns
40-70 mm long and purple.

Flowering January to April. Edge of vleis in sandveld
areas. Common. Biome: Savanna. Angola, Zambia,
Zimbabwe.

Description: Chippindall & Crook 1976 (94). Voucher:
De Winter & Marais 4649. PRECIS code 9901751-00400.

Loudetia pedicellata (Stent) Chippind.

Perennial; tufted; to 1600 mm
tall. Leaf blades to 150 mm long;

4-6 mm wide. Spikelets 20-28
mm long. Ligule a conspicuous
fringe; spikelets in groups of 3(or
rarely paired), pedicels unequal
and 2 mm and 4 mm long respec-
tively; lower glume ovate and 1/2
the length of the upper; lower
lemma 7-nerved; female-fertile (upper) lemma 9-nerved,
lobes awned, 4-5 mm long, central awn 50-70 mm long.

Flowering December to April. Burkea-T erminaha veld.
Locally common. Biome: Savanna. Domestic use (thatch-
ing), or pasture (coarse hay). This species maybe confused
with Tristachya species, which also have the spikelets in

groups of three, but in Tristachya the lower lemma is 3-
nerved and the lower glume is acute, equal or more than
1/2 the length of the upper.

Description: Chippindall 1955 (280). Illustration: Chip-
pindall 1955 (fig. 251). Voucher: De Winter 722. PRECIS
code 9901751-00500.

Fig. 127. Loudetia flavida

207

Loudetia simplex (Nees) C.E. Hubb.

PI. 114.

Common russet grass, stingel-
gras, besemgras.

Perennial; tufted; 400-1500
mm tall. Leaf blades 100-300
mm long; 5 mm wide. Spikelets
7-13 mm long. Lower glume
obtuse or truncate; female-fertile
(upper) lemma lobes acute,

1 mm long, central awn 25-50 mm long, callus clearly two-
toothed in mature specimens; stamens 2.

Llowering throughout the year. Poor coarse, sandy soils
in open grassland or hillsides. Common (widespread).
Biome: Savanna and Grassland. Tropical Africa. Domestic
use (thatching and brooms). This species is exceedingly
variable, especially in panicle shape, hairiness of vegetative
parts and presence or absence of tubercles on the glumes
and lower lemma.

Description: Chippindall & Crook 1976 (95), Chippin-
dall 1955 (282). Illustration: Chippindall 1955 (PI. 8).
Voucher: Du Toil 2355. PRECIS code 9901751-00600.

Megaloprotachne C.E. Hubb.

Annual', caespitose, or decumbent (sometimes rooting at
the lower nodes). Culms 150-900 mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear,
flat. Ligule a fringed membrane. Plants bisexual, with
bisexual spikelets.

Inflorescence of spike-like main branches ( spike -like
racemes or narrow panicles)', digitate or subdigitate to non-
digitate (usually subdigitate, with several racemes from
below the apex); espatheate. Spikelet-bearing axes
persistent.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations (but spikelets homogamous ). Spikelets 4-5 mm
long; abaxial; compressed dorsiventrally; falling with the
glumes. Glumes two; more or less equal', awnless; very dis-
similar (the lower hairless, the upper with four dense rows
of long, green to dark purple hairs between the veins).
Proximal incomplete florets 7; paleate, palea fully
developed (two keeled); male.

Lemale-fertile florets 1. Lemmas decidedly firmer than
the glumes; striate; becoming indurated to not becoming in-
durated (cartilaginous-crustaceous); hairless; having the
margins lying flat and exposed on the palea; with a clear
germination flap; 3 nerved (the nerves obscure); entire;
awnless. Palea present; relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Lruit small (almost
2 mm long); hilum short; embryo large.

Photosynthetic pathway. C4; XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 1 species (supposedly 2). Southern
tropical and South Africa. Mesophytic to xerophytic; in
shade, or in open habitats (in open Acacia and mopane
savanna); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, and Cape Province. 1 or 2 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Roivainen. 1974. Ann. Bot. Fennici IT. 38^-2.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf sheaths of lower leaves with long woolly hairs
for some distance above the node . M. albescens
Leaf sheaths of lower leaves lacking long woolly hairs

except around node at base of sheath

M. glabrescens

Megaloprotachne albescens C.E. Hubb.

Fig. 128. PI. 115.

Erect or decumbent annual; to
800 mm tall. Leaf blades to 150
mm long; 3-4 mm wide. Spike-
lets 4. 0-4. 5 mm long. Long hairs
present on the leaf sheaths and
collar; lower glume as long as
spikelet.

Flowering February to April.
Sandveld. Infrequent. Biome: Sa-

208

vanna. ?Endemic. Easily mistaken for Digitaria, in which
the lower glume is never longer than 1/4 the spikelet length,
and is often absent.

Description: Roivainen 1974 (38-40), Chippindall 1955
(422). Illustration: Chippindall 1955 (fig. 351). Voucher:
Van Vuuren & Giess 1086. PRECIS code 9900881-00100.

Megaloprotachne glabrescens Roiv.

Erect or decumbent annual; to
700 mm tall. Leaf blades 60-100
mm long; 3-4 mm wide. Spike-
lets 3. 5-4.0 mm long. Long hairs
occur only on the lower parts of
the leaf sheaths.

Flowering January to May.

Sandveld. Conservation status not
known. Biome: Savanna.

?Endemic. A less hairy variant of M. albescens , probably
not a distinct species.

Description: Roivainen 1974 (40). Voucher: De Winter
& Giess 6957. PRECIS code 9900881-00200.

Megastachya P. Beauv.

Annual (tall, erect), or perennial (weakly); decumbent
(forming secondary shoots from the rooting nodes). Culms
300-1000 mm high; herbaceous; branched above. Leaf
blades linear-lanceolate to lanceolate; broad ; cordate
(amplexicaul)', flat. Ligule an unfringed membrane. The
spikelets all alike in sexuality.

Inflorescence paniculate; espatheate. Spikelet-bearing
axes persistent.

Spikelets 7-15 mm long; compressed laterally; disartic-
ulating above the glumes. Glumes two; very unequal; decid-
edly shorter than the adjacent lemmas; shortly awned (or
mucronate, from the excurrent mid-nerve); similar
(membranous-herbaceous, broadly ovate). Proximal incom-
plete florets absent.

Female-fertile florets 12-17. Lemmas incised; awnless,
or mucronate (the mucro from between the lobes, via the
excurrent mid-nerve); obscurely 5-7 nerved. Palea present
(narrower than lemma); relatively long; 2-nerved. Stamens
2-3. Ovary glabrous. Fruit small (about 1 mm long);
subglobose; hilum short; embryo small.

Transverse section of leaf blade. Mesophyll without arm
cells (according to Metcalfe), or with arm cells (?); with
fusoids (as represented by laterally extended PBS cells).
Midrib vascularization complex (1 large median with 2 tiny
laterals, all enclosed in a common sheath).

Cytology, classification, distribution. Chromosome base
number, .v = 12. Bambusoideae; Oryzodae; Centotheceae.
1 species. Tropical and southern Africa. Mesophytic; in
shade (in forests); glycophytic. Natal. 1 indigenous species.

References. 1. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

Megastachya mucronata (Poir.) Beauv.

Fig. 129. PI. 116.

Weak perennial, or annual;
sometimes stoloniferous; to 900
mm tall. Leaf blades 60-120 mm
long; 10-25 mm wide (broadly
lanceolate, base clasping, cross-
veins conspicuous). Spikelets
7-15 mm long. Culms decum-
bent, rooting at the lower nodes;
inflorescence an open panicle;

spikelets long-pedicelled, with many florets.

Flowering throughout the year (usually in spring). In
forests, often on sandy soil. Conservation status not known.
Locally common. Biome: forest. Tropical Africa.

Description: Chippindall 1955 (45). Illustration: Chip-
pindall 1955 (fig. 15). Voucher: Ward 8621. PRECIS code
9903881-00100.

Fig. 129 . Megastachya mucronata

209

Melica L.

Beckeria Bernh., Bromelica (Thurber) Farw., Claudia
Opiz, Dalucum Adans., Verinea Merino.

Perennial; long-rhizomatous. Culms 1 00— 1 500(— 2000)
mm high; herbaceous; unbranched above. Sheath margins
joined. Leaf blades linear; flat, or rolled (convolute). Ligule
an unfringed membrane to a fringed membrane, or a fringe
of hairs (rarely).

Inflorescence a single raceme , or paniculate ; open, or
contracted; espatheate. Spikelet-bearing axes persistent.

Spikelets 4-20 mm long; compressed laterally to not
noticeably compressed; disarticulating above the glumes, or
falling with the glumes (sometimes disarticulating both
above and below them). Glumes two; relatively large; very
unequal, or more or less equal; about equalling the spikelets
(usually?); long relative to the adjacent lemmas', awnless;
non-carinate\ very dissimilar, or similar. Incomplete florets
distal to the female-fertile florets, merely underdeveloped,
or clearly specialised and modified ( forming a hall of
successively enveloped lemmas or as a swollen rachilla
extension)', proximal incomplete florets absent.

Female-fertile florets 1-7. Lemmas similar in texture to
the glumes, or decidedly firmer than the glumes (leathery);
5-9 nerved, or rarely 10-15 nerved; entire, or incised;
awnless, or awned. Awns when present /; from the sinus,
or apical; non-geniculate; much shorter than the body of the
lemma to about as long as the body of the lemma. Palea
present; relatively long, or conspicuous but relatively short,
or very reduced. Lodicules 2; joined; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum long-linear;
embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 9. Pooideae; Poodae; Meliceae. About 80
species. North temperate, southern Africa and South
America. Mesophytic to xerophytic; in shade and in open
habitats. Transvaal, Orange Free State, Natal, Lesotho, and
Cape Province. 2 indigenous species.

References. 1. Gibbs Russell & Ellis. 1982. Bothalia
14,1: 37-44.

Species treatment by G.E. Gibbs Russell.

1(0). Lemmas hairy only on margins, glabrous or slightly
scabrous on back; sterile clusters glabrous or with

a few hairs; spikelets 5— 9(— 1 1 )mm long

M. racemosa

Lemmas hairy on the back and margins; sterile

clusters hairy; spikelets 10-15 mm long

M. decumbens

Melica decumbens Thunb.

(=M. neesii Stapf) 1 .

Dronkgras.

Perennial; tufted; 300-500
mm tall. Leaf blades 20-200 mm
long; 1.5-3. 5 mm wide (erect,
usually rolled, strongly scabrous).

Spikelets 10-15 mm long.

Lemmas of female-fertile florets hairy on the back and
margins; lemmas of sterile florets usually hairy.

Flowering October to April. Hillsides and

mountainsides, among rocks or in the shade of trees,
occasionally on roadsides. Infrequent. Biome: Grassland
and Nama-Karoo. Endemic. Poisonous (to horses, cattle and
donkeys).

Description: Gibbs Russell & Ellis 1982 (42), Stapf
1898-1900 (687), Chippindall 1955 (75). Illustration:
Gibbs Russell & Ellis 1982 (fig. 6), Chippindall 1955 (fig.
47). Voucher: Smith 4477. PRECIS code 9903860-00300.

Melica racemosa Thunb.

(=A7. holusii Stapf) 1; (=M.
brevifolia Stapf) 1; (=M.
decumbens sensu Gordon-Gray,
non Thunb.) 1; (=M. ovalis
Nees) 1; ( -M . pumila Stapf) 1.

Perennial; tufted; 300-500
mm tall. Leaf blades 40-300 mm
long; 1 .5-5.0 mm wide (erect, flat

Fig. 130. PI. 117.

210

or rolled, often scabrous). Spikelets 5— 9(— 1 1 ) mm long.
Lemmas of female-fertile florets hairy only on margins;
lemmas of sterile florets glabrous or with only a few hairs.

Flowering September to April. On steep hills and
mountain slopes among rocks and also in lightly shaded
places at edges of bushclumps and dune forest. Infrequent.
Biome: Fynbos, Grassland, and Nama-Karoo. Endemic.
Natal plants tend to have the larger spikelet size.

Description: Gibbs Russell & Ellis 1982 (41), Stapf
1898-1900 (687), Chippindall 1955 (74). Illustration:
Gibbs Russell & Ellis 1982 (fig. 3&4), Chippindall 1955
(fig. 46). Voucher: Edwards 4179. PRECIS code
9903860-00700.

Melinis P. Beauv.

Rhynchelytrum Nees, Suaria Schrank, Tristegis Nees.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 200-1200 mm
high; herbaceous; branched above, or unbranched above.
The shoots aromatic, or not aromatic. Leaf blades linear,
flat, or rolled. Ligide a fringed membrane (sometimes very
narrow) to a fringe of hairs. Plants with hermaphrodite
florets.

Inflorescence paniculate ( often decompound, rarely
composed of secund racemes ): open, or contracted: espathe-
ate. Spikelet-bearing axes persistent.

Spikelets not in distinct ’long-and-short’ combinations;
1-1 1 mm long; compressed laterally (often asymmetric), or
compressed dorsiventrally or not noticeably compressed;
disarticulating above the glumes (the fruiting floret falling
first), or falling with the glumes (falling from the pedicel).
Glumes one or two; very unequal; awned (G2 only,
sometimes awned or mucronate or beaked upwards), or
awnless; very dissimilar (G1 a scale up to 1/3 spikelet
length, or reduced to a vestige or rim, G2 longer, apically
emarginate or bifid, and often awned or mucronate from the
sinus, firmly membranous or papery, straight or curved on
the back). Lower glume 0-1 nerved. Proximal incomplete
florets T, paleate, or epaleate, palea when present fully
developed to reduced; male, or sterile. Proximal lemmas
similar in texture to the female-fertile lemmas.

Female-fertile florets 1 . Lemmas less firm than the
glumes, or similar in texture to the glumes (hyaline or
membranous to papery); smooth (shiny); not becoming in-
durated; hairless (usually glabrous, rarely ciliate); having
the margins lying flat and exposed on the palea; without a
germination flap; 1-5 nerved; entire (truncate), or incised
(emarginate or minutely two-lobed); awnless. Palea
present; relatively long. Lodicules 2; fleshy or
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small, ellipsoid; hilum short; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional. Biochemical type PCK
(M. repens, M. minutiflora); XyMS+. PCR cell chloroplasts
centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae
(Melinideae). About 26 species. Tropical Africa,
Madagascar, Arabia to Indochina, one species in tropical
South America and the West Indies. Mesophytic; in open
habitats (savanna and grassland, often in disturbed ground);
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 10
indigenous species.

References. 1. Clayton. 1978. Kew Bull. 33. 2. Clayton
& Renvoize. 1982. FTEA. 3. Zizka. 1988. Bibliotheca
Botanica 138.

Species treatment by H.M. Anderson.

1(0). Spikelets 1.0-1. 5(— 2.0) mm long 2

Spikelets (2. 2-)3.0-5.0(-12.0)mm long 4

2(1). Plants not sticky; pedicels hairy below the apex, hairs
few, 2-4 mm long; spikelets 1 mm long; upper

glume 3-5-nerved M, tenuissima

Plants sticky; pedicels not hairy below apex; spikelets
1.5-2.0 mm long; upper glume 7-nerved 3

211

3(2). Plants perennial, strongly aromatic; upper glume and
lower lemma grooved between the prominent
nerves; upper glume may be awned, the awn 0-9

mm long M. minutiflora

Plants usually annual and not aromatic; upper glume
and lower lemma not grooved between the

prominent nerves; upper glume not awned

M. macrochaeta

4(1). Leaf sheaths strongly overlapping; leaves rolled;
spikelets densely hairy; awns 1— 2(— 3) mm long;

always perennial M. nerviglumis

Leaf sheaths not strongly overlapping; leaves not
rolled; spikelets glabrous or hairy; awns mainly

longer than 2 mm; perennial or annual 5

5(4). Internode between the glumes 0.7-1. 7 mm long;
spikelets 5-12 mm long; upper glume and lower

lemma always tapering into a long beak

M. repens subsp. grandiflora

Internode between the glumes shorter than 0.7 mm;
spikelets shorter than 5 mm; upper glume and lower

lemma not tapering into a long beak 6

6(5). Awns of upper glume 5-10 mm and lower lemma

8.5-20.0 mm long

M. longiseta subsp. bellespicata

Awns less than 8.5 mm long 7

7(6). Spikelets glabrous or covered with short hairs 0.5 mm
long; internodes between the glumes 0.2-0. 7 mm

long 8

Spikelets covered with hairs 4-6 mm long; internodes
between the glumes usually less than 0.4 mm long

9

8(7). Rhizomes thick and knotty, internodes between the

glumes 0.2-0. 6 mm long M. subglabra

Rhizomes not thick and knotty, internodes between
the glumes 0.4-0.7 mm long . . . M. kallimorpha
9(7). Awns of upper glume 1-6 mm and lower lemma
4. 0-8. 5 mm long . M, longiseta subsp. longiseta
Awns of upper glume and lower lemma 1-3 mm long,
or sometimes absent . . M. repens subsp. repens

NOTE: M. scabrida is not included in the key. It is a
variable grass of possible hybrid origin occuring in
east Africa with only one record from the Transvaal

Melinis drakensbergensis (C.E. Hubb. & Schweick.)

Clayton

(= Rhynchelytrum drakensbergense C.E. Hubb. &
Schweick.) 1.

Zizka 1988 records that this may be a hybrid and that
he was unable to find further specimens. In view of the
uncertain validity of this species it is not treated here.

Description: Zizka 1988 (106). PRECIS code

9901340-00050.

Melinis kallimorpha (Clayton) Zizka

(=Rhynchelytrum
kallimorphon Clayton) 3.

Annual; tufted; 400-1000 mm
tall. Leaf blades 40-160 mm
long; 2-4 mm wide. Spikelets
3. 5-5.0 mm long; 1. 5-2.0 mm
wide. Rhizomes not thick and
knotty; spikelets glabrous or with
hairs 0.5 mm long; intemode between glumes 0.4-0. 7(— 0.9)
mm long; upper glume and lower lemma awned, awn 3-8
mm long, gibbous but not tapering into an elongated beak.

Flowering January to May. Sandy areas, prefers shade.
Infrequent. Biome: Savanna. Angola and east Africa.

Description: Zizka 1988 (64). Illustration: Zizka 1988
(fig. 22). Voucher: Smith 4031. PRECIS code 9901340-
00060.

Melinis longiseta (A. Rich.) Zizka subsp. bellespicata

(Rendle) Zizka

( -Rhychelytrum

bellespicatum (Rendle) Stapf &

C.E. Hubb.) 3.

Perennial; tufted; 200-800
mm tall. Leaf blades (30-)60-200
mm long; 1 .5— 5.0(— 6.0) mm
wide. Spikelets 3.5-5.0(-6.0) mm
long; 2 mm wide. Leaves hairy or glabrous; internode
between the glumes 0.3-0. 5 mm long; lower glume 0.9-1 .4
mm long; upper glume awned, 5-10 mm long; lower lemma
awned, awn 8-20 mm long.

Flowering February to June. Crevices in rocks, usually
in sunny places. Locally common. Biome: Savanna.
Angola, east Africa, Cameroon, Nigeria. Distinguished
from M. longiseta subsp. longiseta , which has shorter awns
(4.0-8. 5 mm long) and shorter internodes between the
glumes (to 0.3 mm long).

Description; Zizka 1988 (78). Illustration; Zizka 1987
(fig. 3 1 ), 1 (fig- 358). Voucher: Smook 5205. PRECIS code
9901340-00070.

Melinis longiseta (A. Rich.) Zizka subsp. longiseta

(= Rhynchelytrum longisetum
(A. Rich.) Stapf & C.E. Hubb.)
3; (=Rhynchelytrum minuti-
florum (Rendle) Stapf & C.E.
Hubb. var. melinoides (Stent)
Stapf & C.E. Hubb.) 2.

Perennial; tufted; 450-1000
mm tall. Leaf blades 40-120 mm
long; 4-9 mm wide. Spikelets

2*3-3. 8(-4. 2) mm long; 1.5-2 mm wide. Leaves hairy, rare-
ly glabrous; internode between the glumes rarely up to 0.3
mm long; lower glume 0.6-1 .0 mm long; upper glume awn-
ed, awn 1-6 mm long; lower lemma awned, awn 4. 0-8. 5
mm long.

Flowering March to July. Sandy areas, open woodland.
Infrequent. Biome: Savanna. Angola and east Africa to
Sudan. Distinguished from M. longiseta subsp. bellespicata,
which has longer awns (8-20 mm) and longer internodes
between the glumes (0.3-0. 5 mm).

Description: Zizka 1988 (75). Voucher: Volk 2159.
PRECIS code 9901340-00080.

Melinis macrochaeta Stapf & C.E. Hubb.

Fig. 132.

Mainly annual; tufted; 500-
1000 mm tall. Leaf blades 50-150
mm long; 5-10 mm wide. Spike-
lets 1. 5-2.0 mm long; 0.5 mm
wide. Culms often with stilt roots
from lower nodes; lower glume
reduced to a scale; upper glume
7-nerved, awnless and not
grooved; female-fertile (upper)
lemma with 3-5 nerves, awn (5— )8— 20 mm long.

Flowering April to June. Sand or loam, mainly
grassland. Infrequent. Biome: Grassland. Mainly southern
tropical Africa.

Description: Zizka 1988 (106). Illustration: Chippindall
1955 (fig. 356). Voucher: Compton 27774. PRECIS code
9901340-00100.

212

Fig. 132. Melinis macrochaeta

Melinis minutiflora Beauv.

PI. 118.

(= M . tenuinervis (Stapf)

Stapf) 2.

Molasses grass.

Perennial; tufted; 800-1500
mm tall. Leaf blades 40-200 mm
long; 5-1 1 mm wide. Spikelets
1. 5-2.0 mm long; 0.5 mm wide.

Leaves strongly aromatic and sticky, hairs finely tubercled,
exuding drops of viscid oil; spikelets glabrous and some-
times shortly hairy; upper glume 7-nerved, often with awn
0-9 mm long; upper glume and lower lemma grooved
between the prominent veins; lower lemma awnless or awn-
ed, awns 5-14 mm long.

Flowering April to June. Sand or near rocks, moist and
shady areas, grassland or savanna, open hillsides. Locally
common. Biome; Savanna and Grassland. Cultivated
throughout tropics. Cultivated pasture. The presence or
absence of awns and hairiness of spikelets is rather variable.

Description: Chippindall 1955 (427). Illustration: Clay-
ton et al. 1970-1982 (fig. 124). Voucher: Killick 1715.
PRECIS code 9901340-00200.

Melinis nerviglumis (Franch.) Zizka

(^Rhynchelytrum nerviglume
(Franch.) Chiov.) 3;

(=Rhynchelytrum nyassanum
(Mez) Stapf & C.E: Hubb.) 2;

(-Rhynchelytrum ramosum
Stapf & C.E. Hubb.) 2:

( =Rhynchelytrum rhodesianum
(Rendle) Stapf & C.E. Hubb.) 3;

(-Rhynchelytrum setifolium (Stapf) Chiov.) 2.

Perennial; tufted; (250-)400-1200(-1500) mm tall. Leaf
blades (30-) 100-300(^440) mm long; ( 1 .3— )2.0— 3.5( — 4.5 )
mm wide. Spikelets (3.2— )3.6 — 5.0(— 5.7) mm long; 2 mm
wide. Basal leaf sheaths strongly overlapping; leaf blades
rolled; spikelets often densely covered with hairs up to 4
mm long, white or purple; intemode between glumes
0.3(-0.6) mm long; lower glume 0.5 mm long and awns
1— 2(— 3) mm long.

Flowering November to September. Open grassland,
stony hillsides. Locally dominant. Biome: Fynbos, Savan-
na, and Grassland. Sub-Saharan Africa, Madagascar,
possibly introduced to southeast Asia. Very similar to M.
repens subsp. repens, mainly distinguished by lacking
strongly overlapping leaf sheaths and rolled leaf blades.

Description: Zizka 1988 (111). Illustration: Chippindall
1955 (fig. 359). Voucher: Smook 5268. PRECIS code
9901340-00250.

Melinis repens (Willd.) Zizka subsp. grandiflora
(Hochst.) Zizka

( —Rhynchelytrum brevipilum
(Hack.) Chiov.) 2;

(=Rhynchelytrum costatum Stapf
& C.E. Hubb.) 2;

( =Rhynchelytrum grandiflorum
Hochst.) 3; (= Rhynchelytrum
villosum (Pari.) Chiov.) 2.

Annual; tufted; 250-900 mm
tall. Leaf blades 40— 1 50(— 1 80) mm long; 2.0-6. 5(— 8.0) mm
wide. Spikelets (4 — )5 — 12 mm long; 2-3 mm wide. Spikelets
glabrous to hairy; intemode between the glumes (0.5-)
0.7—1 .7(— 2.0) mm long; lower glume (0.6 — ) 1 .5— 3.0( — 4.3)
mm long; upper glume and lower lemma gibbous, tapering
into an elongated beak, awns (2-) 12-22 mm long.

Flowering January to July. More often in sunny arid
areas, not a ruderal. Locally dominant. Biome: Savanna and
Nama-Karoo. Most of Africa and as far as India.
Distinguished from M. repens subsp. repens, which is
commonly a ruderal, and has spikelet length mostly 2-4
mm, and intemode between glumes mostly 0.1-0.5(-0.6)
mm long.

Description: Zizka 1988 (60). Voucher: Giess & Van der
Walt 12666. PRECIS code 9901340-00275.

(= Rhynchelytrum repens
(Willd.) C.E. Hubb.) 3.

Natal red top.

Mainly annual, or perennial
(rarely); tufted; 250-1 200(-1500)
mm tall. Leaf blades 40-200
(-270) mm long; 2—1 1 (—1 3) mm

wide. Spikelets 2.2-4.0(-5.0) mm long; 2-3 mm wide.
Spikelets always hairy; intemode between the glumes
0. 1-0.5 (-0.6) mm long; lower glume (0.3— )0.6— 1 .3(— 1 .5)
mm long; upper glume and lower lemma gibbous, rarely
tapering into an elongated beak, mostly shortly awned, to
3 mm long.

213

Flowering September to May. Mainly a ruderal,
common on disturbed ground. Common. Biome: Fynbos,
Savanna, Grassland, and Nama-Karoo. Widely spread in
tropical Africa, a common innocuous weed throughout the
tropics. Erosion control, ornamental, and weed. A very
variable and widely distributed species. Distinguished from
M. repens subsp. grandiflora which is not usually a ruderal,
and has spikelets mainly 5-12 mm long and the internode
betweeen the glumes 1. 5-3.0 mm long.

Description: Zizka 1988 (57). Illustration: Chippindall
1955 (pi. 12). Voucher: Smook 4767. PRECIS code
9901340-00300.

Melinis scabrida (K. Schum.) Hack.

(-Rhynchelytrum scabridum
(K. Schum.) Chiov.) 3.

Perennial; loosely tufted;

400-1000 mm tall. Leaf blades
20-140 mm long; 2. 8-6. 5 mm
wide. Spikelets 2.4-2.8(-3.2) mm
long. Upper glume with awns to
0.3 mm long, 5-7-nerved;
female-fertile (upper) lemma with awns 1. 0-2.4 mm long
and 5-nerved.

Open hillsides. Infrequent. Biome: Grassland. East
Africa. Recorded as of possible hybrid origin by Clayton
& Renvoize 1982 (511), M. ambigua x M. longisetum.

Description: Clayton et al. 1970-1982 (511).

Illustration: Zizka 1988 (fig. 33). No specimens seen.
PRECIS code 9901340-00325.

Melinis subglabra Mez

(- Rhynchelytrum
suberostratum Stapf & C.E.

Hubb.) 2; (= Rhynchelytrum sub-
glabrum (Mez) Stapf & C.E.

Hubb.) 3.

Perennial; tufted; 400-1300
mm tall. Leaf blades 30-170 mm
long; (2.3-)3.0-8.0(-10.0) mm
wide. Spikelets 3. 2-5.0 mm long;

1.5 mm wide. Rhizome thick and knotty; upper glume and
lower lemma with awns up to 3 mm long.

Flowering February to June. Prefers shady areas near
water. Biome: Savanna. Angola and east Africa.

Description: Clayton et al. 1970-1982 (513).

Illustration: Zizka 1988 (fig. 25). Voucher: Codd 1588.
PRECIS code 9901340-00350.

Melinis tenuissima Stapf

Perennial; tufted; 500-1100
mm tall. Leaf blades 20-80 mm*
long; 3-6 mm wide. Spikelets
1 . 1—1 .5(— 1 .6) mm long; 0.5 mm
wide. Inflorescence open and
loose, 80-200 mm long and often
almost as wide; hairs 2-4 mm
long on pedicel at base of spike-
let; lower glume reduced to a
scale 0.1 mm long; upper glume 5-nerved, awnless and not
grooved; female-fertile (upper) lemma 3-5-nerved, white
awn (1.5-)4.0-10.0 mm long.

Flowering April to June. Grassland and bush, often near
water or in cultivation. Rare. Biome: Savanna. Tropical
Africa.

Description: Chippindall 1955 (427). Illustration:
Hooker’s Icon pi. 2660. Voucher: Scheepers 1 153. PRECIS
code 9901340-00400.

Merxmuellera Conert

Sometimes included in Rytidosperma, Danthonia sensu
lato.

Perennial; caespitose. Culms 150-2000 mm high; herba-
ceous; unbranched above. Leaf blades linear, 4-15 mm
wide-, nearly always rolled. Ligule a fringe of hairs.

Inflorescence a single raceme to 60 mm long (rarely —
M. disticha), or paniculate ; contracted ( narrow ,
occasionally spike-like; usually longer than 60 mm, by
contrast with Karroochloaf, espatheate. Spikelet-bearing
axes persistent.

Spikelets 8-25 mm long-, compressed laterally; disarticu-
lating above the glumes. Hairy callus present (0.6-2 mm,
with bearded margins). Glumes two; more or less equal (G2

214

somewhat shorter); about equalling the spikelets to much
exceeding the spikelets-, awnless; similar (papery, margins
and apex hyaline, midnerve percurrent). Incomplete florets
distal to the female-fertile florets, merely underdeveloped;
proximal incomplete florets absent.

Female-fertile florets 3-10. Lemmas similar in texture
to the glumes; hairy, or hairless (rarely glabrous, the hairs
when present in tufts; not in transverse rows); without a ger-
mination flap; (7-)9 nerved; incised; awned. Awns 1 , or i;
median, or median and lateral (the lobes sometimes finely
awn-tipped). The median awn different in form from the
laterals (when laterals present); from the sinus (the lobes
sometimes basally adherent to the median awn); geniculate;
much longer than the body of the lemma. Palea present; rel-
atively long (lanceolate); 2-nerved. Lodicules 2;
membranous; ciliate (often), or glabrous. Stamens 3. Ovary
glabrous. Fruit small (2-3 mm); pericarp fused.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, .v = 6. Arundinoideae; Danthonieae. 17 species.
South and southwest Africa. Mesophytic to xerophytic
(often in mountains); in open habitats; glycophytic.
Namibia, Transvaal, Orange Free State, Natal, Lesotho, and
Cape Province. 17 indigenous species.

References. 1. Conert. 1970. Senck. Biol. 51: 129. 2.
Conert. 1971. Mitt. Bot. Stsamml. Munch. 10: 299. 3. Ellis.
1980a. Bothalia 13: 185-189. 4. Ellis. 1980b. Bothalia 13:
191-198. 5. Ellis. 1981a. Bothalia 13: 487-491. 6. Ellis.
1981b. Bothalia 13: 493-500. 7. Ellis. 1982a. Bothalia 14:
89-93. 8. Ellis. 1982b. Bothalia 14: 95-99. 9. Ellis. 1983.
Bothalia 14: 197-203.

This genus is poorly known, and is in urgent need of
revision. Ellis has studied the taxa anatomically and, where
applicable, his observations have been noted.

Species treatment by N.P. Barker.

1(0). Base of plant densely woolly 2

Base of plant glabrous 4

2(1). Spikelets not densely clustered and panicle branches
and pedicels are partly visible; glumes 15— 22(— 25)
mm long, occasionally pubescent . . . . M. decora
Spikelets densely clustered such that panicle branches
and pedicels are obscured; glumes 7-18 mm long,

never pubescent 3

3(2). Glumes 3-5-nerved; lemmas 7-12 mm long,
including lobes; central awn of lemmas 6-16 mm
long, usually geniculate; spikelets 4-7-flowered .

M. rufa

Glumes l(-3)-nerved; lemmas 6-8 mm long,
including lobes; central awn of lemmas 4-8 mm
long, seldom geniculate; spikelets 2-5-flowered .

M. lupulina

4(1). Lemma backs with hairs in obvious tufts or rows . 5
Lemma backs densely pubescent, hairs not in obvious

tufts M. arundinacea

5(4). Lemma backs with 3 distinct tufts of long, white hairs
on each side of the central nerve, rest of lemma

glabrous or sparsely pubescent 6

Lemma backs with more than 3 or less than 3 distinct
tufts of long, white hairs on each side of the central
nerve, or else with a row of hairs across the lemma

backs 11

6(5). Lemma lobes completely adnate to central awn, with
no lateral bristles; central lemma awn seldom

geniculate M. macowanii

Lemma lobes wholly or partly free from central awn,
each lobe usually terminating into a bristle; central

awn usually geniculate 7

7(6). Tufts of hairs on lemma backs 5-7 mm long, the most
basal and marginal tuft slightly shorter, the rest of

the lemma usually sparsely pubescent

M. papposa

Tufts of hairs on lemma backs up to 3.5 mm long, the

rest of the lemma glabrous 8

8(7). Lemmas 10-13 mm long, including lobes and bristles,
lobes wholly free from the central awn; glumes
15-20 mm long, 3-5-nerved; central awn up to 15

mm long M. aureocephala

Lemmas 6. 5-9.0 mm long, including the lobes and
bristles, lobes partly free from central awn; glumes
9-17 mm long, 1-3-nerved; central awn 9-12 mm

long 9

9(8). Leaf blades cylindrical with an adaxial groove and
pungent apices; spikelets 2-flowered; plant up to

300 mm tall; from Namibia

M. rangei

Leaf blades involute or permanently infolded with
hard but not pungent apices; spikelets 2-8-
flowered; plant usually taller than 300 mm; not
found in Namibia 10

10(9). Glumes 11-13 mm long, 1 -nerved; remains of the
dead leaves curl into a spiral at base of plant; the
three tufts of hairs on each side of the central
nerve on the lemma backs spaced unequally
apart, the marginal tuft being positioned some
distance closer to the base of the lemma than the
other two more central tufts; from the northern

Transvaal Drakensberg and further north

M. davyi

Glumes 13-17 mm long, 1-3-nerved; remains of
dead leaves split, the two halves curling away
from each other; the three tufts of hairs on each
side of the central nerve on the lemma backs
approximately equidistant from each other;
mainly from the Natal Drakensberg, but also
extends into the Cape and eastern Transvaal . .

M. drakensbergensis

11(5). Lemma backs with a row of 5— 7(— 1 2) mm long,
white hairs across the middle, glabrous below

this M. cincta

Lemma backs with tufts of hairs and/or fringed
margins, but never with a row of hairs across the

middle 12

12(11). Lemma margins not fringed, backs with 4 or more
marginal tufts of white hairs near the base, the
remainder of the lemma surface glabrous or

pubescent 13

Lemma margins with a fringe of hairs, with or
without 1 tuft of white hairs on each side of the
central nerve, the remainder of the lemma surface

glabrous 14

13(12). Lemma pubescent at least down central nerve,
sometimes completely pubescent basally
between the tufts; glumes 9-14 mm long;
spikelets 3-4(-5)-flowered . M. guillarmodiae
Lemma glabrous in centre of back between tufts;
glumes 1 1-22 mm long; spikelets 5-7-flowered
M. stricta

14(12). Lemma margins fringed with hairs 15

Lemma margins not fringed with hairs 16

15(14). Inflorescence a spike-like panicle, 20-100 mm
long; spikelets distichous; lemmas with 1 basal
tuft of hairs on each side of the central nerve,
with a marginal fringe running from this tuft

upwards M. disticha

Inflorescence a contracted, shortly branched
panicle up to 180 mm long; spikelets not
distichous; lemma margins fringed from base
upwards, fringe ending in an apical tuft of white

hairs M. stereophylla

16(14). Glumes 13-18 mm long, 3-5-nerved; spikelets 4-7-
flowered; plant base not bulbous .... M. dura
Glumes 9-13 mm long, 1-nerved; spikelets 3( — 4)-

flowered; plant base bulbous

M. sp. (=EIlis 5500)

215

Merxmuellera arundinacea (Berg.) Conert

PI. 120.

( =Danthonia arundinacea
(Berg.) Schweick.) 1.

Tall, reedlike perennial;
densely tufted; 1000-1200 mm
tall. Leaf blades to 600 mm long;

4-7 mm wide. Spikelets (9.0-)

13.5-16.5 mm long; 9-11 mm
wide. Panicle densely contrac-
ted, 120-250 mm long; spikelets (2-)3-4-flowered; glumes
10-15 mm long, 1-nerved and usually pubescent; lemmas
6-8 mm long, including the lemma lobe which extends into
a short, soft bristle; lemma back completely covered with
white hairs; central awn 9-13 mm long.

Flowering August to November. Xeric areas on north
facing slopes of the Cape fold mountains. Locally common
(warm slopes with northern aspect). Biome: Fynbos.
Endemic. Similar in habit to Merxmuellera cincta , which
has tufted hairs on the lemma. Studied anatomically by Ellis
(1982a).

Description: Stapf 1898-1900 (524), Chippindall 1955
(429). Voucher: Ellis 2474. PRECIS code 9902043-00150.

Merxmuellera aureocephala (J.G. Anders.) Conert

(-Danthonia aureocephala
J.G. Anders.) 1.

Perennial; densely tufted; 900
mm tall. Leaf blades 400 mm
long; 1.5 mm wide. Spikelets to
23 mm long (including awns); to
10 mm wide (including awns).

Panicle contracted, interrupted, to
170 mm long; spikelets 3^4-flowered; glumes 15-20 mm
long, 3-5-nerved, golden-brown; lemmas 10-13 mm long,
including lobes and bristles which are 5. 5-7.0 mm long and
free from central awn; lemma backs with 3 tufts of white
hairs on either side of middle nerve, lowermost tuft margin-
al and somewhat distant from other 2 tufts; awn to 15 mm
long, sometimes strongly geniculate.

Flowering July to August. High Drakensberg mountains
in xeric areas such as steep grassy slopes above 2000 m.
Locally common (Giants Castle and Cathkin Peak). Biome:
Afromontane. Endemic. This species flowers in the winter
months and is therefore reproductively isolated from the
other alpine Merxmuellera species which tend to flower
from August to January.

Description: Anderson 1962 Bothalia 8: 170-172.
Voucher: Edwards 2453. PRECIS code 9902043-00250.

Merxmuellera cincta (Nees) Conert

( =Danthonia cincta Nees) 1.

Tall, reedlike perennial;
densely tufted; to 2000 mm tall.

Leaf blades 1000 mm long (or
more); 5-15 mm wide. Spikelets
to 14 mm long; to 8 mm wide.

Panicle dense, contracted, 200-
400 mm long; spikelets 3-4-flow-
ered; glumes 10—1 3(— 1 8) mm long, 1-nerved; lemmas 7-8
mm long, including 3. 5^4.0 mm long lobes which are free
from the central awn; lemma backs with a tufted row of
5— 7(— 1 2) mm long, white hairs across the middle; central
awn 5-14 mm long, sometimes longer.

Flowering September to February. Moist areas such as

seeps and stream banks on the south facing mountain
slopes. Locally common (in damp areas). Biome: Fynbos.
Endemic. Ellis(1982a)has noted the presence of lacunae in
the leaf blades of this species, an apparent adaptation to
aquatic environments. There are two forms of this species.
The less common form is found only in very sandy habitats
and has larger floral parts (e.g. glumes 18 mm long) and
very long tufts of hairs (10 mm or more) on the lemmas.
The more common form has glumes 10-13 mm long and
lemma hair tufts 5-7 mm long.

Description: Stapf 1898-1900 (526), Chippindall 1955
(250). Voucher: Burger 82. PRECIS code 9902043-00300.

Merxmuellera davyi (C.E. Hubb.) Conert

( -Danthonia davyi C.E.

Hubb.) 1.

Perennial; tufted; to 1000 mm
tall. Leaf blades to 600 mm long;
to 1.5 mm wide. Spikelets 15-17
mm long (including awns); 6-8
mm wide (including awns). Dead
leaves break off above sheath
mouth and the remains curl into a tight spiral; panicle dense,
150-250 mm long; spikelets 2-4-flowered; glumes 1 1-13
mm long, 1-nerved; lemmas 7. 5-9.0 mm long, including
lemma lobes which are adnate to the central awn for 1/4-3/4
of their length, ending in a short bristle; lemma body with
3 unequally spaced tufts of white hairs on either side of cen-
tral nerve, the most basal tuft being marginal and closer to
the base of the lemma than the other two tufts which are
equidistant, or almost so, from the lemma base; central awn
12 mm long.

Flowering September. Xeric, rocky, mountainous areas.
Locally common (in the eastern Transvaal Drakensberg).
Biome: Afromontane. Northwards to central Africa. This
species is anatomically very similar to Merxmuellera
macowanii (Ellis 1981b).

Description: Hubbard 1937 FTA 10: 137. Voucher:
Davidson & Mogg 33315. PRECIS code 9902043-00350.

Merxmuellera decora (Nees) Conert

(-Danthonia zeyheriana
Steud. var. trichostachya Stapf)

2; (=Danthonia zeyheriana
Steud. var. zeyheriana ) 2.

Perennial; tufted; 250-700
mm tall. Leaf blades to 200 mm
long; to 1.5 mm wide. Spikelets
18-25 mm long; to 12 mm wide.

Culm bases bulbous, surrounded by old, woolly, persistent
leaf sheaths; panicle loosely contracted, interrupted so that
panicle branches are visible, 50-130 mm long; spikelets
4-7-flowered; glumes 15-22(-25) mm long, 5-7-nerved,
sometimes densely pubescent; lemmas 9-15 mm long, in-
cluding 5-8 mm long lobes which are adnate to the central
awn for 1/3-1/2 their length, ending in a short, soft bristle;
lemma backs covered from base to middle in short, dense
hairs, with a row of white hairs across middle of back,
glabrous above this; central awn 12-20 mm long.

Flowering August to December. Sandy soils of mountain
slopes of southwestern Cape. Locally common (in naturally
burnt areas and firebreaks). Biome: Fynbos. Endemic. Ellis
(1983) distinguishes three anatomical forms. Variety
trichostachya Stapf was not recognised by Conert, but it is
easily distinguishable by the pubescent glumes and woolly
upper sheaths.

Description: Stapf 1898-1900 (521), Chippindall 1955
(245). Illustration: Chippindall 1955 (fig. 216). Voucher:
Ellis 2543. PRECIS code 9902043-00400.

216

Merxmuellera disticha (Nees) Conert

(= Danthonia disticha
Nees) 1.

Perennial; tufted; 150-700
mm tall. Leaf blades 100-500
mm long; to 3.5 mm wide. Spike-
lets to 18 mm long (including
awns); to 3 mm wide. Panicle
spike-like, 20-100 mm long, with
spikelets distichous, 2(-5)-flowered; glumes 9-20 mm
long, 1-3-nerved; lemmas 10-15 mm long including 7-9
mm long lobes which attenuate into a long bristle; lemmas
fringed along the margin with a tuft of white hairs each side
near the base; central awn geniculate, 10-16 mm long.

Flowering October to May. A variety of habitats, from
coastal regions to high altitude montane bogs. Common (in
certain veld types such as Karroid Merxmuellera Mountain
Veld). Biome: Fynbos, Grassland, Nama-Karoo_and Afro-
montane. Zimbabwe. Weed (can usurp valuable grazing
grasses in some areas). There are 3 anatomical forms: a
‘typical form’, an ‘alpine bog form’ and a ‘Drakensberg
form’. These forms may also be distinguished
morphologically, as outlined by Ellis (1980a). This taxon
may be confused with Pentaschistis hasutorum, which has
long, lax hairs covering the lemma.

Description: Stapf 1898-1900 (529), Chippindall 1955
(249). Illustration: Chippindall 1955 (fig. 220). Voucher:
Acocks 1 1961. PRECIS' code 9902043-00500.

Merxmuellera drakensbergensis (Schweick.) Conert

Fig. 133.

( =Danthonia drakensberg-
ensis Schweick.) 1.

Perennial; tufted; to 1000 mm
tall. Leaf blades to 300 mm long;
to 1.3 mm wide. Spikelets 12-17
mm long (including awns); 8-13
mm wide. Old leaves break off
close to the sheath mouth, and the
remaining leaf bases split along the midrib, the two halves
curling away from each other; panicle 80-180 mm long,
loosely contracted and interrupted; spikelets (5— )6— 8-
flowered; glumes 13-17 mm long, 1-3-nerved; lemmas
6. 5-9.0 mm long, including lobes 3. 5-5.0 mm long, adnate
to the central awn fot approximately half their length; lem-
ma backs with 3 tufts of equally spaced white hairs on either
side of central nerve, the marginal tuft the most b^sal; centr-
al awn 9-12 mm long, geniculate at point where lemma
lobes detatch from awn.

Flowering October to March. Mesic sites in
streambanks, mud patch communities and rocky outcrops
of the alpine belt, in areas where the soil is deeper than in
the surrounding areas. Common (streambanks and seeps).
Biome: Afromontane. Endemic. This species is anatomical-
ly very similar to Merxmuellera stereophyllai Ellis 1981a).

Description: Schweickerdt 1938 Fed. Rep. 43:88-89,
Chippindall 1955 (248). Illustration: Chippindall 1955
(fig. 219). Voucher: Ellis 3304. PRECIS code 9902043-
00600.

Merxmuellera dura (Stapf) Conert

Fig. 134.

( =Danthonia dura Stapf) 1.

Perennial; shortly rhizomat-
ous; 600-900 mm tall. Leaf
blades to 600 mm long; to 1 .5 mm
wide. Spikelets 20-25 mm long
(including awns); 6-8 mm wide.

Panicle 100-180 mm long,
loosely contracted, slightly nod-
ding; spikelets 4-7-flowered; glumes 13-18 mm long, 3-5-
nerved; lemmas 7-12 mm long, including 4—7 mm long
lobes terminating abruptly into a short, soft bristle; lemma

back glabrous except for a marginal tuft of white hairs by
the lemma base and a smaller tuft of hairs at the base of
the central awn; central awn 10-15 mm long.

Flowering July to November. Stony or sandy soils in
arid areas. Locally common (Carnavon and Calvinia
districts). Biome: Nama-Karoo and Succulent Karoo.
Endemic. Ellis (1982b) considers this species anatomically
distinct from M. stricta, despite some morphological
similarities.

Description: Stapf 1898-1900 (527), Chippindall 1955
(248). Voucher: Ellis 2464. PRECIS code 9902043-00700.

Merxmuellera guillarmodiae Conert

Perennial; tufted; 1 20 — 400
(-700) mm tall. Leaf blades
200^100 mm long; 0.4— 0.6 mm
wide. Spikelets 12-15 mm long
(including awns); 6-8 mm wide.

Panicle 40-90 mm long, inter-
rupted; spikelets 3^t(-5)-flower-
ed; glumes 9-14 mm long, 3-5-
nerved; lemmas 6-7 mm long
including 3 mm long lobes which are adnate to the central
awn for most of their length and which terminate in a short,
soft bristle; lemma backs with 4 or more, sometimes in-
distinct, tufts of white hairs along each margin near the
base, the rest of the lemma surface varying in pubescence
from a quite dense, basal pubescence to almost glabrous
with a few hairs each side of central nerve; central awn
5-1 1 mm long.

Flowering November to February. Grassland and rocky
areas above about 2000 m. The alpine form is asscoiated
with moist habitats. Locally common (Drakensberg).
Biome: Afromontane. Endemic. Ellis (1980a) recognises
two anatomical forms; the ‘Cathkin Peak form’ and the
‘alpine form’, which are morphologically separable:
‘Alpine form’ - lemmas sparsely pubescent, hairs short,
awns 5. 0-6. 5 mm long. ‘Cathkin Peak’ form - lemmas
densely pubescent, awn 8-1 1 mm long.

217

Description: Conert 1975 Senck. Biol. 56 (145).
Illustration: Conert 1975 Senck. Biol. 56 (145). Voucher:
Jacot Guillarmod 3727. PRECIS code 9902043-00750.

Merxmuellera lupulina (Thunb.) Conert

(^Danthonia lupulina
(Thunb.) Beauv. ex Roem. &

Schult.) 2.

Perennial; tufted; 400 mm tall.

Leaf blades 75-150 mm long; to
3 mm wide. Spikelets 9-12 mm
long; 6-8 mm wide. Culm bases
bulbous, covered in old, per-
sistent, densely woolly leaf sheaths; panicle densely
contracted such that panicle branches are obscured, 20-35
mm long; spikelets 2-5-flowered; glumes 7-10 mm long,
l(-3)-nerved; lemmas 6-8 mm long, including 2. 5-3. 5 mm
long lemma lobes adnate to the central awn for part of their
length, usually ending in a short, soft bristle; lemma backs
covered from base to middle by short, dense hairs, with a
row of long, white hairs across the middle above which the
lemma is glabrous; central awn 4-8 mm long, seldom
geniculate.

Flowering October to January. Sandy mountain slopes
of southwestern Cape. Locally common (in naturally burnt
areas and firebreaks). Biome: Fynbos. Endemic. This
species is anatomically similar to M. rufa, Pentaschistis
argentea and P. viscidula (Ellis 1983).

Description: Stapf 1898-1900 (523), Chippindall 1955
(245). Voucher: Taylor 5477. PRECIS code 9902043-
00800.

Merxmuellera macowanii (Stapf) Conert

( =Danthonia macowanii
Stapf) 1.

Perennial; tufted; 700-1300
mm tall. Leaf blades to 650 mm
long; to 1.3 mm wide. Spikelets
to 13 mm long (including awns);
to 6 mm wide. Old leaves break
off above sheath mouth, the
remaining leaf bases sometimes split along midrib and the
two halves curl away from each other; panicle 170-270 mm
long, loosely contracted, interrupted and shortly branched;
spikelets (2-)3-4-flowered; glumes 9-14 mm long, 1-3-
nerved; lemmas 10 mm long, including central awn to which
lemma lobes are usually fully adnate, lateral bristles absent;
lemma backs with 3 equally spaced tufts of white hairs on
either side of central nerve, the tuft on the margin being
most basal; central awn 5-8 mm long, seldom geniculate.

Flowering July to January. In montane and subalpine
regions at altitudes between 1500 and 3000 m. Locally
dominant (stream banks and marshy areas of the montane
Drakensberg). Biome: Afromontane. Endemic. Domestic
use (used for making brooms in Lesotho). Anatomically
very similar to M. aureocephala and M. davyi, with which
this species is closely allied and not consistently separable
(Ellis 1981b).

Description: Stapf 1898-1900 (527), Chippindall 1955
(248). Voucher: Ellis 3282. PRECIS code 9902043-00900.

Merxmuellera papposa (Nees) Conert

(-Danthonia papposa
Nees) 1.

Perennial; tufted; to 500 mm
tall. Leaf blades 120-300 mm
long. Spikelets 20-25 mm long
(including awns); 5-7 mm wide
(excluding awns). Panicle 1 20—

150 mm long, dense; spikelets
2-3-flowered; glumes 13-18 mm long, 3-nerved; lemmas

9-10 mm long including 5 mm long lobes which are free
from the central awn; backs of lemmas with 3 tufts of long
(5-7 mm) white hairs on each side of the central nerve, the
most basal being shorter, marginal and situated some
distance from the other two more apical tufts, with the rest
of the lemma surface below these tufts being sparsely
pubescent; central awn 15-18 mm long, geniculate.

Flowering December and January. Infrequent (known
from only a few fragments at PRE, all apparently from the
Uitenhage area). Biome: Fynbos. Endemic. Despite the
poor quality of specimens seen, the very long tufts of hairs
on the lemmas distinguish this species from other

Description: Stapf 1898-1900 (527). Voucher: No
voucher given as there is no proper material. PRECIS code
9902043-01000.

Merxmuellera rangei (Pilg.) Conert

(-Danthonia rangei Pilg.) 1.

Perennial; tufted; 120-300
mm tall. Leaf blades 35-140 mm
long; to 1.3 mm wide. Spikelets
1 1-14 mm long (including awns);
to 2 mm wide. Upper 2 nodes of
culm often conspicuously dark
brown or black; basal sheaths
persistent and papery; leaf blades cylindrical with small
adaxial groove, pungent; panicle 40-60 mm long, contract-
ed and partially enclosed by uppermost sheath; spikelets 2-
flowered; glumes 9-12 mm long, 1-3-nerved; lemmas 7
mm long including 4 mm long lobes; lemma backs with 3
tufts of white hairs on either side of the central nerve;
central awn geniculate, 9-10 mm long.

Flowering August to October. Dry, sandy areas between
hills and koppies. Conservation status not known. Biome:
Nama-Karoo. Endemic. Natural pasture (for sheep and
goats). Ellis (1982b) postulates that this species has an
affinity with Dregeochloa.

Description: Pilger 1909 Bot. Jb. 43: (386), Launert
1970 (160:128). Voucher: De Winter & Giess 6323.
PRECIS code 9902043-01 100.

Merxmuellera rufa (Nees) Conert

(=Danthonia lanata (Schrad.)

Schrad. var. lanata ) 2;

(=Danthonia lanata (Schrad.)

Schrad. var. maior Nees) 2;

(=Danthonia macrocephala
Stapf) 2.

Perennial; tufted; to 400 mm
tall. Leaf blades to 200 mm long;
to 4.5 mm wide. Spikelets to 25 mm long (including awns);
to 15 mm wide. Base bulbous, covered in old, persistent,
woolly sheaths; panicle 30-70 mm long, globose or cylin-
drical, with spikelets densely clustered; spikelets 4-7-
flowered; glumes 10-18 mm long, 3-5-nerved; lemmas
7-12 mm long, including 3. 0-6. 5 mm long lobes which are
adnate to the central awn for part of their length and usually
end in a short, soft bristle; lemma backs shortly and
densely pubescent from base to middle region, with row
of white hairs across the middle above which the lemma is
glabrous; central awn 6-16 mm long, geniculate.

Flowering September to December. Sandy soils on
mountain slopes in southern and southwestern Cape. Lo-
cally common (in naturally burnt areas and fire breaks).
Biome: Fynbos. Endemic. Ellis (1983) described three
anatomical forms of this species, one of which is actually
a variety of M. decora. The two remaining forms which
constitute M. rufa are morphologically distinguishable, and
account for the wide range of variation in this taxon. The
first form has shorter glumes (10-12 mm) with 3 indistinct
nerves near the base, and a central lemma awn 6-9 mm
long. The second form is larger, with glumes 16-18 mm

218

long, 3-5 obvious basal nerves, and a central lemma awn
1 1-16 mm long. Both forms are anatomically similar to M.
lupulina.

Description: Stapf 1898-1900 (522), Chippindall 1955
(244). Voucher: Ellis 2517. PRECIS code 9902043-01200.

Merxmuellera stereophylla (J.G. Anders.) Conert

(=Danthonia stereophylla
J.G. Anders.) 1 .

Perennial; tufted; 800 mm tall.

Leaf blades to 360 mm long; to
1 .5 mm wide. Leaves rigid, erect;
panicle contracted, shortly bran-
ched, to 180 mm long; spikelets
(3-)4-5-flowered; glumes 1 1-18
mm long, 1-nerved; lemmaslO-16 mm long, including 5-7
mm long lobes which are free from central awn; lemma
backs glabrous with only the margins fringed with short
hairs from base to middle, this fringe terminating in a mar-
ginal tuft of white hairs; central awn 13-18 mm long,
geniculate close to the base, twisted basally, the apical
portion much longer than base and protruding from spike-
lets for some length.

Flowering December to April. Xeric alpine grasslands
and crevices in basaltic cliffs of the Drakensberg above
2000 m. Common (in alpine grassland). Biome: Afromont-
ane. Endemic. Anatomically and morphologically very
similar to M. drakensbergensis (Ellis 1981a). which has
smaller floral parts (especially awns), usually more florets
and a different pattern of hair tufts on the back of the lem-
mas.

Description: Anderson 1960 Bothalia 7: (419). Voucher:
Killick 2349. PRECIS code 9902043-01300.

Merxmuellera stricta (Schrad.) Conert

PI. 121.

(=Danthonia stricta
Schrad.) 1 .

Perennial; tufted; 300-800
mm tall. Leaf blades 100-450
mm long; to 0.5 mm wide. Spike-
lets to 23 mm long (including
awns); to 10 mm wide. Panicle
loosely contracted, interrupted,

30-130 mm long; spikelets 5-7-flowered; glumes 11-22
mm long, 3-7-nerved; lemmas 6-9 mm long, including
3. 5-5. 5 mm long lobes which terminate in a short, soft
bristle; lemma backs with 4, sometimes more, occasionally
indistinct tufts of white hairs along each margin near the
base; central awn 6— 1 2(— 1 7) mm long.

Flowering August to March. A variety of habitats. Com-
mon (in Fynbos and Renosterbosveld veld types). Biome:
Fynbos, Nama-Karoo, and Afromontane. Endemic. This
taxon consists of two anatomical forms (Ellis 1980a). The
‘Drakensberg form’ has dense tufts of long hairs (to 3.5
mm) on the lemmas and longer glumes (15-22 mm) which
are often partially dark brown. The ‘typical form' occuring
from Cape Town to the eastern Cape, has tufts of shorter
hairs on the lemmas, shorter glumes (1 1-16 mm) which are
straw-coloured. The Drakensberg form also appears to
flower later, from December to January.

Description: Stapf 1898-1900 (528), Chippindall 1955
(247). Voucher: Ellis 2242. PRECIS code 9902043-01400.

Merxmuellera sp. (= Ellis 5500)

Perennial; shortly rhizomat-
ous; 730-1000 mm tall. Leaf
blades to 150 mm long; about 1
mm wide. Spikelets 12-15 mm
long (excluding awns); 4-6 mm
wide (excluding awns). Culm
bases bulbous, glabrous, straw-
coloured; leaf blades basal, short
and pungent; panicle contracted.

shortly branched, to 150 mm long; spikelets 3(— 4)-
flowered; glumes 9-13 mm long, 1-nerved; lemmas
8.5-1 1 .0 mm long including 5-7 mm long lobes attenuating
into a long (to 5 mm) bristle; lemma backs glabrous except
for one tuft of white hairs on each margin halfway up lem-
ma body; central awn 1 1-15 mm long, geniculate near base.

Flowering November. Seeps and streambanks. Rare.
Biome: Fynbos. Endemic. Ellis (pers. comm.) considers this
species to be anatomically similar to M. decora.

Voucher: Ellis 5500. PRECIS code 9902043-99999.

Microchloa R.Br.

Micropogon Pfeiffer.

Annual (rarely), or perennial; caespitose (low), or
decumbent (mat-forming). Culms 50-600 mm high; herba-
ceous; unbranched above. Ligule a fringed membrane
(narrow), or a fringe of hairs.

Inflorescence a single spike (slender, often curved)',
espatheate (but often embraced by the uppermost sheath).
Spikelet-bearing axes persistent (tough, narrow).

Spikelets solitary; biseriate; 1.7-5. 5 mm long; com-
pressed dorsiventrally, disarticulating above the glumes.
Glumes two; more or less equal; long relative to the adja-
cent lemmas (exceeding the floret); awnless; very dissimilar

Fig. 135. Microchloa caffra

219

(the lower asymmetric, cymbiform, keeled, twisted at the
base; the upper flat). All florets female-fertile; proximal in-
complete florets absent.

Female-fertile florets 1 . Lemmas without a germination
flap; 2 nerved ; entire, or incised; awnless; or mucronate.
Palea present. Lodicules 2; fleshy; glabrous. Stamens 3
(anthers relatively long). Ovary glabrous. Fruit small
(0.9-1. 5 mm); ellipsoid; hilum short; pericarp fused;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven, or even. PCR sheath
extensions absent. PCR cell chloroplasts centrifugal/
peripheral (usually), or centripetal (in some individuals of
M. caffraT).

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 4
species. 3 in Africa, 1 pantropical. Mesophytic to xero-
phytic; in open habitats (savanna, in shallow hard soils);
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 3
indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

1(0). Plants annual, loosely tufted; leaves usually cauline,
old leaf sheaths not breaking into fibres; anthers

0.3-0. 7 mm long M. indica

Plants perennial, densely tufted; leaves usually basal,
old leaf sheaths splitting into a tuft of dense fibres;

anthers 0. 5-2.0 mm long 2

2(1). Spikelets 2. 5-4.0 mm long; anthers 0.5-1. 2 mm long;
spikes rarely more than 1 mm wide . . M. kunthii
Spikelets 3. 0-5. 5 mm long; anthers 1.2-2. 0 mm long;
spikes 1-2 mm wide M. caffra

Microchloa caffra Nees

Fig. 135. PI. 122.

Elsgras, pincushion grass.

Perennial; densely tufted ( with
most leaves basal); 100-500 mm
tall. Leaf blades 20-100 mm
long. Spikelets 3.0-5. 5 mm long.

Old leaf sheaths splitting into
fibres; spike 40-150 mm long,

1-2 mm wide; anthers 1. 2-2.0
mm long; caryopsis terete, over 1 mm long.

Flowering October to April. Shallow soils on rocky
outcrops. Common. Biome: Savanna, Grassland, and
Nama-Karoo. Africa south of the equator. The
circumscription of this species is not clear-cut. It
intergrades with M. kunthii, which generally has shorter
spikelets and anthers.

Description: Hitchcock & Chase 1950 (636), Chippin-
dall 1955 (203), Clayton et al. 1970-1982 (316).
Illustration: Chippindall 1955 (fig. 179). Voucher: Smook
4450. PRECIS code 9902940-00100.

Microchloa indica (L. f.) Beauv.

(=M. setacea R. Br.) 1.

Annual; loosely tufted (with
leaves usually cauline); 90-200
mm tall. Leaf blades 10-80 mm
long; 0.3-1. 8 mm wide. Spikelets
1. 7-2.9 mm long. Old leaf
sheaths not splitting into fibres;
anthers 0. 3-0.7 mm long;
caryopsis dorsally compressed, 1

Flowering January to May. In semi-shade on bare hard
ground. Rare (in South Africa). Biome: Savanna. Africa
south of the Sahara and Mexico. Very similar to M. kunthii
in all characters except its annual habit.

Description: Stapf 1898-1900 (636), Chippindall 1955
(204), Clayton et al. 1970-1982 (314). Voucher: Volk 1013.
PRECIS code 9902940-00200.

Microchloa kunthii Desv.

Elsgras, pincushion grass.

Perennial (growing in com-
pact mats); densely tufted (with
most leaves basal); 100-430 mm
tall. Leaf blades 10-80 mm long.

Spikelets 2. 5-4.0 mm long. Basal
sheaths splitting into fibres;
spikes 20—1 50(— 250) mm long,
rarely more than 1 mm wide; anthers 0. 5-1.2 mm long;
caryopsis 1.5 mm long.

Flowering November to April. Shallow soil on rocky
outcrops, open sandy patches or sometimes even on
waterlogged, clayey soil. Infrequent. Biome: Savanna and
Grassland. Africa south of the Sahara. Intergrades with M.
caffra, which generally has longer spikelets and anthers.

Description: Clayton et al. 1970-1982 (314).

Illustration: Clayton et al. 1970-1982 (fig. 88). Voucher:
De Winter & Codd 557. PRECIS code 9902940-00300.

Microlaena R.Br.

Sometimes included in Ehrharta.

Perennial; long-stoloniferous and caespitose. Culms
300-2000 mm high; woody and persistent, or herbaceous.
Leaf blades linear to linear-lanceolate; flat (or concave).
Ligule an unfringed membrane to a fringed membrane {a
hyaline rim, with caducous cilia). Plants bisexual, with
bisexual spikelets. The spikelets all alike in sexuality (but
often cleistogamous, leading to reduced paleas and
lodicules, and indehiscent stamens ).

Inflorescence a single raceme, or paniculate', espathe-
ate. Spikelet-bearing axes persistent.

Spikelets compressed laterally; disarticulating above the
glumes', with a distinctly elongated rachilla internode above
the glumes (i.e., beneath the empty lemmas). Hairy callus
present. Glumes two; minute', very unequal (G2 longer); de-
cidedly shorter than the adjacent lemmas; awnless; similar
(membranous). Proximal incomplete florets 2 (similar);
epaleate; sterile. The proximal lemmas awned (long
acuminate, tapered into long slender awns).

Female-fertile florets 1. Lemmas 5-7 nerved; entire;
awnless, or mucronate, or awned (tapered into the stout
awn). Awns 1; median. Awns apical; non-geniculate; much
shorter than the body of the lemma. Palea present (usually);
when present relatively long, or conspicuous but relatively
short, or very reduced; 1 -nerved (or nerveless). Lodicules
2; membranous; glabrous. Stamens 2-6. Ovary glabrous.
Fruit medium sized; oblong-linear; hilum long-linear;
embryo small.

Transverse section of leaf blade. Mesophyll without arm
cells; without fusoids. Midrib with one bundle only.

Cytology, classification, distribution. Chromosome base
number, x = 10. Bambusoideae; Oryzodae; Ehrharteae. 10
species. Philippines, Java to Australasia. Helophytic to
mesophytic; in shade and in open habitats. Natal. 1
naturalized species.

References. 1. Willemse, L.P.M. 1982. Blumea 28:
181-194.

Species treatment by G.E. Gibbs Russell.

220

Microlaena stipoides (Labill.) R. Br.

. • . ,, PI. 123. PI. 124.

Slender perennial; weakly
tufted; 300-500 mm tall. Leaf
blades 40-150 mm long; 1. 5-3.0
mm wide. Spikelets 20-30 mm
long (including stipe and awn).

Sterile lemmas subtended by a
long stipe with a fine tuft of hairs
at its base, lemma tips drawn out
into a long scabrous awn; stamens
usually 4.

Semi-shade in forests. Rare. Naturalized from
Australasia. Malesia, New Zealand and Australia. Recently
Microlaena has been united with Ehrharta (Willemse
1982), on the grounds that the characters supporting four
genera in Ehrharteae are too variable within the genera to
be used for generic differences.

Voucher: Gordon-Gray s.n. PRECIS code 9901610-
00300.

Microstegium Nees

Coelarthron Hook.f ..Ephebopogon Steud .,Leptatherum
Nees , Nemastachys Steud ., Psilopogon Hochst.

Annual, or perennial (rambling); decumbent. Culms
300-600 mm high; herbaceous; unbranched above. Leaf
blades linear to lanceolate; flat. Ligule an unfringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets homomorphic.

Inflorescence of spike-like main branches (flexuous,
fragile racemes, these not villous)', digitate or subdigitate
(usually digitate), or non-digitate (sometimes scattered on
a short axis); espatheate; not comprising ‘partial inflores-
cences’ and foliar organs. Spikelet-bearing axes with very
slender rachides; disarticulating at the joints.

Spikelets in pairs; consistently in Mong-and-short' com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets hermaphrodite. Female-fertile spikelets com-
pressed dorsiventrally; falling with the glumes (the
pedicellate spikelet falling from its pedicel, the sessile
falling with the adjacent intemode and pedicel). Glumes
two; more or less equal; awned, or awnless; very dissimilar
(lower bicarinate, chanelled; upper laterally compressed,
naviculate). Lower glume sulcate on the back. Proximal in-
complete florets 1 (often very reduced); epaleate, or paleate,
palea reduced; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline or membranous); incised; awned. Awns 1;
median; from the sinus (usually); geniculate; much longer
than the body of the lemma. Palea present, or absent; rela-
tively long (always small, but sometimes exceeding the
body of the L2), or conspicuous but relatively short to very
reduced. Lodicules 2; fleshy; glabrous. Stamens 2-3. Ovary
glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. About 15 species. Tropical and
subtropical Africa and Asia. Mesophytic; in shade;
glycophytic. Transvaal, Natal, and Cape Province. 1
indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Microstegium nudum (Trin.) A. Camus

Fig. 136. PI. 125.

(=M. capense (Hochst.) A.

Camus) 1.

Trailing annual (forming
tangled mats); to 600 mm tall.

Leaf blades to 80 mm long; 2-7
mm wide. Spikelets 3.5 — 4.5 mm
long (sessile and pedicellate
alike). Inflorescence of 3^1
slender racemes, solitary or paired on central axis; lower
glume of sessile spikelets concave on back.

Flowering January to May. Moist shady places in
forests. Infrequent. Biome: Forest. Tropical Africa east to
Japan and Australia.

Description: C'hippindall 1955 (484), Clayton et al.
1970-1982 (717). Illustration: Chippindall 1955 (fig. 396).
Voucher: Fisher 131. PRECIS code 9900550-00100.

221

Miscanthus Anderss.

Sometimes included in Miscanthidium Stapf.

Perennial; sometimes long-rhizomatous. Culms
1000-4000 mm high', herbaceous (erect); unbranched
above. Leaf blades linear; flat (or terete). Ligule an
unfringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets homomorphic.

Inflorescence of spike-like main branches, or paniculate
(the panicle often large, branched, silky, red or brown: the
central axis longer and the racemes shorter in the species
previously assigned to Miscanthidium)', open, or contracted;
espatheate; not comprising ‘partial inflorescences’ and
foliar organs. Spikelet-bearing axes ‘racemes’ (slender,
flexuous); with very slender rachides; disarticulating at the
joints (but tardily).

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; unequally pedicellate in each combination.
Pedicels free of the rachis. The short-pedicellate spikelets
hermaphrodite. The long-pedicellate spikelets hermaphro-
dite. Female-fertile spikelets compressed dorsiventrally;
falling with the glumes (disarticulating from the pedicels
before break-up of the rachis). Glumes two; more or less
equal; awnless; very dissimilar (papery to leathery: G1 flat-
backed, 2-keeled with inflexed margins and nerves between
the keels, G2 naviculate). Proximal incomplete florets /;
epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire (seemingly); awned. Awns 1;
median; apical; geniculate (twisted, slightly bent); about as
long as the body of the lemma to much longer than the body
of the lemma. Lemmas hairy ( marginally ). Palea present
(but small); conspicuous but relatively short. Lodicules 2;
fleshy. Stamens 3. Hilum short; embryo large.

Fig. 137. Miscanthus capensis

Cytology, classification, distribution. Chromosome base
number, x = 15. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 6-7 species. Tropical and
southern Africa. Helophytic; in shade, or in open habitats
(streamsides and forest margins); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
Lesotho, and Cape Province. 2 indigenous species.

Intergeneric hybrids procured with Saccharum.

References. 1. Launert. 1970. FSWA. 2. Clayton &
Renvoize. 1982. FTEA. 3. PRE Herbarium practice,
following Gibbs Russell.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf blades terete, reduced to a midrib . M. junceus
Leaf blades expanded, sometimes folded or
sometimes narrowed to the midrib near base ....

M. capensis

Miscanthus capensis (Nees) Anderss.

Fig. 137. PI. 126.

( =Miscanthidium capense
(Nees) Stapf var. capense) 2;

(= Miscanthidium capense
(Nees) Stapf var. villosa Stapf)

3; (-Miscanthidium sorghum
(Nees) Stapf) 3;

(-Miscanthidium erectum Stent
& C.E. Hubb.) 3.

Ruigtegras, dabagrass.

Perennial (often robust); tufted; to 2400 mm tall. Leaf
blades to 90 mm long; to 16 mm wide (expanded). Spikelets
4-6 mm long.

Flowering November to April. Riverbanks, forest
margins and wet places. Infrequent. Biome: Fynbos, Savan-
na, and Grassland. Southern Africa. All three broad-leaved
species formerly recognized in Miscanthidium, M.
sorghum, M. erectum and M. capense, are combined here
because of great variability in the characters upon which
separation has been attempted, including leaf blade
reduction, leaf blade hairiness, ligule length, and spikelet
length and hairiness. In addition, these characters
apparently do not correlate with major habitat differences
between streamsides and forest margins.

Description: Chippindall 1955 (478). Illustration: Chip-
pindall 1955 (fig. 393). Voucher: Moll 1667. PRECIS code
9900380-00100.

Miscanthus junceus (Stapf) Pilg.

(= Miscanthidium junceum
Stapf) 3; (= Miscanthidium
teretifolium (Stapf) Stapf) 1, 2.

Besemgras, ruigtegras.

Perennial; tufted; 1000-18C
mm tall. Leaf blades 500-10C
mm long; to 3 mm wide (terete).

Spikelets 4-5 mm long.

Flowering November to June. Riverbanks and vleis. In-
frequent. Biome: Savanna and Grassland. Southern tropical
Africa.

Description: Chippindall 1955 (480). Voucher: Edwards
2053. PRECIS code 9900380-00500.

222

Monelytrum Hack.

Annual, or perennial; long-stoloniferous (each ‘stolon’
being a single, bare internode), or caespitose, or decumbent.
Culms 80-800 mm high; herbaceous; branched above, or
unbranched above. Leaf blades 2-7 mm wide (their margins
thickened, with tubercle-based hairs); somewhat cordate.,
flat, or rolled (convolute). Ligule a fringed membrane. The
spikelets of sexually distinct forms on the same plant (there
being 1-3 sterile spikelets at the tips of the reduced
inflorescence branches).

Inflorescence bristly, a false spike, with clusters of
spikelets on reduced axes ; espatheate. Spikelet-bearing axes
disarticulating; falling entire (i.e., the clusters shed).

Female-fertile spikelets solitary; 3-4 mm long; com-
pressed dorsiventrally; falling with the glumes (with the
glomerules). Hairy callus present (at base of cluster).
Glumes one per spikelet (G 1 sometimes absent), or two; rel-
atively large (G2); very unequal; long relative to the adja-
cent lemmas (i.e., G2); awned (G2 with an awn at least as
long as itself); very dissimilar (G1 reduced to a minute
scale, G2 flat, elliptic-lanceolate, herbaceous). All florets
female-fertile; proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas less firm than the
glumes (membranous); without a germination flap; 3
nerved; entire to incised; mucronate to awned (from the
mid-nerve). Awns when present 1 ; median; apical; non-gen-
iculate; much shorter than the body of the lemma. Palea
present (broadly lanceolate); relatively long. Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit small (2
mm long); ellipsoid; hilum short (the hilum elliptical); peri-
carp fused; embryo large (about 1/3 the length of the fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Fig. 138. Monelytrum luederitzianum

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Southwest Africa to
southern Angola. Xerophytic; in open habitats (in
seasonally moist locations?); glycophytic. Namibia. 1
indigenous species.

References. 1. Launert. 1970. FSWA.

Species treatment by G.E. Gibbs Russell.

Monelytrum luederitzianum Hack.

Fig. 138. PI. 127.

(=M. annuum Goossens) 1.

Perennial, or annual; stolon-
iferous, or tufted; to 800 mm tall.

Leaf blades to 500 mm long; to 8
mm wide. Spikelets 3-4 mm long.

Pedicels and spikelet bases
woolly; upper glume and lemma
awned; bisexual floret one.

Flowering December to June. Rocky hillslopes and often
by ephemeral water on sandy or calcareous soils. Infre-
quent. Biome: Savanna and Nama-Karoo. Into southern
Angola.

Description: Chippindall 1955 (1 10). Illustration: Chip-
pindall 1955 (fig. 83). Voucher: De Winter 2558. PRECIS
code 9902750-00200.

Monocymbium Stapf

Perennial; caespitose. Culms 300-1200 mm high; herba-
ceous; branched above, or unbranched above. Leaf blades
linear; flat (tapering to a sharp point). Ligule an unfringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets of sexually distinct forms on the same plant ;
overtly heteromorphic (the pedicellate spikelets awnless);
all in heterogamous combinations.

Inflorescence paniculate ( the 'racemes’ loosely
gathered into a false panicle)', spatheate', a complex of
‘partial inflorescences’ and intervening foliar organs.
Spikelet-bearing axes ‘racemes’ (with at least 6 spikelet
pairs); solitary; with very slender rachides (filiform); disar-
ticulating at the joints.

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, similar in form to the sessile, except
that they are awnless. Female-fertile spikelets compressed
dorsiventrally (flattened dorsally, the sides rounded);
falling with the glumes (deciduous with the adjacent joint
and pedicel). Glumes two; more or less equal; awned (G2,
from a notch); very dissimilar (thinly cartilaginous: G2
awned). Lower glume not two-keeled (naviculate, laterally
compressed and keeled over upper 1 13). Proximal incom-
plete florets 1\ epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (except for the cartilaginous median zone); incised;
awned. Awns 1; median; from the sinus; geniculate; much
longer than the body of the lemma. Palea absent. Lodicules
2; fleshy; glabrous. Stamens 3. Ovary glabrous. Hilum
short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5, or 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 4 species. Tropical and
southern Africa. Mesophytic; in open habitats (savanna);
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 1 in-
digenous species.

223

Fig. 139. Monocymbium ceresiiforme

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

Monocymbium ceresiiforme (Nees) Stapf

Wild oatgrass, wildehawer-
gras.

Graceful perennial; sometimes
shortly rhizomatous and tufted
(loosely or densely); 300-1000
mm tall. Leaf blades 50-1 80mm
long; 2-6 mm wide. Spikelets
(sessile and pedicellate) 3.5^t.O
mm long. Plant reddish or purple-tinged when flowering;
racemes solitary, partly enclosed in the reddish-brown boat-
shaped spatheole.

Flowering January to June. Open grassland and hillsides,
often in wet places. Common. Biome: Savanna and Grass-
land. Tropical Africa. Indicator (acid soils). Some
Hyparrhenia species have similar ovate reddish spatheoles,
but they all are larger plants and have paired racemes.
Schizachyrium spp. and Andropogon fastigiatus also have
solitary spatheate racemes, but in Schizachyrium the
spatheoles are narrow, and A. fastigiatus is annual.

Description: Chippindall 1955 (515), Clayton et al.
1970-1982 (825). Illustration: Chippindall 1955 (fig. 411),
Clayton et al. 1970-1982 (fig. 190). Voucher: Ward 6477.
PRECIS code 9900750-00100.

Mosdenia Stent

Perennial; long-stoloniferous (the stolons with densely
imbricate cataphylls). Culms 100-900 mm high; herba-
ceous; unbranched above. Leaf blades linear to linear-
lanceolate. Ligule an unfringed membrane (laciniate). The
spikelets of sexually distinct forms on the same plant (those
at the tip of the inflorescence sometimes reduced), or all
alike in sexuality.

Inflorescence a single spike (dense, continuous,
elongated, the spikelets spreading at right angles to the
axis)-, espatheate. Spikelet-bearing axes persistent.

224

Female-fertile spike lets solitary; not two-ranked (in
whorls or spirals)', not in distinct ‘long-and-short’ combina-
tions; 2.5-3.75 mm long (sub-falcate); falling with the
glumes. Glumes two; more or less equal (G1 slightly longer
and broader); about equalling the spikelets (or somewhat
longer); awnless', very dissimilar (G2 flat-backed). All
florets female-fertile; proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes; without a germination flap; 1 nerved, or 3 nerved;
entire; awnless. Palea present; relatively long. Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit small
(about 1.5 mm long); ellipsoid; hilum short (this elliptical);
pericarp fused; embryo large (about 1/3 the length of the
fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae.se/Jsw lato. 1 species. South Africa. Mesophytic;
in open habitats (dry savanna); glycophytic. Transvaal. 1 in-
digenous species.

References. 1. Clayton. 1971. Kew Bull. 25: 250.

Species treatment by G.E. Gibbs Russell.

Mosdenia leptostachys (Fical. & Hiern) Clayton

Fig. 140. PI. 129.

(=M. phleoides (Hack.)

Stent) 1.

Perennial; rhizomatous (rhi-
zome creeping); to 900 mm tall.

Leaf blades 20-80 mm long; 2-3
mm wide. Spikelets 2.50-3.75
mm long. Inflorescence narrowly
spikelike; spikelets awnless, not
clustered; glumes glabrous, 1 -nerved; bisexual floret one.

Flowering January to April. Bushveld, usually on sandy
soil. Infrequent. Biome: Savanna. Endemic.

Description: Chippindall 1955 (108). Illustration: Chip-
pindall 1955 (fig. 81). Voucher: Codd 825. PRECIS code
9902741-00100.

Nassella Desv.

Sometimes included in Stipa p.p.

Perennial; caespitose. Culms 250-650 mm high; herba-
ceous. Ligule an unfringed membrane.

Inflorescence paniculate; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets 1-3 mm long; compressed laterally (but plump
and gibbous)', disarticulating above the glumes. Glumes
two; more or less equal; much exceeding the spikelets;
awned (acuminate into an awn), or awnless; similar. All
florets female-fertile; proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas saccate (above); decid-
edly firmer than the glumes; hairy, or hairless; without a
germination flap; 3 nerved (obscurely); entire; awned. Awns
1; located asymmetrically, dorsal; geniculate; much longer
than the body of the lemma. Palea present; conspicuous but
relatively short; nerveless. Lodicules 2 (in material seen);
fleshy (‘stipoid’); glabrous. Stamens 3. Ovary glabrous.
Fruit small; oblong to pyriform; hilum long-linear; pericarp
fused; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Stipeae. 15 species. Andes. Mesophytic to xerophytic; in
open habitats; glycophytic. Cape Province. 1 naturalized
species.

References. 1. Caro. 1966. Kurtziana. 3: 79.2. Clayton
& Renvoize. 1986. Gen. Gram.

Species treatment by G.E. Gibbs Russell.

Nassella trichotoma (Nees) Hack, ex Arech.

Fig. 141. PI. 130.

( =Stipa trichotoma Nees) 2.

Nassella tussock, serrated
tussock.

Perennial; densely tufted;

250-650 mm tall. Leaf blades
1 50-450 mm long; 0.25-0.50 mm
wide (setaceous). Spikelets 6.0-

225

8.5 mm long (excluding awns to 35 mm long). Glumes
swollen around floret at base; floret asymmetrical, upper
end rounded, awn not centrally placed.

Flowering August to January. Mountain grasslands and
disturbed places. Locally dominant. Naturalized and inva-
der (very serious) from South America. Southern
hemisphere. Declared weed. This species is sometimes
classified in Stipa and is very similar vegetatively to S.
tenuissima but is distinguished by its asymmetric floret with
the awn arising from one side.

Description; Wells 1986 Mem. Bot. Surv. S. Afr.
(53:502). Illustration: Henderson & Anderson Mem. Bot.
Surv. S. Afr. (37:40). Voucher: Theron 1856. PRECIS code
9902650-00100.

Odontelytrum Hack.

Perennial; long-stoloniferous. Culms 600-1000 mm
high (standing 300-400 mm above the water); herbaceous;
branched above. Leaf blades linear, or linear-lanceolate;
flat, or rolled. Ligule an unfringed membrane to a fringed
membrane. The spikelets all alike in sexuality.

Inflorescence a false spike, with clusters of spikelets on
reduced axes, or a single raceme (a coarse, cylindrical
'raceme' , apparently representing a raceme of reduced
‘ glomerules’ , each glomerule shortly pedunculate,
comprising a single spikelet subtended by a lobed scale
forming an involucre-plus-bristle)', espatheate (but
enveloped below by the uppermost leaf sheath, whose blade
is at least as long as the inflorescence). Spikelet-bearing
axes disarticulating; falling entire (i.e., the reduced
‘glomerules’ deciduous — the main axis persistent).

Spikelets associated with bractiform involucres and (at
least some of them) subtended by solitary ‘bristles’ (each
spikelet with a purplish, irregularly 4-6 lobed involucre.

this being herbaceous except for one lobe, which is almost
free, awnlike, scabrid and 12-25 mm long); solitary.
Spikelets 10-14 mm long; abaxial; compressed dorsiven-
trally; falling with the glumes. Glumes one per spikelet (G 1
missing), or two; very unequal (G 1 when present very
small); awnless. Proximal incomplete florets 1; paleate,
palea fully developed (as long as the lemma); male.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (cartilaginous below, herbaceous above);
smooth; not becoming indurated; hairless; having the
margins lying flat and exposed on the palea; without a ger-
mination flap; 7 nerved; entire; awnless (the tip caudate,
membranous). Palea present; relatively long (equalling the
lemma). Stamens 3. Embryo large (about 1/3 the length of
the fruit).

Photosynthetic pathway. C4; XyMS+. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Panicoideae; Pani:
codae; Paniceae. 1 species ( O . abyssinicum). Abyssinia and
southern Africa. Helophytic (in flowing or standing water);
in open habitats; glycophytic. Transvaal and Orange Free
State. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

Odontelytrum abyssinicum Hack.

Fig. 142. PL 131.

Perennial; hydrophyte; 600-
1000 mm tall. Leaf blades 100-
200 mm long; 7 mm wide. Spike-
lets to 12 mm long; 3 mm wide.

Culms soft, spongy; inflorescence
a raceme, embraced below by the
uppermost leaf sheath; spikelets
solitary, subtended by a lobed
herbaceous scale, with one lobe free and awnlike, to 20 mm
long.

Flowering December to February. In stagnant and
running water. Rare. Highlands of eastern Africa.

Description: Du Toit, Bothalia 12, 2 (258). Illustration:
Clayton et al. 1970-1982 (fig. 154). Voucher: P.V.C. du
Toit 1083. PRECIS code 9901430-00100.

Odyssea Stapf

Perennial (glaucous); long-rhizomatous (sand binding).
Culms 50-750 mm high (and creeping); herbaceous',
branched above, or unbranched above. Plants conspicu-
ously armed ( leaf blades short, rigid and very pungent-
tipped). Leaf blades flat and rolled (inrolled from the flat
base); hard, woody, needle-like . Ligule a fringe of hairs.

Inflorescence paniculate; contracted (fairly to very);
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 5-9 mm long; compressed laterally;
disarticulating above the glumes; disarticulating between
the florets. Glumes two; very unequal; markedly shorter
than the spikelets; awnless (but sometimes with the nerve
tip constituting a tiny mucro); similar (thinly membranous
to hyaline). Incomplete florets distal to the female-fertile
florets, merely underdeveloped; proximal incomplete
florets absent.

Female-fertile florets 2-8. Lemmas similar in texture to
the glumes to decidedly firmer than the glumes
(membranous with scarious margins, or scarious); without
a germination flap; 3 nerved; incised; mucronate. Palea
present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3 (the anthers long). Ovary glabrous. Fruit small
(1.1-1. 5 mm); ellipsoid; hilum short; pericarp free; embryo
large (around 1/3 grain length).

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

226

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Coastal Red Sea, tropical
and southern Africa. Xerophytic; in open habitats;
halophytic. Namibia, Botswana, Transvaal, and Cape
Province. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Odyssea paucinervis (Nees) Stapf

( =Diplachne cinerea
Hack.) 1.

Steekriet, prickly brack grass.

Mat-forming perennial; rhizo-
matous (with dense tufts of spiny
glaucous shoots at the nodes);

100-750 mm tall. Leaf blades
10-60 mm long; 1-5 mm wide. Spikelets 5-9 mm long.
Rhizomes very long, well developed, deeply buried; panicle
15-70 mm long; spikelets fewer than 15, 4-9-flowered.

Flowering October to May. Brackish or saline soil, in
or near pans or rivers. Locally common. Biome: Savanna,
Nama-Karoo, Succulent Karoo, and Desert. Tropical Africa
south of the Congo River. Natural pasture (eaten by stock
because of salty deposits on leaves). Tufts tend to grow in
rows, due to the long rhizomes.

Description: Chippindall 1955 (118), Clayton et al.
1970-1982 (288). Illustration: Chippindall 1955 (fig. 89),
Clayton et al. 1970-1982 (fig. 79). Voucher: Acocks 15602.
PRECIS code 9903451-00100.

Fig. 143. PI. 132.

Olyra L.

Mapira Adans.

Perennial. Culms 500-5000 mm high; woody and
persistent; scandent (twining), or not scandent; branched
above. Leaf blades ovate; pseudopetiolate\ disarticulating
from the sheaths. Ligule an unfringed membrane. Plants
monoecious with all the fertile spikelets unisexual (the male
spikelets immediately beneath the female, or the lower parts
of the panicle exclusively male). The spikelets of sexually
distinct forms on the same plant.

Inflorescence paniculate; spatheate, or espatheate (?).
Spikelet-bearing axes persistent. The male spikelets with 3
free stamens. Female-fertile spikelets 5-10 mm long; com-
pressed dorsiventrally; falling with the glumes (?). Glumes
one per spikelet; long relative to the adjacent lemmas;
awnless, or awned (often caudate-acuminate). Proximal in-
complete florets 1; sterile.

Female-fertile florets 1. Lemmas becoming indurated;
entire; awnless. Palea present; relatively long; 2-nerved.
Lodicules 3. Stamens 0. Ovary glabrous, or hairy. Hilum
long-linear; embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells; with fusoids. Midrib with one bundle only, or with a
conventional arc of bundles, or vascularization complex.

Cytology, classification, distribution. Bambusoideae;
Oryzodae; Olyreae. 23 species. Tropical America, Africa.
Mesophytic; in shade (of forests); glycophytic. Natal and
Cape Province. 1 species, indigenous or possibly
naturalized.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Olyra latifolia L.

Fig. 144. PI. 133.

Perennial (bamboo-like);
scrambler (erect or straggling);

900-3000 mm tall. Leaf blades to
170 mm long; 25-70 mm wide
(flat, pseudopetiolate, broadly
lanceolate, cross-veins visible).

Spikelets (female-fertile) 7-10
mm long (excluding awns, the
male spikelets smaller). Inflores-
cence a scanty whitish panicle.

Flowering December to May. Wet forests, climbing over
shrubs. Rare. Locally common. Possibly naturalized from

227

tropical America. Biome: Forest. Tropical America, Africa
and Madagascar.

Description: Chippindall 1955 (453), Clayton et al.
1970-1982 (17). Illustration: Chippindall 1955 (fig. 376),
Clayton et al. 1970-1982 (fig. 6). Voucher: Smook 5527.
PRECIS code 9901660-00100.

Oplismenus P. Beauv.

Hekaterosachne Steud., Hippagvostis Kuntze,
Orthopogon R. Br.

Annual, or perennial; decumbent. Culms 100-1000 mm
high; herbaceous; freely branched above. Leaf blades linear
to ovate; flat (thin). Ligule a fringed membrane (very short),
or a fringe of hairs.

Inflorescence of spike-like main branches (short
racemes)', open; espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary, paired or in clusters, distant or
approximate; biseriate; not in distinct ‘long-and-short’
combinations', abaxial ; compressed laterally (weakly), or
not noticeably compressed to compressed dorsiventrally;
falling with the glumes. Hairy callus present. Glumes two;
more or less equal; awned (both or at least the lower, awn
of lower always longer, the awns often viscid)', similar
(herbaceous). Proximal incomplete florets 7; paleate, palea
fully developed to reduced; male, or sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes (papery to
coriaceous); smooth (shining); becoming indurated, or not
becoming indurated; hairless (smooth, glossy); having the
margins tucked in onto the palea; with a clear germination
flap; 3-5 nerved; entire; awnless. Palea present; relatively
long. Lodicules 2; fleshy. Stamens 3. Ovary glabrous. Fruit
ellipsoid; hilum short to long-linear (oblong, up to a half
as long as the fruit); embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x = 9, 10, and 1 1. Panicoideae; Panicodae; Pani-
ceae. 5 species. Tropical and subtropical. Mesophytic; in
shade (forest); glycophytic. Namibia, Botswana, Transvaal,
Swaziland, Natal, and Cape Province. 3 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Awns minutely scabrid; plant annual

O. burmannii

Awns smooth, sticky; plant perennial 2

2( 1 ). Spikelets 6-20 per inflorescence, arranged in racemes

O. hirtellus

Spikelets 2-6 per inflorescence, arranged in fascicles
O. undulatifolius

Oplismenus burmannii (Retz.) Beauv.

Prostrate annual; 100-250 mm
tall. Leaf blades 10-60 mm long;

5-20 mm wide. Spikelets 2. 5-3. 5
mm long. Spikelets hairy; glumes
with minutely scabrid awns 3-20
mm long.

Flowering February to April.

In forest shade. Rare (in southern
Africa). Biome: Savanna and For-
est. Tropical Africa, Asia, America.

Description: Clayton et al. 1970-1982 (542). Voucher:
PA. Smith 583. PRECIS code 9901150-00100.

Oplismenus hirtellus (L.) Beauv

Prostrate perennial (some-
times climbing in undergrowth);

150-800 mm tall. Leaf blades
to 130 mm long; 4—20 mm wide.

Spikelets 2-4 mm long. Inflores-
cence of racemes, with 6-20
spikelets; glumes with smooth
sticky awns 3-14 mm long.

Flowering January to June

Fig. 145. PI. 134.

228

Fig. 145. Oplismenus hirtellus

(rarely at other times). In forest shade. Locally common.
Biome: Savanna and Forest. Throughout tropics except
southeastern Asia. Variable in leaf and inflorescence,
intergrading with O. undulatifolius , which has clumped
spikelets. Depauperate specimens may be difficult to place.

Description: Chippindall 1955 (362), Clayton et al.
1970-1982 (543). Illustration: Chippindall 1955 (fig. 313).
Voucher: Liebenberg 8035. PRECIS code 9901 150-00200.

Oplismenus undulatifolius (Ard.) Roem. & Schult.

Trailing perennial; 150-500
mm tall. Leaf blades 10-70 mm
long; 4-15 mm wide. Spikelets
2. 5 — 4.0 mm long. Inflorescence
of 2-6 fascicled spikelets in
wedge-shaped clumps; glumes
with smooth, sticky awns 7-14
mm long.

Flowering January to July. In
forest shade. Locally common. Biome: Savanna and Forest.
Temperate areas in northern hemisphere and upland areas
in Africa. Intergrades with O. hirtellus, which has spikelets
in racemes.

Description: Stapf 1919 (495). Voucher: Davidse 5827.
PRECIS code 9901 150-00300.

Oropetium Trin.

Annual, or perennial; caespitose (dwarf, cushion-
forming). Culms 20— 1 50(— 1 70) mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear;
flat, or folded, or rolled. Ligule an unfringed membrane, or
a fringed membrane.

Inflorescence a single spike (straight, curved, sinuous
or coiled)', espatheate. Spikelet-bearing axes persistent, or
disarticulating; when fragile disarticulating at the joints (or
fracturing into segments of 1—4 spikelets).

Spikelets solitary; distichous; 2.5-3.S mm long', com-
pressed laterally, disarticulating above the glumes, or
falling with the glumes (and with the joint). Glumes two,
or one per spikelet (G 1 sometimes vestigial or missing); rel-
atively large (G2); very unequal (except in terminal
spikelets)', much exceeding the spikelets; awnless; very dis-
similar (G1 reduced and scarious or missing, G2 covering
the florets, hardened). All florets female-fertile, or one
distal incomplete floret present, merely underdeveloped
(male or sterile); proximal incomplete florets absent.

Female-fertile florets 1 (rarely, the second floret also
hermaphrodite?). Lemmas less firm than the glumes
(hyaline); without a germination flap; incised; 3 nerved;
mucronate, or awned. Awns when present 1 ; from the sinus
(or mucronate); non-geniculate; much shorter than the body
of the lemma. Palea present; relatively long (oblong). Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small (about 1.5 mm long); fusiform; hilum short; pericarp
loosely adherent (removable when soaked); embryo small
(about 1/4 the length of the fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 3^1
species. Arid subtropical Africa and mountains. Mesophytic
to xerophytic; in open habitats (in shallow soil between or
over rocks and in outwashes); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Natal, and Cape
Province. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

229

Oropetium capense Stapf

Haasgras, dwarf grass.

Dwarf perennial; densely tuft-
ed; 25-100 mm tall (rarely to 170
mm). Leaf blades 10-40 mm
long; to 1.2 mm wide. Spikelets
2. 5-4.0 mm long. Spikes solitary,
straight or curved; spikelets sunk
into the rachis; upper glume 2-3
mm long.

Flowering December to May. In shallow soil in open
places or rocky outcrops or in crevices and hollows on
exposed rocks, often in badly grazed or disturbed veld. Lo-
cally common. Biome: Savanna, Grassland, and Nama-
Karoo. Eastern tropical Africa to Chad and Somalia.

Description: Stapf 1898-1900 (742), Chippindall 1955
(204), Clayton et al. 1970-1982 (306). Illustration: Chip-
pindall 1955 (fig. 180). Voucher: Van Rooyen 3130.
PRECIS code 9903200-00100.

Fig. 146. PI. 135.

Fig. 146. Oropetium capense

Oryza L.

Padia Moritzi.

Annual, or perennial; long-rhizomatous, or caespitose.
Culms 300-3000 mm high; herbaceous. Leaves usually au-
riculate. Leaf blades flat; pseudopetiolate, or not pseudo-
petiolate. Ligule an unfringed membrane. Plants bisexual,
with bisexual spikelets.

Inflorescence paniculate', espatheate. Spikelet-bearing
axes persistent.

Spikelets 4-12 mm long; compressed laterally; disartic-
ulating above the glumes (if the pedicel cup is interpreted
as glumes). Hairy callus absent. Glumes present to absent
(represented only by a small 2-lobed cupule); if present

two; minute; more or less equal; awnless. Proximal incom-
plete florets 2 (small, vestigial, sometimes only bristles);
epaleate; sterile.

Female-fertile florets 1. Lemmas 3-9 nerved; entire;
awnless, or mucronate, or awned. Awns when present 1.
Awns apical; non-geniculate; much shorter than the body
of the lemma, to much longer than the body of the lemma.
Palea present; relatively long (but narrower than the
lemma); with several nerves. Lodicules 2; membranous (but
the membranous flange may be narrow); glabrous. Stamens
6. Ovary glabrous. Fruit small, or medium sized, or large;
hilum long-linear; embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells; without fusoids. Midrib vascularization complex.

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Oryzodae; Oryzeae to
Olyreae. 25 species. Tropical. Hydrophytic or helophytic;
in shade (wet forests) or open habitats (swamps);
glycophytic. Namibia, Botswana, Transvaal, and
Swaziland. Indigenous species (3), cultivated species (1).

Intergeneric hybrid claimed with Triticum : X Oryticum
Wang & Tang in Acta Phytotax. Sin. 20: 179 (1982).

References. 1. Clayton. 1970. FTEA. 2. Launert. 1971.
FZ. 10(1).

Species treatment by G.E. Gibbs Russell.

1(0). Ligule of lowest leaves longer than 15 mm, apex
acute; plants perennial, with long rhizomes ....

O. longistaminata

Ligule of lowest leaves shorter than 10 mm, truncate
or rounded; plants annual 2

2(1). Spikelets 7-1 1 mm long O. barthii

Spikelets 5-6 mm long O. punctata

Oryza barthii A. Chev.

Robust annual; hydrophyte; to
1 500 mm tall. Leaf blades to 450
mm long; to 15 mm wide. Spike-
lets 7-11 mm long(awns40-160
mm long). Ligule 2-6 mm long,
truncate.

Flowering February to March.

Floodplain pans. Rare. Tropical
Africa.

Description: Clayton et al. 1970-1982 (30). Voucher:
P.A. Smith 1937. PRECIS code 9901580-00050.

Oryza longistaminata A. Chev. & Roehr.

Fig. 147. PI. 136.

(=0. barthii auctt., non A.

Chev.) 1.

Wild rice.

Perennial; hydrophyte and
rhizomatous (rhizomes extensive,
branched); to 1200 mm tall
(culms spongy). Leaf blades to
450 mm long; to 1 5 mm wide. Spikelets 7-9 mm long (awns
40-80 mm long). Ligule 15-45 mm long, acute.

Flowering October to May. Swamps and floodplains,
often in deep water. Locally common. Throughout tropical
Africa and Madagascar. O. sativa, the cultivated rice, is
grown in southern Africa. It is an annual with a long ligule
and the spikelets are awnless. Escapes from cultivation have
not been reported.

Description: Chippindall 1955 (32), Clayton et al.
1970-1982 (30). Illustration: Clayton et al. 1970-1982 (fig.
10(8)). Voucher: Killick & Leistner 3029. PRECIS code
9901580-00100.

230

Fig. 147. Oryza longistaminata

Oryza punctata Steud.

Annual; hydrophyte; 600-
1200 mm tall (culms spongy).

Leaf blades to 300 mm long; to 1 0
mm wide. Spikelets 5-6 mm long
(awns 10-70 mm long). Ligule
3-10 mm long, truncate.

Flowering November to April.

Floodplain pans, rice paddies.

Rare. Tropical Africa and Madagascar, Thailand. Weed (in
rice fields).

Description: Clayton et al. 1970-1982 (31 ). Illustration:
Clayton et al. 1970-1982 (fig. 10). Voucher: Ward 2054.
PRECIS code 9901580-00200.

Oryzidium C.E. Hubb. & Schweick.

Perennial; long-stoloniferous. Culms 400-1200 mm
high (the lower internodes trailing in water or floating); her-
baceous; branched above. Leaf blades linear; flat. Ligule a
fringe of hairs. Plants without hermaphrodite florets (the
lower floret male, the upper female).

Inflorescence paniculate; narrow, the branches nearly
erect; espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 8-10 mm long; compressed dorsiven-
trally; falling with the glumes. Glumes two; very unequal;
awned (upper glume attenuate into a long straight awn);
very dissimilar (the G1 a small, membranous, truncate
scale, the G2 large, firm, awned). Proximal incomplete
florets I : paleate, palea fully developed; male (with 3
stamens).

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (thinly
coriaceous); smooth; not becoming indurated; hairless;
having the margins lying flat and exposed on the palea; with
a clear germination flap; 7 nerved; entire; mucronate (or
mucronulate). Palea present; relatively long. Lodicules 2;
fleshy; glabrous. Stamens 0. Ovary glabrous. Fruit small
(3-3.5 mm); ellipsoid. Hilum short; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional. XyMS+. PCR cell chloroplasts
seemingly centripetal.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 1 species. Southern tropical Africa.
Hydrophytic (in permanent water); in open habitats;
glycophytic. Namibia and Botswana. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA.

Species treatment by H.M. Anderson.

Oryzidium barnardii C.E. Hubb. & Schweick.

Fig. 148. PI. 137.

Perennial; hydrophyte; float-
ing culms to 1200 mm tall. Leaf
blades 150-200 mm long; 6-8
mm wide. Spikelets 8-10 mm
long; 1.5 mm wide. Culms root-
ing and branching at the lower
nodes; leaf sheaths broad, papery
and straw coloured; lower glume
an ovate, white scale 1-2 mm
long; upper glume 8-10 mm long with scabrid awn 10-18
mm long; female-fertile (upper) floret separated from the
lower by a rachilla internode 1 mm long.

Flowering October to May. Pans and dams. Infrequent.
Biome: Savanna. Zambia and Zimbabwe.

Description: Chippindall 1955 (425). Illustration: Chip-
pindall 1955 (fig. 354). Voucher: Smith 1944. PRECIS
code 9901142-00100.

231

Fig. 148. Oryzidium barnardii

Oxyrhachis Pilg.

Perennial; caespitose. Culms 200-800 mm high; herba-
ceous; unbranched above. Leaf blades linear; folded, or
rolled. Ligule a fringed membrane, or a fringe of hairs
(short). Plants bisexual, with bisexual spikelets.

Inflorescence a single spike (narrow, cylindrical,
terminating the culm)', espatheate; not comprising ‘partial
inflorescences’ and foliar organs. Spikelet-bearing axes
cylindrical spikes; solitary; with substantial rachides; disar-
ticulating at the joints. ‘Articles’ without a basal callus-
knob.

Spikelets solitary (or theoretically in pairs, the ‘pedicel’
fused with and indistinguishable from the rachis). Female-

fertile spikelets 4-6 mm long; compressed dorsiventrally;
falling with the glumes (and with the adjacent joint).
Glumes two; more or less equal; awnless; very dissimilar
(G1 obtuse, leathery, G2 apically notched or entire,
membranous-hyaline). Proximal incomplete florets 1\
epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes; entire; awnless. Palea present, or absent; when
present very reduced (adherent to the lodicules). Lodicules
2; fleshy; glabrous. Stamens 3. Ovary glabrous. Hilum
short; embryo large.

Fig. 149. Oxyrhachis gracillima

232

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Rottboelliinae. 1
species. Tropical Africa, Madagascar. Helophytic; in open
habitats (streamsides and marshy places); glycophytic.
Natal and Cape Province (Transkei). 1 indigenous species.
References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Oxyrhachis gracillima (Bak.) C.E. Hubb.

Fig. 149. PI. 138.

Perennial; densely tufted;

200-600 mm tall. Leaf blades
50-300 mm long, filiform. Spike-
lets (sessile) 3-6 mm long (pedi-
cellate spikelets completely ab-
sent, no pedicel). Inflorescence
very slender with sunken spike-
lets; glumes smooth.

Flowering June. Wet places.

Rare and conservation status not known. Tropical Africa.

Description: Clayton et al. 1970-1982 (855).

Illustration: Clayton et al. 1970-1982 (fig. 204). Voucher:
Huntley 791. PRECIS code 9900341-00100.

Oxytenanthera Munro

Houzeaubambus Mattei, Scirpobambus Kuntze.
Sometimes included in Dendrocalamus.

Perennial; caespitose. Culms 3000-13000 mm high
(somewhat crooked, bending over to the ground); woody
and persistent (forming dense clumps). Culms reaching
20-100 mm in diameter. Culms branched above (at the
nodal line). Leaf blades linear-lanceolate to lanceolate; flat;
pseudopetiolate; disarticulating from the sheaths. Ligule an
unfringed membrane. The spikelets of sexually distinct
forms on the same plant (there being numerous sterile
spikelets).

Inflorescence a false spike, with clusters of spikelets on
reduced axes (often reduced to a single terminal cluster );
spatheate (each spikelet cluster subtended by a papery
sheath, and individual female-fertile spikelets by several
short, papery 'bracts’).

Spikelets 1 5 — 45 mm long; associated with bractiform in-
volucres; compressed laterally to not noticeably com-
pressed; falling with the glumes. Glumes two (cross-
veined); very unequal; decidedly shorter than the adjacent
lemmas; awnless; similar (papery to leathery). Proximal in-
complete florets 1-3; male, or sterile (the paleas when
present two-keeled).

Female-fertile florets 1. Lemmas 11-23 nerved (with
cross-nerves); entire; mucronate to awned. Awns 1; median.
Awns apical; non-geniculate; to 7 mm long. Palea present;
relatively long (may exceed the lemma); with several
nerves (16-19). Stamens 6. Ovary glabrous (but the style
mostly shortly hairy); with a conspicuous apical ap-
pendage', the appendage long, stiff and tapering. Stigmas 3.
Fruit large; hilum long-linear; embryo small.

Transverse section of leaf blade. Mesophyll without arm
cells; with fusoids. Midrib vascularization complex.

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Bambusodae; Bambuseae.

1 species. Africa. Mesophytic; in shade (growing in the
protection of larger trees); glycophytic. Transvaal. 1 indige-
nous species.

References. 1. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

Fig. 150. Oxytenanthera abyssinica

Oxytenanthera abyssinica (A. Rich.) Munro

Fig. 150.

Bamboo; rhizomatous; to
10000 mm tall (culms 50-100
mm in diameter). Leaf blades
50-250 mm long; 10-30 mm
wide. Bamboo with drooping
culms and clustered leaf-bearing
branches; sheaths of culm leaves
with dense hairs on the inner
surface.

Flowering unknown in southern Africa. In shade of
larger trees. Rare. Biome: Savanna. Tropical Africa. Do-
mestic use (flutes). Maintained in semi-cultivation by the
Venda.

Voucher: Smook & Soderstrom 1983. PRECIS code
9904770-00100.

233

Panicum L.

Chasea Nieuw., Coleataenia Griseb., DUeucaden (Raf.)
Steud., Eatonia Raf., Eriolytrum Kunth, Milium Adans.,
Monachne P. Beauv., Phanopyrum (Raf.) Nash, Polyneura
Peter, Psilochloa Launert, Setiacis S.L. Chen and Y.X. Jin
(? — original description inadequate).

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 200-4000 mm
high; woody and persistent, or herbaceous; branched above,
or unbranched above. Leaf blades flat (usually); not disar-
ticulating. Ligule an unfringed membrane, or a fringed
membrane to a fringe of hairs. Plants with hermaphrodite
florets.

Inflorescence paniculate ( except in the Stolonifera
group, where it consists of racemes and the dis fiction from
Brachiaria breaks down)\ open, or contracted; espatheate.
Spikelet-bearing axes persistent. Spikelets not secund
(except the American Agrostoidea group, 'Psilochloa' ,
etc.). Pedicel apices cupuliform.

Spikelets not in distinct ‘long-and-short’ combinations;
1.4-6 mm long (narrowly elliptic, usually more or less
acute); adaxial (in the few cases where the orientation is
ascertainable); compressed dor siventr ally (with very few
exceptions: e.g. P. hemitomum)', falling with the glumes, or
not disarticulating. Glumes two; nearly always very
unequal; nearly always awnless (the G2 truncate to pointed,
very rarely shortly awn-tipped); very dissimilar, or similar
(herbaceous-membranous, the lower sometimes very short
and nerveless). Proximal incomplete florets 1 (rarely 2);
paleate, or epaleate, palea when present fully developed to
reduced; male, or sterile. Proximal lemmas less firm than
the female-fertile lemmas.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes; smooth (rarely
rugose: subgenus Megathyrsus (P. maximum)); becoming
indurated, or not becoming indurated (coriaceous, bony or
cartilaginous); hairless; having the margins tucked in onto
the palea; with a clear germination flap; 3-1 1 nerved;
entire; awnless (rarely minutely apiculate). Palea present;
relatively long. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small; hilum short (linear in (e.g.) P.
glutinosum, P. macranthum, P. pilgerianum = Psilochloa );
embryo large.

Photosynthetic pathway. C4, or C3 (with a very few
species intermediate). The anatomical organization when
C4 conventional, or unconventional. Organization of PCR
tissue in a few C4 species Alloteropsis type. Biochemical
type PCK (5 species), or NAD-ME (14 species), or
NADP-ME (4 species); XyMS+ (C3, or C4 NAD-ME or
PCK), or XyMS-. PCR cell chloroplasts centrifugal/
peripheral, or centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 7, 9, and 10. Panicoideae; Panicodae; Paniceae.
About 370 species. Tropical, subtropical and warm
temperate. Mesophytic, or xerophytic; in shade and in open
habitats (diverse habitats); maritime-arenicolous
(occasionally sandbinding — e.g. P. pinifolium), or glyco-
phytic. Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. Indigenous
species (40), naturalized species (1).

References. 1 . Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA. 3. Clayton & Renvoize. 1982. FTEA. 4.
Clayton & Renvoize. 1986. Genera Graminum. 5.
Renvoize. 1989. Kew Bull. 44: 544.

Species treatment by L. Smook.

1(0). Glumes and lemma of the lower floret pectinate at the

apex P. ecklonii

Glumes and lemma of the lower floret entire .... 2
2(1). Inflorescence branches with flat glandular patches .

3

Inflorescence branches eglandular, or if glandular the
patches not flat 4

Fig. 151. Panicum maximum

3(2). Leaves thin; plant usually erect; female-fertile (upper)
lemma densely verruculose . P. heterostachyum
Leaves thick and slightly leathery; plant usually
trailing; female-fertile (upper) lemma smooth and
shiny, sometimes with scattered papillae at the base
and apex P. glandulopaniculatum

234

4(2). Female-fertile lemma conspicuously transversely
rugose (in P obumbratum the female-fertile
(upper) lemma is minutely rugose, but the spikelets

are over 3 mm long) 5

Female-fertile lemma sculpturing not conspicuously
rugose (if minutely rugose, spikelets less than 3 mm

long) 7

5(4). Plants trailing, inflorescence narrow, with a few
spikelets only; female-fertile (upper) lemma

minutely rugose P. obumbratum

Plants usually erect, sometimes geniculate and rooting
at the lower nodes; inflorescence lax or contracted,
with many spikelets; female-fertile (upper) lemma

conspicuously rugose 6

6(5). Spikelets 2.5-3.0(-4.0) mm long, cartilaginous;
inflorescence usually much branched, secondary
branches usually flexible; most of the nerves on the
lemma of the lower floret are clearly visible on the

closed spikelet P. maximum

Spikelets 3. 2-3. 5 mm long, somewhat leathery;
inflorescence sparsely branched because the
secondary branches are usually absent; only the
central nerve on the lemma of the lower floret is
usually clearly visible on the closed spikelet ....

P. infestum

7(4). Palea of lower floret absent or reduced in length,
being conspicuously shorter than the lower lemma;

lower floret always sterile 8

Palea of lower floret well developed, or if reduced
usually reduced in width only (if reduced in length
then the spikelets with lower floret male); lower
floret usually male, occasionally sterile (sometimes

mixed on the same inflorescence) 20

8(7). Upper glume (10-) 1 1-14-nerved; lemma of the lower
floret (9-) 10-1 1 -nerved; inflorescence enclosed in
the two uppermost leaves, not extending beyond the

leaf apex P. gilvum

Upper glume 3-9-nerved; lemma of the lower floret
5-9-nerved; inflorescence well exserted from the
uppermost leaf, or if the base is enclosed then the

inflorescence extends beyond the leaf tip 9

9(8). Spikelets 5.0-6.5(-7.0) mm long P. volutans

Spikelets to 4 mm long 10

10(9). Lower glume as long as the spikelet; plant
scrambling, sometimes rooting at the lower

nodes; spikelets pubescent P. aequinerve

Lower glume to 3/4 the length of the spikelet, or if
as long as the spikelet then the plant not

scrambling and the spikelets glabrous 11

1 1(10). Upper glume 3-5-nerved 12

Upper glume 7-9-nerved 15

12(11). Upper glume 3-nerved; female-fertile (upper)

lemma with a green spot at the apex

P. comorense

Upper glume 5-nerved; female-fertile (upper)
lemma lacking a green spot at the apex .... 13
13(12). Spikelets 1.5-2. 5 mm long; female-fertile (upper)

floret dull and granulose P. laticomum

Spikelets 2. 6-4.0 mm long (occasionally less),
female-fertile (upper) floret shiny and smooth

14

14( 13). Plant shrub-like, culms hard and wiry, erect; lower
glume narrowly ovate, 3-nerved, 3/4 as long as

the spikelet; leaves up to 4.5 mm wide

P. dewinteri

Plant usually soft, decumbent or trailing; lower
glume broadly ovate, 0-1 -nerved, 1/4— 1/2 the
length of the spikelet; leaves (5—) 10— 25 mm wide

P. monticola

15(1 1). Perennial, shrub-like; culms hard, wiry and
branching, particularly in the upper portion . . .

P. dewinteri

Annuals or short-lived perennials; culms not hard
and wiry, rarely branching in the upper portion

16

16( 15). Inflorescence branches with spikelets appressed
and appearing close together 17

Inflorescence branches with spikelets spreading and

therefore distant from each other 18

17(16). Upper leaf surface densely covered with large
prickles which are usually white (always visible

on upper leaves) and minute papillae

P. subalbidum

Upper leaf surface without prickles, densely

covered with papillae P. impeditum

18(16). Spikelets 1.8-2. 2 mm long, often entirely tinged
purple when mature; inflorescence branches
modestly covered with prickles less than 0.05
mm long just below the spikelets; mature
inflorescence well exserted from uppermost leaf

P. atrosanguineum

Spikelets 2-3 mm long, only tinged purple at the
apex of glumes and lemmas; inflorescence
branches densely scabrid with prickles 0. 1 0-0. 1 5
mm long just below the spikelets; base of
inflorescence usually enclosed in the uppermost

leaf 19

19(18). Inflorescence broadly ovate, branches long and

flexible P. novemnerve

Inflorescence obovate, branches usually short and

rigid P. arcurameum

20(7). Lemma of the lower floret 5-nerved 21

Lemma of the lower floret 7-1 l(-14)-nerved . 28
21(20). Spikelets 3. 3-5. 5 mm long, clavellate hairs usually
present on inflorescence branches . P. deustum
Spikelets up to 3 mm long, clavellate hairs absent

22

22(21). Female-fertile floret sparsely to densely granulose

or papillose 23

Female-fertile floret smooth 24

23(22). Lower glume to 1/2 the length of the spikelet; lower
floret palea coriaceous, longer than the lemma;
spikelets oblong, appressed on and hiding the

inflorescence branches P. hians

Lower glume 1/2-3/4 the length of the spikelet;
lower floret palea membranous, never longer
than the lemma; spikelets nearly rounded in
outline, not appressed to inflorescence branches

which are visible between the spikelets

P. natalense

24(22). Spikelets 1. 3-1.8 mm long; glume apex not
recurved or mucronate; inflorescence 10-60 mm

long 25

Spikelets 2-3 mm long; glume apex recurved,
usually shortly mucronate; inflorescence 80-150

mm long 26

25(24). Leaves not reflexed, linear-lanceolate, tapering to
a long acuminate tip, base straight; lower glume

1/3 the length of the spikelet

P. subflabellatum

Leaves often reflexed at maturity, lanceolate to
narrowly ovate, tip acute, base cordate; lower

glume 1/2-2/3 the length of the spikelet

P. parvifolium

26(24). Basal sheaths silky pubescent ... P. dregeanum
Basal sheaths glabrous or sparsely hispid, not silky

pubescent 27

27(26). Culms usually stout, (2.0-)3.5-7.0 mm wide at the

base P. fluviicola

Culms usually slender, 1-2 mm wide at the base

P. genuflexum

28(20). Lower leaf sheaths densely covered with matted

woolly hairs P. lanipes

Lower leaf sheaths glabrous to densely hairy, hairs

not matted and woolly 29

29(28). Lower glume narrowly ovate; clavellate hairs
usually present on inflorescence branches ....

P. hymeniochilum

Lower glume ovate to broadly ovate; clavellate
hairs never present on inflorescence branches .

30

30(29). Lower floret lemma with interspaces between the
nerves broadest adjacent to the central nerve . .
31

235

Lower floret lemma with the broadest interspaces
between nerves not confined to those by the

central nerve 32

31(30). Spikelets 2. 0-3. 5 mm long P. repentellum

Spikelets 4-6 mm long P. pilgerianum

32(30). Rhizomes stout, long and creeping . . . P. repens
Rhizomes absent, or if present, short and compact

33

33(32). Upper glume 5-nerved; spikelets to 1.8 mm long.

spherical P. subflabellatum

Upper glume (5— )7— 1 1 -nerved; spikelets2 mm or

longer, not spherical 34

34(33). Plants annual 35

Plants perennial 38

35(34). Spikelets 4-6 mm long P. pilgerianum

Spikelets 2. 0-3. 5 mm long 36

36(35). Inflorescence with branches appressed and
ascending, and closely associated with and
enclosed on the one side by the upper leaf blade;

lower florets always sterile

P. sp. 1 (=Smook 3463)

Inflorescence with branches open, not appressed,
not closely associated with uppermost leaf blade;
lower florets male or sterile in the same

inflorescence 37

37(36). Inflorescence obovate to broadly obovate; spikelets
acute; upper glume apex usually with a small
mucro-point and slightly recurved backwards;
plants yellowish green . P. sp. 2 (=Giess 8605)
Inflorescence oblanceolate to narrowly obovate,
spikelets blunt; upper glume apex not mucronate

or recurved backwards; plants green

P. schinzii

38(34). Spikelets 3. 4-4. 5 mm long; upper leaf surface
densely covered with short hairs, abaxial surface
at sheath mouth densely covered with long,

woolly hairs P. kalaharense

Spikelets up to 3.2 mm long; upper leaf surface
smooth to densely papillate or with scattered
hairs; abaxial surface of the sheath mouth
glabrous or with short, appressed hairs .... 39
39(38). Culms hard, brittle, branching after the lower 1/4,
nodes thickened and swollen, often bulbous;

plants shrub-like P. arbusculum

Plants not as above 40

40(39). Nodes densely covered with appressed hairs; lower

glume to 1/4 the length of the spikelet

P. trichonode

Nodes glabrous or sparsely hairy; lower glume

1/2-2/3 the length of the spikelet

P. coloratum / P. stapfianum complex

(including also P. bechuanense and P. merkeri)

NOTE: Panicum miliaceum L., a cultivated species
commonly known as ‘proso’, is occasionally
found as an escape. It has a dense, often drooping
inflorescence, spikelets (4.0-)4.5-5.5 mm long
clustered towards the upper parts of the branches
and a lower palea reduced to a small scale.

Panicum aequinerve Nees

Bosbuffelsgras.

Shortlived perennial, or an-
nual; scrambler (trailing, decum-
bent and rooting at the nodes);
culms to 1000 mm long. Leaf
blades 30-110 mm long; 3-10
(-12) mm wide. Spikelets 2. 5-3. 5
(-4.0) mm long. Inflorescence

sparsely branched, usually spreading at maturity, sometim-
es contracted, branches naked for a long distance, with 2-5
spikelets crowded at the apex, exserted from uppermost
leaf; spikelets acuminate, pubescent, lower glume as long
as the spikelet, 5(-7)-nerved; upper glume 7-nerved; lower
floret sterile, the lemma 5-nerved, the palea reduced;
female-fertile (upper) lemma pale and shiny.

Flowering September and January to June. Clay or sand
on shallow soils of forest margins or open grasslands, main-
ly in damp places and around boulders. Infrequent to locally
common. Biome: Grassland and Forest. Northwards to
Uganda, Ethiopia and in Madagascar. Shows much
variation in the inflorescence shape and spikelet size.
Spikelets are often infected with fungi. Similar to P.
inaequilatum Stapf & C.E. Hubb. from Zimbabwe and
Mozambique, which has a 3-nerved lower glume.

Description: Stapf 1898-1900 (399), Chippindall 1955
(324), Clayton et al. 1970-1982 (495). Illustration: Chip-
pindall 1955 (fig. 281). Voucher: Smook 5486; Smook
5634. PRECIS code 9901 160-00100.

Panicum arbusculum Mez

Struikpanicum.

Perennial; tufted (erect); to
800 mm tall. Leaf blades to 100
mm long; to 6 mm wide. Spike-
lets 2. 5-3.0 mm long. Plant
shrub-like, glaucous, culms hard
and brittle; rhizomes short and
strong; culms much branched
with nodes thickened and swollen, often bulbous; inflores-
cence open, sparsely branched, primary branches with long
naked bases and bearing spikelets crowded towards the
apex; spikelets often flushed with purple; lower glume
broadly ovate, up to 1/2 the length of the spikelet; upper
glume 7-9-nerved; lower floret male or sterile, lemma 9-
nerved, palea well developed; female-fertile (upper) lemma
pale-yellow to brown, shiny.

Flowering October to May. Stony places in mountainous
areas. Infrequent to locally common. Biome: Nama-Karoo.
Endemic. Pasture (good grazing), or erosion control
(effective in blocking water drainage).

Description: Muller 1984 (194), Chippindall 1955 (338).
Illustration: Muller 1984 (fig. 96). Voucher: Giess 10356.
PRECIS code 9901 160-00300.

Panicum arcurameum Stapf

Annual; tufted (erect or geni-
culate); to 600 mm tall. Leaf blad-
es to 1 00 mm long; to 7 mm wide.

Spikelets 2. 0-2. 5 mm long. In-
florescence obovate, base usually
enclosed by uppermost leaf,
branches usually short, rigid and
ascending, densely scabrid with
prickles which are 0. 1 0-0. 1 5 mm
long just below point of attachment of the spikelets, which
are spreading and distant from one another; the apex of low-
er glume and lower lemma often tinged purple; lower glume
to 2/3 the length of the spikelet; upper glume and lower
lemma (7-)9-nerved; lower floret sterile with a reduced
palea; female-fertile (upper) lemma pale to dark, shiny.

Flowering January. Sandy soils, black turf in disturbed
areas. Infrequent. Biome: Savanna and Nama-Karoo.
Southern tropical Africa. Barely distinguishable from P.
novemnerve and a detailed study is needed in this group.
Similar to P. atrosanguineum, which has the inflorescence
branches moderately scabrid with prickles less than 0.05
mm long just below the spikelet.

Description: Stapf 1920 (704), Chippindall 1955 (327),
Clayton et al. 1970-1982 (488). Voucher: Smook 4404.
PRECIS code 9901 160-00400.

236

Panicum atrosanguineum A. Rich.

Annual; tufted; 100-400 mm
tall. Leaf blades to 60 mm long;
to 5 mm wide. Spikelets 1.8-2. 2
mm long. Inflorescence open,
mature inflorescences well ex-
serted from the uppermost leaf,
branches moderately scabrid,
with prickles less than 0.05 mm
long just below the point
of attachment of the spikelets, which are distant from one
another; spikelets usually strongly flushed with purple;
lower glume 3/4 the length of the spikelet, broadly ovate;
upper glume and lower lemma 5-7-nerved; lower floret
sterile with a reduced palea; female-fertile (upper) lemma
usually dark at maturity, shiny.

Flowering February to May. Old farmlands and other
disturbed places. Infrequent. Biome: Savanna. Northwards
through Zimbabwe, Zaire to tropical east Africa. Also in
northwest India. Similar to P. novemnerve and P.
arcurameum, which have the inflorescence branches
densely covered with prickles 0.10-0.15 mm long just
below the spikelet attachment.

Description: Stapf 1920 (703), Chippindall 1955 (328),
Clayton et al. 1970-1982 (488). Voucher: Smith 2366.
PRECIS code 9901 160-00450.

Panicum bechuanense Brem. & Oberm.

Perennial; tufted (erect to gen-
iculate); to 600 mm tall. Leaf
blades to 100 mm long; 3-5 mm
wide. Spikelets to 2.4 mm long.

Upper leaf surface densely
papillate; leaves and sheaths with
bulbous-based hairs; lower glume
ovate, 1/2-2/3 the length of the
spikelet; upper glume and lower
lemma 9-nerved; lower floret male, with the palea well
developed; female-fertile (upper) lemma pale to dark,
shiny.

Flowering March. Seepage areas in river beds and pans,
also in disturbed areas. Infrequent. Biome: Savanna.
Endemic. This species, of which only two specimens have
been seen for this treatment, belongs to the P. coloratum-
P. stapfianum complex which needs a more detailed study.
One of these, Ellis 4366, is anatomically different from P.
coloratum.

Description: Bremerkamp & Obermeyer 1935 Ann.
Trans. Mus. 16 (403), Chippindall 1955 (336). Voucher:
Ellis 4366. PRECIS code 9901 160-00500.

Panicum coloratum L. var. coloratum

(-P. coloratum L. var.
makarikariense Goossens) 3.

Witbuffelgras, white buffalo
grass.

Perennial; tufted (erect, geni-
culate or occasionally decum-
bent); to 1000 mm tall. Leaf blad-
es to 300 mm long; 5-10 mm wide. Spikelets 2. 5-3.0 mm
long. Lower leaf sheaths glabrous to densely appressed-
hairy, bulbous-based hairs present or absent; leaves usually
mainly cauline, broad; inflorescence branches spreading
with the spikelets distant; lower glume ovate, 1/2-2/3 the
length of the spikelet; upper glume and lower lemma 7(— 9)-
nerved; lower floret male, with the palea well developed;
female-fertile (upper) lemma pale to dark, shiny.

Flowering October to May. Sandy or clay soils in river
beds, drainage courses, around pans or in depressions.
Common. Biome: Savanna, Grassland, and Nama-Karoo.
Tropical and subtropical Africa, introduced elsewhere.
Palatable and drought-resistant pasture. A highly variable
species belonging to the P . coloratum - P. stapfianum

complex, which also includes P. bechuanense and P.
merkeri. Detailed study is needed to elucidate species
limits. There are many distinct ecotypes, a number of which
have been selected for pastures such as var. makarikariense ,
a tall, robust, glaucous plant.

Description: Stapf 1920 (713), Chippindall & Crook
1976 (46), Stapf 1898-1900 (409), Chippindall 1955 (335),
Clayton et al. 1970-1982 (485). Illustration: Muller 1984
(fig. 97), Chippindall 1955 (fig. 291 ). Voucher: Merxmuell-
er & Giess 30162, Theron
9901 160-00800.

Panicum comorense Mez

Annual; tufted (trailing, de-
cumbent or erect and rooting at
the nodes); culms to 1000 mm
long, or sometimes longer. Leaf
blades 60-150 mm long; 10-15
mm wide. Spikelets 1.8-2. 2 mm
long. Leaf apex abruptly acumin-
ate; inflorescence sparsely bran-
ched, the secondary branches
appressed; spikelets oblong, blunt; lower glume 1/4— 1/3 as
long as the spikelet; upper glume 3-nerved; lower floret
sterile, the lemma 5-nerved, the palea absent; female-fertile
(upper) lemma pale, shiny, minutely scaberulous with a tiny
green spot at the apex.

Flowering March. Forest shade. Rare (in the FSA
region). Biome: Forest. Throughout tropical Africa;
Comoro Is. and Madagascar. Resembles P. monticola,
which has the upper glume 5-nerved and lacks the green
spot at the tip of the female-fertile (upper) lemma.

Description: Clayton et al. 1970-1982 (492). Voucher:
Culverwell 757. PRECIS code 9901 160-00950.

Panicum deustum Thunb.

Reed panicum, broad-leaved
panicum.

Perennial; shortly rhizomatous
and tufted (sometimes rooting at
lower nodes); to 2000(-2400)
mm tall. Leaf blades 150^180
mm long; 5-35(-45) mm wide.

Spikelets 3.5-5.0(-5.5) mm long.

Culms slender or robust, branched or unbranched; leaves
mainly cauline, cordate or straight at the base; inflorescence
branches usually with clavellate hairs; lower glume 1/2-2/3
the length of the spikelet, separated by a short intemode
from the rest of the spikelet; upper glume 7 -nerved; lower
floret male, lemma 5-nerved and palea well developed;
female-fertile (upper) lemma pale, dull or shiny.

Flowering September to April. Often in moist soils,
shady places or rocky hillsides on clay, loam or sandy soils.
Common (but scattered). Biome: Savanna and Forest.
Northwards to Ethiopia and Sudan. Domestic use (grass
mats), or pasture (palatable and nutrituous, staying green
well into dry periods). Variable in size, hairiness and
habitat.

Description: Stapf 1920 (651), Chippindall & Crook
1976 (37). Stapf 1898-1900 (403), Chippindall 1955 (328),
Clayton et al. 1970-1982 (468). Voucher: Crompton 26639;
Godfrey & Acocks SH 1652. PRECIS code 9901160-
01000.

Panicum dewinteri J.G. Anders.

Perennial; tufted (erect, some-
times rooting at the lower nodes);
to 1000 mm tall. Leaf blades
(young leaves) 200-500 mm
long; 1-6 mm wide. Spikelets
3. 5-4.0 mm long. Plant shrub-
like, culms hard and wiry, well
branched, particularly in the up-
per part, leaf blades of the older

2091. PRECIS code

237

portion of the culms falling off early; inflorescence sparsely
branched with secondary branches appressed; lower glume
narrowly ovate, 3-nerved, 2/3 the length of the spikelet;
upper glume and lower lemma 5-7-nerved; lower floret
sterile, with the palea reduced; female-fertile (upper) lem-
ma pale to light brown, shiny.

Flowering January to May. Rocky outcrops, in crevices,
along forest margins and on wooded rocky slopes. Locally
common. Biome: Savanna. Endemic.

Description: Anderson 1967 Bothalia 9,2 (344).
Voucher: Raal 377, Raal 143. PRECIS code 9901160-
01 100.

Panicum dregeanum Nees

Perennial; tufted; to 1100 mm
tall. Leaf blades 140-350(-500)
mm long; 1. 5-3.0 mm wide.

Spikelets 2.0-2. 5(-3.0) mm long.

Basal sheaths silky pubescent,
leaves mostly basal; inflores-
cence 80-150 mm long; spikelets
usually strongly flushed with
purple; glume tips shortly mucro-
nate and recurved; lower glume 1/2-3/4 the length of the
spikelet; upper glume 7-nerved; lower floret male, lemma
5-nerved and the palea well developed; female-fertile
(upper) lemma pale and shiny.

Flowering November to April. Usually in wet places,
frequently in vleis, sometimes on hillsides. Infrequent to lo-
cally common. Biome: Savanna and Grassland. Throughout
tropical Africa. Pasture (grazed by cattle). Resembles P.
genuflexion and P. fluviicola, which have glabrous or
sparsely hispid basal sheaths.

Description: Chippindall & Crook 1976 (36), Stapf 1920
(684), Stapf 1898-1900 (411), Chippindall 1955 (332),
Clayton et al. 1970-1982 (478). Voucher: Smook 1891.
PRECIS code 9901160-01200.

Panicum ecklonii Nees

Perennial; shortly rhizomatous
and tufted; to 800 mm tall. Leaf
blades 60-200(-260) mm long;

3-8 mm wide. Spikelets 2. 5-3. 5
mm long. Leaves mainly basal,
flat, bright green, usually densely
hairy with long tubercle-based
hairs; spikelets usually flushed
with purple; glumes and lower
lemma pectinate at the apex; lower floret sterile with the
palea absent; female-fertile (upper) lemma dull, often flush-
ed purple and shortly hairy towards the apex.

Flowering September to April. Sandy soils and often in
moist areas in mountainous regions that are subjected to
burning. Locally common. Biome: Grassland. Northwards
to Zaire and Tanzania and west Africa. Botha et al. 1988.
S. Afr. J. Bot. 54: 89-93, report that P. ecklonii has both
C3 and C4 forms.

Description: Chippindall & Crook 1976 (47), Stapf
1898-1900 (413), Chippindall 1955 (332), Clayton et al.
1970-1982 (466). Illustration: Chippindall 1955 (fig. 289).
Voucher: Kluge 1968, Hoener 1903. PRECIS code
9901160-01300.

Panicum fluviicola Steud.

{-P. aphanoneurum
Steud.) 3.

Perennial; tufted (erect to gen-
iculate); (300-)600-2300 mm
tall. Leaf blades 130-500 mm
long; 3-12 mm wide. Spikelets
2. 0-2. 5 mm long. Plants often
flushed with purple, basal sheaths

glabrous, sometimes sparsely hispid; culms usually stout,
(2.0-)3.5-7.0 mm wide at the base; inflorescence 80-150
mm long; spikelets green with purple tips to glumes and
lower lemma; glumes acuminate to mucronate, recurved;
lower glume up to 2/3 the length of the spikelet; lower floret
male, lemma 5-nerved, and the palea well developed;
female-fertile (upper) lemma pale, smooth and shiny.

Flowering December to May. Sandy loam, sand or heavy
clays in seasonally wet open areas. Locally common (where
occuring). Biome: Savanna. Scattered throughout tropical
Africa. Barely distinct from P. genuflexum, which has
slender culms that are 1-2 mm wide at the base; also
resembles P. dregeanum , which has silky-pubescent basal

Description: Stapf 1920 (689), Clayton et al. 1970-1982
(478). Voucher: De Winter 4264. PRECIS code

9901160-01450.

Panicum genuflexum Stapf

Perennial; tufted (loosely); to
750(-1000) mm tall. Leaf blades
150-300 mm long; 4-5 mm wide.

Spikelets 2.0-2. 5 mm long. Basal
sheaths glabrous; culms slender,
wiry, 1-2 mm wide at the base;
inflorescence 80-150 mm long;
spikelets often strongly flushed
purple; glume tips usually shortly
mucronate and recurved; lower glume 2/3— 3/4 the length of
the spikelet; lower floret usually male, lemma 5-nerved and
the palea well developed; female-fertile (upper) lemma
pale, smooth and shiny.

Flowering January to March. Usually in sandy soils in
marshy areas and grassy clearings. Rare (in FSA area). Lo-
cally common. Biome: Savanna. Mozambique to Zaire and
east Africa. Barely distinct from P. fluviicola , which has
stout culms (2.0-)3.5-7.0 mm wide at the base, and P.
dregeanum, which has silky pubescent basal sheaths.

Description: Stapf 1920 (689), Clayton et al. 1970-1982
(479). Voucher: Smook 1931. PRECIS code

9901160-01500.

Panicum gilvum Launert

(-P. laevifolium Hack. var.
contractum Pilg.) 2.

Annual; hydrophyte and tufted
(geniculate, rarely erect); to 650
mm tall. Leaf blades 30-150 mm
long; 3-8 mm wide. Spikelets
2. 8-3. 4 mm long. Inflorescence
not exserted beyond the upper-
most leaf which is usually about 8 mm wide. Often there
are two leaves closely associated with the inflorescence.
Secondary inflorescence branches are usually absent and
the spikelets are appressed to the branches; lower glume up
to 1/3 the length of the spikelet; upper glume ( 10— )1 1-14-
nerved; lower floret sterile, lemma (9-) 10-1 1 -nerved, the
palea absent or reduced; female-fertile (upper) lemma pale
to yellow, often flushed dark, smooth and shiny.

Flowering January to April. Sandy soils, margin of vleis,
dams and waterholes, in ephemeral water, and in disturbed
areas. Locally common. Biome: Savanna. Endemic.
Resembles a number of other taxa associated with moist
habitats: P. impeditum andP. subalbidum, which have upp-
er glumes with 3-9 nerves, the lemmas of the lower florets
with 5-9 nerves, and P. sp. I, which has the lower floret
always sterile but a well developed palea.

Description: Launert 1970 Mitt. Bot. Munchen. 8 (153),
Chippindall 1 955 (334). Voucher: De Winter & Giess 6911,
Smith 3300. PRECIS code 9901 160-01600.

238

Panicum glandulopaniculatum Renvoize

Panicum hymeniochilum Nees

Annual; trailing or rambling,
often rooting at the nodes; to
1 000 mm long. Leaf blades to 1 00
mm long; 10-25 mm wide. Spike-
lets 2.0— 2.5(— 3 .0) mm long.

Leaves thick and slightly lea-
thery, flat with cordate bases; in-
florescence branches with flat
glandular patches; the spikelets
asymmetrical; glumes as long as the spikelet, pilose; upper
glume and lower lemma 5-nerved; lower floret male or ster-
ile, with the palea well developed; female-fertile (upper)
lemma shiny, smooth except for scattered papillae occuring
sometimes at the base and/or apex.

Flowering sporadically November to June. Forest shade,
in sand. Locally common. Zambia, Zimbabwe and
Mozambique. Resembles P. heterostachyum, which has
thinner leaves and a densely verruculose upper lemma, and
P. brevifolium , which has eglandular inflorescence
branches and occurs further north.

Description: Renvoize 1989 Kew Bull. 44: 544.
Voucher: Strey 8222. PRECIS code 9901 160-01650.

Panicum heterostachyum Hack.

Annual; erect, loosely tufted;

200-800 mm tall. Leaf blades
80-120 mm long; 10-25 mm
wide. Spikelets to 1.5 mm long.

Plant base may be decumbent;
leaves thin, amplexicaul; inflor-
escence branches with flat gland-
ular patches; spikelets asymmet-
rical, ovate; glumes as long as
the spikelets, sparsely to densely pubescent; upper glume
and lower lemma 5-nerved; lower floret male with the palea
well developed; female-fertile (upper) lemma pale, densely
verruculose.

Flowering January to May (and August). Poor sandy
soils in wooded grassland, seasonally Hooded pans, rocky
hills and in disturbed areas. Locally common. Biome: Sa-
vanna. Throughout tropical Africa, also recorded from
Guyana and Trinidad. Resembles P. glandulopaniculatum ,
which has thick, leathery leaves, and a smooth female-
fertile (upper) lemma which occasionally has scattered
papillae on the apex and base.

Description: Chippindall & Crook 1976 (35), Stapf 1920
(733), Chippindall 1955 (327), Clayton et al. 1970-1982
(496). Illustration: Chippindall 1955 (fig. 285). Voucher;
Merxmueller & Giess 1963. PRECIS code 9901 160-01800.

Panicum hians Ell.

Perennial; tufted (erect, some-
times decumbent to procumbent);
to 600 mm tall. Leaf blades to 200
mm long; to 2.5 mm wide. Spike-
lets 2. 2-2.4 mm long. Inflores-
cence sparsely branched, secon-
dary branches short with the spi-
kelets appressed; lower glume to
1/2 the length of the spikelet,
membranous; lower floret male, the lemma 5-nerved, the
palea coriaceaous, well developed and longer than the lem-
ma; female-fertile (upper) lemma pale, granulose.

Flowering November to January. Damp soils in disturb-
ed places around ponds and streams. Locally common
(where growing in the FSA area). Naturalized from North
America. Biome: Grassland. North America.

Description: Hitchcock & Chase 1950 (703). Voucher:
Wells 1011. PRECIS code 9901160-01900.

(=P. filiculme Schinz) 3; (=P.
hymeniochilum Nees var.
glandulosum Nees) 3; (=P.
hymeniochilum Nees var.
hymeniochilum) 3.

Scrambler and hydrophyte
(often rooting at lower nodes);
culms 140-2000 mm long. Leaf
blades 12-70 mm long; 1 .2-5. 0(-10.0) mm wide. Spikelets
2.0-2. 5(-3.0) mm long. Inflorescence branches sparsely
branched, with clavellate hairs usually present, rarely
absent; spikelets often purple-tinged; lower glume narrowly
ovate, 1/2-2/3 the length of the spikelet; upper glume 7-9-
nerved; lower floret male or sterile, the lemma (7— )9— 1 1-
nerved. the palea usually well developed, sometimes reduc-
ed; female-fertile (upper) lemma granulose, especially
towards the apex.

Flowering December to May. Moist organically rich
soils of river margins and perennial swamps, in or near
water. Locally common. Biome: Savanna, Grassland, and
Forest. Northwards to east Africa, Ethiopia and Guinea;
also in Madagascar. A few specimens from St. Lucia area
are more robust and may represent a new taxon (eg. Feely,
Tinley & Ward 22).

Description: Chippindall & Crook 1976 (79), Chippin-
dall 1955 (324), Clayton et al. 1970-1982 (470). Voucher:
Ward 5518. PRECIS code 9901 160-02100.

Panicum impeditum Launert

Annual; hydrophyte and tufted
(geniculate, erect to prostrate); to
500 mm tall. Leaf blades 20-80
mm long; 3-6 mm wide. Spike-
lets 2. 7-3. 3 mm long. Upper leaf
surface densely covered with
papillae; the base of the inflores-
cence is enclosed by the upper-
most leaf but the inflorescence
extends beyond tip of the leaf; spikelets are crowded and
appressed to the inflorescence branches; lower glume up to
1/2 the length of the spikelet; upper glume and lower lemma
7-9-nerved; lower floret sterile with the palea reduced;
female-fertile (upper) lemma pale, smooth and shiny.

Flowering November and January to May. Moist sandy
or clay soils around water holes, vleis and pans. Locally
common. Biome: Savanna and Nama-Karoo. Endemic.
Resembles P. gilvum, which has the upper glume
( 1 0 — ) 1 1-14-nerved and the lower lemma of the lower floret
(9-) 10-1 1 -nerved, P . sp. I , which has the palea of the lower
floret well developed, and P. subalbidum, which has the
upper leaf surface covered with prickles.

Description: Launert 1970 Mitt. Bot. Munchen. 8(150).
Voucher: Leistner 2229, Leistner 3152. PRECIS code
9901160-02200.

Panicum infestum Peters

Perennial; shortly rhizomatous
and tufted (erect); to 2000 mm
tall. Leaf blades 150-500 mm
long; 2-10 mm wide. Spikelets
3.5( — 4.0) mm long. Plant usually
hairy with tubercle-based hairs,
rarely glabrous; inflorescence
branched sparsely, primary
branches ascending, secondary
branches usually absent; spikelets acute, somewhat
leathery, often with a groove on the back; the closed spike-
lets with only the central nerve visible on the lemma of the
lower floret; lower floret male and the palea well develop-
ed; female-fertile (upper) lemma conspicuously transverse-
ly rugose.

Flowering November, January, February, and May. Clay
or sandy soils in seasonally damp places, rocky hillsides

239

and disturbed areas. Infrequent. Biome: Savanna and Grass-
land. Northwards to Zaire, east Africa and Somalia. Similar
to P. maximum , which has a much more branched inflores-
cence, cartilaginous spikelets, and the closed spikelets with
many nerves clearly visible on the lemma of the lower
floret.

Description: Chippindall & Crook 1976 (34), Chippin-
dall 1955 (330), Clayton et al. 1970-1982 (472). Voucher:
Acocks 16687. PRECIS code 9901 160-02300.

Panicum kalaharense Mez

finely and usually profusely branched; lower glume 1/2 the
length of the spikelet, separated from the rest of the spikelet
by a short internode; upper glume 5-nerved; lower floret
sterile, the lemma 5-7-nerved and the palea reduced;
female-fertile (upper) lemma pale and granulose.

Flowering January to April (June and July). Sandy soils
in dense shade and wet areas in forests. Locally common.
Biome: Forest. South central to east Africa.

Description: Stapf 1920 (736), Chippindall 1955 (325),
Clayton et al. 1970-1982 (498). Voucher: Culverwell 736,
Galpin 2896. PRECIS code 9901 160-02700.

Robust perennial; shortly
rhizomatous and tufted (hard and
dense); to 2500 mm tall. Leaf
blades to 350 mm long; to 8 mm
wide. Spikelets 3. 2-4. 2 mm long.

Basal sheaths densely hairy,
leaves mainly basal, flat or rolled,
upper leaf surface densely cover-
ed with short hairs, sheath mouth
with dense, long to woolly hairs on the abaxial side; lower
glume broadly ovate, up to 1/2 the length of the spikelet,
3-5-nerved; upper glume and lower lemma 7-9-nerved;
lower floret male with the palea well developed; female-
fertile (upper) lemma pale and dull.

Flowering December to April. Kalahari sands and
disturbed places such as roadsides. Infrequent, or locally
common (in dune streets). Biome: Savanna. Zimbabwe.
Food and drink (seeds eaten by Wambos), or pasture
(reasonably palatable when young, drought resistant).
Plants from Namibia that were previously referred to P.
phragmitoides Stapf are a more broad-leaved form found
in the northern part of the distribution range.

Description: Launert 1970 (160:136), Chippindall 1955
(338). Voucher: Ellis 2677, Acocks 12488, Story 6373.
PRECIS code 990 1 1 60-02400.

Panicum lanipes Mez

Wolvoet panicum.

Perennial; densely tufted (e-
rect to geniculate); to 800 mm
tall. Leaf blades to 300 mm long;
to 6 mm wide. Spikelets 2. 0-2. 5
mm long. Usually glaucous, basal
leaf sheaths densely covered with
matted woolly hairs; spikelets
light green, sometimes tinged with purple; lower glume up
to 2/3 the length of the spikelet; upper glume and lower
lemma 7-9-nerved; lower floret male with the palea well
developed; female-fertile (upper) lemma pale yellow to
brown, shiny.

Flowering September to May. Stony, sandy or calcrete
soils in vleis, on mountain slopes or in dry river beds. Lo-
cally common. Biome: Savanna and Nama-Karoo.

Endemic. Pasture (valuable fodder). Similar to P. pearsonii
Bol. f., which has a woolly base but longer spikelets
(3. 0-3. 5 mm long). As far as known, P. pearsonii has never
been recollected since the type specimen. It also resembles
P. coloratum and P. stapfianum, which have glabrous or
hairy bases, but then the hairs are not woolly and matted.

Description: Launert 1970 (160:136), Chippindall 1955
(337). Illustration: Muller 1984 (fig. 99), Chippindall 1955
(fig. 294). Voucher: Muller 1389, Giess, Volk & Bleissner
6104. PRECIS code 9901160-02600.

Panicum laticomum Nees

Annual; scrambling, decum-
bent to semi-erect, rooting at the
nodes; culms to 2000 mm long.

Leaf blades to 100 mm long;

(5— )8— 28 mm wide. Spikelets
1.5-2. 5 mm long. Leaves abrupt-
ly or asymmetrically narrowed at
the base; inflorescence extending
beyond the uppermost leaf, open.

Panicum maximum Jacq.

Guinea grass, blousaad soet-
gras.

Usual'y perennial, or annual
(occasionally); loosely to densely
tufted (erect and geniculate, root-
ing at the nodes); to 2000 mm tall.

Leaf blades 60-400(-1000) mm
long; 4— 1 2(— 35 ) mm wide. Spike-
lets 2.5— 3.0(— 4.0) mm long. Inflorescence usually much
branched with secondary branches well developed and
flexuous; spikelets blunt or acute, rounded on the back,
cartilaginous; the closed spikelet has many nerves clearly
visible on the lemma of the lower floret; lower floret usual-
ly male with the palea well developed; female-fertile
(upper) lemma pale and conspicuously transversely rugose.

Flowering November to July. In shady places, especially
under canopy of trees, in cultivated areas and along river
banks, but well adapted to a variety of conditions. Widely
common. Biome: Fynbos, Savanna and Nama-Karoo. To
tropical Africa, and in Madagascar. Widely introduced
throughout tropics. Extensively planted hay and pasture (of-
ten grown in the form of selected agricultural strains in the
tropics). There is considerable variation in size and
indumentum of culms, leaves and inflorescences. Resembl-
es P infestum , which has the inflorescence sparsely branch-
ed, secondary branches usually absent, spikelets leathery
and the closed spikelets with only the central nerve visible
on the lemma of the lower floret.

Description: Stapf 1920 (655), Chippindall & Crook
1976 (33), Chippindall 1955 (329), Clayton et al.
1970-1982 (471). Voucher: De Winter 9167, Godfrey SH
1709. PRECIS code 9901160-02800.

Panicum merkeri Mez

( -P . radula Mez )

Perennial; stout, shortly rhizomatous and tufted (erect);
to 1600 mm tall. Leaf blades to 350 mm long; 7-15 mm
wide. Spikelets 2. 0-2. 5 mm long. Plant generally hispid;
lower glume 1/4-1/3 the length of the spikelet; upper glume
and lower lemma 9(— 1 l)-nerved; lower floret male with the
palea well developed; female-fertile (upper) floret pale and
shiny.

Swamps and seasonally damp places in heavy clay soils.
Infrequent. Biome: Savanna. Reported from Namibia.
Angola and north to east Africa. No specimen definitely
referred to this species has been seen, but it belongs to the
P. coloratum -P . stapfianum complex which needs study.

Description: Clayton et al. 1970-1982 (486). Voucher:
photo of type ofP. radula, Morgenstein (B). PRECIS code
9901160-02850.

Panicum monticola Hook. f.

Perennial; trailing, decum-
bent, rooting at the nodes; culms
300-1000 mm long. Leaf blades
to 150 mm long; ( 5—) 1 0—25 mm
wide. Spikelets 2.2-3.5(-4.0) mm
long. Inflorescence sparsely and
irregularly branched, exserted
from the uppermost leaf; lower
glume broadly ovate, 1/4- 1/2 the
spikelet length, 0-1 -nerved; upper glume and lower lemma

Fig. 151. Pl. 139.

240

5-nerved; lower floret sterile with the palea absent or
rudimentary; female-fertile (upper) lemma pale, smooth
and shiny.

Flowering January, April and June. In the shade of
forests. Rare (in the FSA area). Biome: Forest. Throughout
tropical Africa.

Description: Clayton et al. 1970-1982 (494). Voucher:
Scheepers 399. PRECIS code 9901 160-03000.

Panicum natalense Hochst.

Fig. 152. PI. 140.

(=P . fulgens auctt., non
Stapf) 3.

Natal buffalo grass, Natal-
buffelsgras.

Perennial; shortly rhizomatous
and tufted (densely); to 500
(-800) mm tall. Leaf blades to
500 mm long; to 3.5 mm wide. Spikelets 1.7-2. 2 mm long.
Plant base knotty, leaves mainly basal, tightly folded or flat;
spikelets nearly rounded in outline; lower glume 1/2-3/4
the spikelet length; upper glume and lower lemma 5-nerved;
lower floret male or sterile, the palea membranous, well
developed or reduced in width only, not longer than the
lemma; female-fertile (upper) lemma pale, sparsely to
densely papillose.

Flowering October to April. Sandy loam or sandy soils
in well drained or shallow soils in rocky areas, often in
burnt veld. Infrequent to common. Biome: Savanna and
Grassland. Angola and Zimbabwe. Pasture (normally only
grazed in spring after burning). A few specimens in the
Transvaal may represent a new species (eg. Codd 2736).
They have longer rhizomes and the plants are more loosely
tufted and the leaves are not basal, but cauline.

Description: Chippindall & Crook 1976 (40). Stapf
1898-1900 (412), Chippindall 1955 (333). Voucher:
Smook 1 154, Smook 5005. PRECIS code 9901 160-03100.

Panicum novemnerve Stapf

Annual; loosely tufted (geni-
culate); to 600 mm tall. Leaf blad-
es to 200 mm long; to 15 mm
wide. Spikelets 2. 0-2. 5 mm long.

Bulbous-based hairs at least on
the leaf sheaths; inflorescence
extends beyond the tip of the
uppermost leaf, but the base is
enclosed in the uppermost leaf,
broadly ovate, branches long, flexuous, spreading, densely
scabrid with prickles that are 0.10-0.15 mm long just below
the spikelets; lower glume to 2/3 the length of the spikelet;
upper glume and lower lemma (7-)9-nerved; lower floret
sterile with the palea conspicuously reduced; female-fertile
(upper) floret pale to dark, smooth and shiny.

Flowering December to April. Moist clayey loams,
brackish soils along drainage lines and in depressions where
water collects. Infrequent. Biome: Savanna. Southern
tropical Africa. Barely distinguishable front P. arcurameum
and a detailed study is needed. Resembles P.
atrosanguineum, which has the inflorescence branches only
moderately scabrid with prickles.

Description: Stapf 1920 (702), Chippindall & Crook
1976 (39), Chippindall 1955 (327). Voucher: Freyer 36.
PRECIS code 9901 160-03200.

Panicum obumbratum Stapf

Perennial; trailing, prostrate
base, rooting at nodes; culms to
500 mm long. Leaf blades to 40
mm long; 3-6 mm wide. Spike-
lets 4 mm long. Inflorescence
narrow, sparsely branched with
few spikelets; lower glume 1/3
the length of the spikelet; upper
glume 7-nerved; lower floret

male, the lemma 5-nerved, the palea well developed;
female-fertile (upper) lemma minutely transversely rugose.

Flowering December to January. Shady places around
streams and forests. Rare. Biome: Forest. Endemic.
Resembles Brachiaria chusqueoides, which has
pseudopetiolate leaves.

Description: Stapf 1898-1900 (401), Chippindall 1955
(326). Voucher: Acocks 17887. PRECIS code 9901160-
03300.

241

Panicum parvifolium Lam.

Perennial; scrambler and hy-
drophyte (rooting at lower
nodes); culms 80-500 mm long.

Leaf blades (1 3-) 15-30 mm long;

2-7 mm wide. Spikelets 1-2 mm
long. Leaves often reflexed at
maturity, lanceolate to narrowly
ovate, acute, base cordate; inflor-
escence small, 10-50 mm long,
open, often barely exserted from the uppermost leaf sheath;
lower glume 1/2-2/3 the length of the spikelet, glume tips
not recurved; upper glume and lower lemma 5-nerved; low-
er floret usually male, the palea well developed; female-
fertile (upper) lemma pale, smooth and shiny.

Flowering December to June. Organically rich sandy
soils in water or along streams or in swamps. Infrequent.
Biome: Grassland and Forest. Throughout tropical Africa,
and in Madagascar and tropical America. Resembles P.
subflabellatum , which has longer leaves (30-70 mm long)
and a shorter lower glume (1/3 the length of the spikelet).

Description: Stapf 1920 (726), Chippindall 1955 (325),
Clayton et al. 1970-1982 (490). Voucher: Acocks 13340.
PRECIS code 9901 160-03500.

Panicum pilgerianum (Schweick.) Clayton

(=Acroceras pilgeranum
Schweick.) 2; ( ^Psilochloa
pilgerana (Schweick.)

Launert) 4.

Annual; hydrophyte and tuft-
ed; to 2000 mm tall. Leaf blades
to 320 mm long; to 10 mm wide.

Spikelets 4. 5-6.0 mm long. In-
florescence branches ascending, with large spikelets densely
appressed to the branches; upper glume 9-nerved; lower
floret sterile or male, or both on the same inflorescence, the
lemma 7-nerved, the broadest interspaces between the nerv-
es are adjacent to. the central nerve, and the palea well
developed or slightly shorter and narrower; female-fertile
(upper) lemma pale, dull, with a rough surface.

Flowering February to June. Growing in clay soils, in
water of dams and pans, also in vleis. Locally common.
Biome: Savanna. Endemic. Psilochloa has been placed in
synonomy with Panicum since the characters on which it
is based do not separate it from the large genus Panicum.

Description: Chippindall 1955 (386). Voucher: Smith
1899. PRECIS code 9901 160-03750.

Panicum repens L.

Couch panicum, kruipgras.

Perennial; occasional hydro-
phyte, rhizomatous, and tufted (e-
rect to decumbent, sometimes
floating); to 1000 mm tall. Leaf
blades 70-250 mm long; 2-8 mm
wide. Spikelets 2-3 mm long.

Leaves mainly cauline, distich-
ous, ascending, usually pungent; inflorescence narrowly
oblong, sparsely to moderately branched, usually ascend-
ing; lower glume broadly ovate, to 1/2 the length of the
spikelet; upper glume and lower lemma 7-9-nerved; lower
floret usually male, the lemma with the broadest interspaces
between nerves not confined to adjacent to the central
nerve, the palea well developed; female-fertile (upper) lem-
ma pale to yellowish, shiny.

Flowering October to June. Wet sandy soils, sometimes
in either fresh or brackish water. Locally common. Biome:
Fynbos, Savanna, Grassland, and Desert. Throughout
tropics and subtropics. Pasture (grazed by game and
domestic livestock in Botswana), or erosion control (plant-
ed around dams in Zimbabwe).

Description: Stapf 1920 (708), Chippindall & Crook
1976 (38), Chippindall 1955 (333), Clayton et al.
1970-1982 (481). Illustration: Clayton et al. 1970-1982
(fig. 121). Voucher: Smook 4211. PRECIS code

9901160-03800.

Panicum repentellum Napper

Perennial (to subperennial);
hydrophyte, rhizomatous, and
stoloniferous (erect or decum-
bent); to 600 mm tall. Leaf blades
50-120 mm long; 3-5 mm wide.

Spikelets 2. 0-3. 5 mm long.

Slender leaves mainly cauline; in-
florescence closely associated
with uppermost leaf, narrow and
sparsely branched, ascending; lower glume broadly ovate;
upper glume and lower lemma 7-nerved; lower floret usual-
ly male, the lemma with the broadest interspaces of the
nerves adjacent to the central nerve, and the palea well
developed; female-fertile (upper) lemma pale, shiny.

Flowering January to May. Black or grey soils around
vleis and lakes or in water. Infrequent. Biome: Savanna.
Malawi, Zambia, Zaire to east Africa, Ethiopia and Sudan.

Description: Clayton et al. 1970-1982 (482). Voucher:
Chippindall 346, Smith 3812. PRECIS code
9901160-03850.

Panicum schinzii Hack.

( -P . laevifolium Hack. var.
laevifolium) 3.

Land grass, blousaadgras.

Annual; tufted (erect to
sprawling, occasionally rooting at
the nodes); to 900 mm tall. Leaf
blades to 300 mm long; to 20 mm
wide. Spikelets 2.0-2. 5(-3. 5) mm long. Lower sheaths gla-
brous or sparsely hairy at the extreme base, leaves dark
green; inflorescence well exserted from the uppermost leaf,
open, oblanceolate to narrowly obovate, profusely branched
with many spikelets; spikelets blunt; tips of the glumes not
recurved; lower glume broadly ovate; upper glume and
lower lemma 9-1 1 -nerved; lower floret male or both male
and sterile occuring in the same inflorescence, the palea
well developed; female-fertile (upper) lemma pale to
yellow, shiny.

Flowering November to May. Moisture loving, in sandy
and clay soils of seepage areas, depressions where water
collects and in cultivated lands. Common. Biome: Fynbos,
Savanna, Grassland, and Nama-Karoo. Zimbabwe and Mo-
zambique. Good hay, palatable pasture and weed (of
cultivated lands). Resembles P. sp. 2, which is yellowish
green, with inflorescence obovate to broadly obovate,
spikelets acute and occurs in northern Namibia; also P
subalbidum , which has appressed spikelets and palea of the
lower floret conspicuously reduced in length.

Description: Chippindall 1955. Voucher: Smook 3163,
Galpin 401. PRECIS code 9901 160-03900.

Panicum stapfianum Fourc.

Perennial; tufted; to 900 mm
tall. Leaf blades to 400 mm long
(but usually shorter); to 5 mm
wide. Spikelets to 3 mm long.

Plants with bulbous-based hairs
present or absent, leaves mainly
basal, upper leaf surface usually
densely papillate; lower glume
1/2-2/3 the length of the spikelet;
upper glume and lower lemma 7-9-nerved; lower floret
male with the palea well developed; female-fertile (upper)
lemma pale yellow to flushed greyish, smooth.

242

Flowering November to May. Damp area on heavy or
sandy soils, disturbed or calcrete areas and occasionally
rocky dry areas. Locally common. Biome: Fynbos,
Savanna, Grassland, and Nama-Karoo. Endemic. Highly
palatable pasture. Part of the P. coloratum-P . stapfianum
complex, which also includes P. bechuanense and P.
merkeri. Detailed study of species limits is needed.

Description: Chippindall 1955 (336). Illustration:
Muller 1984 (fig. 102), Chippindall 1955 (fig. 293).
Voucher: Smook 2357. PRECIS code 9901 160-04000.

Panicum subalbidum Kunth

of plant usually strongly flushed with purple; inflorescence
moderately branched, ascending, lower part of the branches
naked for a distance below the spikelets; lower glume
broadly ovate, to 1/4 the length of the spikelet; upper glume
and lower lemma 9-11-nerved; lower floret male with the
palea well developed; female-fertile (upper) lemma pale to
yellow, shiny.

Flowering January to May. Black clays or sand in
seasonally flooded areas, vleis and pan edges. Infrequent.
Biome: Savanna. Zambia. Domestic use (thatching).

Description: Launert 1970 (160:138). Voucher: Rodin
9098. PRECIS code 9901 160-04300.

( -P . glabrescens Steud.) 3.

Elbow buffalo grass.

Shortlived perennial, or an-
nual; hydrophyte (occasionally),
or tufted (erect to decumbent and
rooting at the nodes); to 2000 mm
tall. Leaf blades 200-500 mm
long; 6-18 mm wide. Spikelets 2. 5-3. 5 mm long. Plant
slender to robust; upper leaf surface with large, usually
white prickles always visible on the upper leaves, papillae
sometimes present; inflorescence sparsely to moderately
branched with spikelets appressed to the branches; spikelets
acuminate; lower glume broadly ovate, to 1/3 the length
of the spikelet; upper glume and lower lemma 7-9-nerved;
lower floret sterile with the palea absent or poorly develop-
ed; female-fertile (upper) lemma pale to yellowish, shiny.

Flowering October to April. Usually on clay soils in
disturbed areas, in or near water, along rivers and swamps.
Infrequent. Biome: Savanna and Grassland. Throughout
tropical Africa. This species is very variable in size. It
resembles P impeditum, which has papillae only, and lacks
prickles on the upper leaf surface.

Description: Chippindall & Crook 1976 (48), Chippin-
dall 1955 (333), Clayton et al. 1970-1982 (484). Voucher:
Smith 1438, Theron 1978. PRECIS code 9901160-04050.

Panicum subflabellatum Stapf

Perennial; sometimes shortly
rhizomatous, stoloniferous, and
tufted (erect or decumbent); to
500 mm tall. Leaf blades 30-70
mm long; 2-3 mm wide. Spike-
lets 1 .3-1.7 mm long. Lower leaf
sheaths glabrous, leaves not
reflexed, linear-lanceolate, taper-
ing to a long acuminate tip, base
straight; inflorescence moderately branched, 10-60 mm
long; spikelets spherical, strongly flushed with purple;
glume apex not recurved; lower glume 1/3 the length of the
spikelet, broadly ovate; upper glume 5-nerved; lower floret
usually male, lemma 5-7-nerved, and the palea well
developed; female-fertile (upper) lemma pale, shiny or dull.

Flowering December and March. Sand dunes and
swampy areas along coast. Locally common. Biome: Sa-
vanna. Mozambique and Tanzania. Resembles P. parvi-
folium , which has shorter leaves (15-30 mm long) and a
longer lower glume (1/2-2/3 the length of the spikelet).

Description: Stapf 1920 (711), Clayton et al. 1970-1982
(481). Voucher: Ward 8822. PRECIS code 990 1 1 60-04 1 00.

Panicum trichonode Launert & Renvoize

Perennial; shortly rhizomatous
and tufted (densely, erect or
occasionally geniculate); to 900
mm tall. Leaf blades 100-300
mm long; 2-6 mm wide. Spike-
lets (2.0-)2.5-3.0 mm long.

Plants not appearing leafy, leaves
mainly cauline, upper leaf surface
densely papillate; culm nodes
densely covered with short appressed hairs; lower portion

Panicum volutans J.G. Anders.

Rolling grass, tumble weed.

Annual; loosely tufted (erect
to decumbent and rooting at the
nodes); to 750 mm tall. Leaf blad-
es 230 mm long; 0.5-10.0 mm
wide. Spikelets (5.5-)6.0-6.5
(-7.0) mm long. Leaves hispid
with bulbous-based hairs; inflor-
escence large, open, spreading and extending beyond the tip
of the uppermost leaf while the base is often enclosed in
uppermost leaf, branches naked for long distance before the
1-3 spikelets at the tips; spikelets long-acuminate; upper
glume 7-9-nerved; lower floret sterile, lemma 7-nerved,
and the palea reduced; female-fertile (upper) lemma
yellowish grey with very conspicuous nerves.

Flowering January to March. Mainly in black turf in
cultivated and disturbed areas and areas of high moisture.
Locally common. Biome: Savanna, Grassland. Endemic.
Chippindall (1955) treated this species as P. sp. aff. P.
hippothrix K. Schum. It is the only Panicum species in the
FSA area in which the whole inflorescence breaks off at
maturity and is rolled about by the wind.

Description: Anderson 1960 Bothalia 7 (420), Chippin-
dall 1955 (328). Voucher: Scheepers 1478. PRECIS code
9901160-04400.

Panicum sp. 1 (=Smook 3463)

Annual; tufted (often genicu-
late); 150-180 mm tall. Leaf
blades to 120 mm long; 3-6 mm
wide. Spikelets 2. 4-2. 8 mm long.

Plant often Hushed with purple;
inflorescence always exserted out
to one side of the uppermost leaf,
shorter or longer than the leaf tip,
moderately branched, branches
appressed, ascending; spikelets appressed to the inflores-
cence branches; lower glume broadly ovate, to 1/3 the
length of the spikelet; upper glume and lower lemma 9-1 1-
nerved; lower floret always sterile with the palea either well
developed or only slightly reduced in width; female-fertile
(upper) lemma pale yellow to dark, shiny.

Flowering November to April. Moist soils around vleis,
dams, pans, and depressions, and in disturbed areas. Locally
common. Biome: Grassland and Nama-Karoo. Endemic.
Resembles P. gilvum and P. impeditum , which have the
palea of the lower floret greatly reduced in length and
width.

Voucher: Smook 3463. PRECIS code 9901 160-99999.
Panicum sp. 2 (=Giess 8605)

Annual; tufted (erect, some-
times geniculate); to 1200 mm
tall. Leaf blades 100-250 mm
long; 5-15 mm wide. Spikelets
2. 2-2. 8 mm long. Plant yellowish
green, lower sheaths glabrous to
sparsely hairy; inflorescence obo-
vate to broadly obovate, open,
profusely branched, well exserted

243

from the uppermost leaf; spikelets acute, often strongly
flushed with purple; lower glume ovate; upper glume usual-
ly with small recurved mucro at the tip; upper glume and
lower lemma 7-9-nerved; lower floret always male with the
palea usually well developed, occasionally slightly reduced
in length; female-fertile (upper) lemma pale to yellow to
dark brown.

Flowering February to May. Moist sand to clayey loams
(often overlaying calcrete) around vleis, pans, and dams and
often in disturbed moist areas. Locally common. Biome: Sa-
vanna. Endemic. Resembles P. schinzii , which probably do-
es not occur in Namibia, but is usually green, with the in-
florescence oblanceolate to narrowly obovate, and the
spikelets blunt-tipped. The plants assigned to this new
taxon have previously been referred to P. novemnerve ,
which has the palea of the lower floret poorly developed.
Until further study, these specimens will be kept as a
separate entity.

Voucher: Giess 10799, Giess 8605. PRECIS code
9901160-99999.

Parapholis C.E. Hubb.

Lepidurus Janchen.

Annual (erect or more or less prostrate); caespitose.
Culms 20-500 mm high; herbaceous; branched above, or
unbranched above. Leaf blades linear; flat, or rolled
(convolute). Ligule an unfringed membrane .

Inflorescence a single spike ( cylindrical ); espatheate.
Spikelet-bearing axes disarticulating at the joints.

Spikelets solitary; distichous; 4-7 mm long ; compressed
laterally, falling with the glumes (shed with rachis joints).
Glumes two; more or less equal; displaced (abaxial, side by

side); awnless; similar (leathery). All florets female-fertile;
proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes (membranous, side-on to the rachis); 3 nerved (the
laterals very short); awnless. Palea present; relatively long.
Lodicules 2; membranous; glabrous. Stamens 3. Ovary
glabrous. Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7, 9, and 19. Pooideae; Poodae; Poeae. 6
species. Western Europe, Mediterranean to India.
Maritime-arenicolous to halophytic, or glycophytic. Cape
Province. 1 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Parapholis incurva (L.) C.E. Hubb.

Fig. 153. PI. 141.

Annual; culms erect or
decumbent; 60-300 mm tall. Leaf
blades inconspicuous, to 30 mm
long; 1 .5-2.0 mm wide. Spikelets
4. 5-5. 5 mm long. Spikes cylin-
drical, solitary, usually curved;
the two glumes lying side by side,
keel not winged.

Flowering August to October.

Weedy at roadsides and in moist places. Infrequent. Nat-
uralized from Europe. Biome: Fynbos. Widely introduced
in temperate areas. Weed. Similar to Hainardia cylindrica,
which has a single glume.

Description: Tutin 1980 FI. Europ. 5 (243), Chippindall
1955 (73). Voucher: Acocks 17786. PRECIS code
9904431-00100.

Paratheria Griseb.

Perennial; geniculate ascending, rooting at the nodes.
Culms 150-800 mm high; herbaceous; branched above.
Leaf blades linear; flat. Ligule a fringed membrane (very
narrow ), or a fringe of hairs. Plants with hermaphrodite
florets. The spikelets all alike in sexuality (but sometimes
with cleistogamous spikelets lacking bristles at the base of
the inflorescence). Plants with hidden cleistogenes (in the
upper sheaths).

Inflorescence paniculate ; contracted; espatheate.
Spikelet-bearing axes disarticulating; falling entire (i.e., the
lateral branch disarticulates, carrying with it the spikelet
and bristle, and constituting a pointed callus beneath the
spikelet).

Spikelets (at least some of them) subtended by solitary
‘bristles’-, solitary (and 1 spikelet per branch). Spikelets
8-13 mm long; abaxial; compressed dorsiventrally; falling
with the glumes (and the branch). Glumes two; minute;
more or less equal; awnless; similar (hyaline). Proximal in-
complete florets I ; epaleate; sterile.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes; smooth; not becoming indurated; hairless;
having the margins lying flat and exposed on the palea;
without a germination flap; 7 nerved; entire; awnless (but
acuminate-subulate, like the LI). Palea present; relatively
long (about equalling the lemma). Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Hilum short; embryo
large.

Photosynthetic pathway. C4; XyMS-.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 2 species. Africa, Madagascar, Cuba,
Brazil. Hydrophytic, or helophytic; in open habitats
(swamps and lakes); glycophytic. Namibia. 1 indigenous
species.

References. 1. Launert. 1970. FSWA.

Species treatment by H.M. Anderson.

244

Paratheria prostrata Griseb.

PI. 142.

Perennial; tufted; 150-450
mm tall. Leaf blades 20-60 mm
long; 2^4 mm wide. Spikelets
about 9 mm long; 1 mm wide.

Culm nodes bearded; spikelet
with a little tuft of hairs at base
and a single bristle 20 mm long;
the spikelet and bristle disarticu-
late with the lateral branch
which becomes a pointed callus beneath the spikelet.

Flowering December to January. Growing in and near
water. Infrequent. Biome: Savanna. Tropical west Africa,
Cuba, Brazil and Madagascar.

Description: Clayton 1972 FTWA (457), Launert 1970
(160:143). Voucher: De Winter 4049. PRECIS code
9901420-00100.

Paspalidium Stapf

Somewhat marginally separable from Setaria P. Beauv.

Annual, or perennial (often aquatic); long-rhizomatous,
or caespitose to decumbent. Culms herbaceous. Nodes
glabrous. Leaf blades flat, or rolled. Ligule a fringed
membrane (very narrow), or a fringe of hairs.

Inflorescence of spike-like main branches ( the branches
generally oppressed to the rachis, and sometimes greatly
reduced)', espatheate. Spikelet-bearing axes persistent.

Spikelets unaccompanied by bractiform involucres, not
associated with setiform vestigial branches (this being the
‘distiction from Setaria: however, the terminal spikelet of
each branch is associated with the branch-tip bristle, and
since the ‘ branches' may be reduced to single spikelets, the
separation is scarcely adequate)', solitary, or in pairs;
biseriate. Spikelets abaxial', compressed dorsiventrally;
falling with the glumes. Glumes two; very unequal;
awnless. Proximal incomplete florets l ; paleate, or epaleate,
palea when present fully developed to reduced: male, or
sterile.

Female-fertile florets I . Lemmas decidedly firmer than
the glumes; rugose; becoming indurated (crustaceous);
hairless; having the margins tucked in onto the palea; with
a clear germination flap; 5 nerved; entire; awnless (often
apiculate). Palea present; relatively long. Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit small;
ellipsoid to subglobose; hilum short; embryo large.

Photosynthetic pathway. C4; XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. Abaut 40
species. In warm regions. Hydrophytic to mesophytic; in
shade and in open habitats (swamps, forests, dry slopes);
glycophytic. Namibia, Botswana, Transvaal, Natal, and
Lesotho. 2 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by M. Koekemoer.

1(0). Spikelets 1.6-2. 6 mm long, ovate; central axis of
inflorescence 0. 5-1.0 mm wide, narrowly winged;
leaf blades setaceously acuminate . P. geminatum
Spikelets 3. 0-3. 5 mm long, narrowly ovate; central
axis of inflorescence 3-5 mm wide, broadly winged
and ribbon-like; leaf blades bluntly acute to broadly
obtuse and then often notched at the apex and
splitting along the midrib P. obtusifolium

Fig. 154. Paspalidium obtusifolium

Paspalidium geminatum (Forssk.) Stapf

Perennial; hydrophyte and
rhizomatous (rhizome floating
and spongy), or stoloniferous
(culms prostrate and rooting at
the nodes); 100-600 mm tall.

Leaf blades 50-350 mm long;

2-13 mm wide. Spikelets 1.6-2. 6
mm long. Leaf blades acuminate;
central axis of inflorescence
narrowly winged, 0. 5-1.0 mm wide; spikelets ovate.

Flowering March to June. In water up to 2 m deep but
also extending to wet marshy soils on the edges of rivers,
pans or vleis. Infrequent to locally common. Biome: Savan-
na, Grassland, and Nama-Karoo. Old world tropics. Easily
distinguished from P. obtusifolium, which has larger spike-
lets and blunt leaf tips.

245

Description: Stapf 1920 (582), Chippindall 1955 (366),
Clayton et al. 1970-1982 (552). Illustration: Clayton et al.
1970-1982 (fig. 133). Voucher: Leistner, Oliver,

Steenkamp & Vorster 253, Giess 3132. PRECIS code
9901090-00100.

Paspalidium obtusifolium (Del.) Simpson

Fig. 154. PI. 143.

(=P. platyrrhachis C.E.

Hubb.) 2.

Perennial; hydrophyte and
rhizomatous (rhizome floating
and spongy), or stoloniferous
(culms prostrate and rooting at
the nodes); 300-600 mm tall.

Leaf blades 30-200 mm long;

4-12 mm wide. Spikelets 3. 0-3. 5 mm long. Leaf blades
bluntly acute to broadly obtuse, often notched at the
rounded apex and splitting along the midrib; central axis of
inflorescence broadly winged and ribbon-like, 3-5 mm
wide; spikelets acute and narrowly ovate.

Flowering September to May. On marshy soils or
shallow water in pans, often with culms floating. Infrequent
to locally common. Apparently indigenous, but possibly
brought from Egypt by waterbirds. Biome: Savanna.
Northwards to Algeria and Egypt. Easily distinguished from
P. geminatum, which has smaller spikelets and acuminate
leaf tips. Vegetatively similar to Stenotaphrum secundatum,
which is not aquatic and has minute spikes sunk in the
rachis.

Description: Chippindall 1955 (366), Clayton et al.
1970-1982 (551). Illustration: Chippindall 1955 (fig. 315).
Voucher: Davidse 5868. PRECIS code 9901090-00200.

Paspalum L.

Anachyris Nees, Cerea Schlecht., Ceresia Pers.,
Cleachne Roland, ex Rottb., Cymotochloa Schlecht.,
Dichromus Schlecht., Digitaria Fabric., Dimorphostachys
Fourn., Maizilla Schlecht., Moenchia Steud.,
P aspalanthium Desv., Reimaria Fluegge, Sabsab Adans.,
Wirtgenia Doell.

Perennial; long-rhizomatous, or long-stoloniferous, or
caespitose, or decumbent. Culms 100-3000 mm high
(rarely taller, sometimes with culms trailing to 2 m or
more); herbaceous. Leaf blades linear, or linear to linear-
lanceolate; flat, or folded, or rolled. Ligule an unfringed
membrane to a fringe of hairs. Plants bisexual, with
bisexual spikelets.

Inflorescence of spike-like main branches', digitate or
subdigitate, or non-digitate; espatheate. Spikeiet-bearing
axes disarticulating (e.g., P. repens), or persistent; when
disarticulating falling entire.

Spikelets biseriate; consistently in ‘long-and-short’
combinations, or not in distinct ‘long-and-short’ combina-
tions. Spikelets ( 1 .2-) 1 .5 — 4.2( — 4.5) mm long; abaxiab, com-
pressed dorsiventrally; falling with the glumes; awnless,
muticous. Glumes present (usually), or absent (in Section
Anachyris)', when present one per spikelet (in species with
an ‘andropogonoid’ spikelet arrangement), or two; very
unequal; awnless; very dissimilar (G1 usually much
reduced). Lower glume 0-1 nerved. Proximal incomplete
florets 7; epaleate; sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes (papery to
crustaceous); smooth to striate; becoming indurated, or not
becoming indurated; hairless; having the margins tucked in
onto the palea; with a clear germination flap; 3-5 nerved;
entire; blunt', awnless. Palea present; relatively long. Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; embryo large, or small (rarely).

Photosynthetic pathway. C4; NADP-ME (notatum,
dilatatum)', XyMS-. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x - 10 and 12. Panicoideae; Panicodae; Paniceae.
320 species. In warm regions. Mostly helophytic, or meso-
phytic, or xerophytic; mostly in open habitats (diverse
habitats — savanna, damp places, forest margins, weedy
ground, coastal and inland saltmarshes); maritime-
arenicolous (a few — P . vaginatum being a useful sand
stabilizer), or halophytic (e.g. P . distichum ), or glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. Indigenous
species (3), naturalized species (3).

Fig. 155. Paspalum dilatatum

246

References. 1 . De Winter & Vorster. 1974. Bothalia 1 1 :
295. 2. Clayton & Renvoize. 1982. FTEA. 3. Clayton &
Renvoize. 1980. Taxon 29: 337. 4. Webster. 1987. The
Australian Paniceae (Poaceae).

Species treatment by M. Koekemoer.

1(0). Spikelets hairy or fringed with hairs, matt, arranged

in four rows on one side of the rachis 2

Spikelets glabrous or minutely hairy on the body,
usually glossy, arranged in two rows on one side

of the rachis 3

2(1). Spikelets longer than 3 mm; racemes 4-9, scattered
on a central axis 30-200 mm long; basal sheaths

glabrous or sparsely hairy P. dilatatum

Spikelets shorter than 3 mm; racemes 10-30, closely
arranged on a central axis 120-300 mm long; basal

sheaths densely hairy P. urvillei

3(1). Spikelet tips rounded; spikelets ovate to elliptic,

1 .0- 1 .5 times as long as wide 4

Spikelet tips acute; spikelets broadly lanceolate, more

than 2 times longer than wide 5

4(3). Spikelets 2. 8-3. 7 mm long; rachis often zig-zag,
about half the width of the raceme; rhizome very

well developed and horizontally creeping

P. notatum

Spikelets 2. 0-2. 5 mm long; rachis flat and linear,
almost leaf-like, as wide as the raceme; rhizome

short and not horizontally creeping

P. scrobiculatum

5(3). Lower glume usually developed as a small triangular
scale or up to half the spikelet length; upper glume
minutely hairy; leaves more than 3 mm wide;
spikelets 2.5— 3.5 mm long, narrow obovate ....

P. distichum

Lower glume absent or reduced to a rim; upper glume
glabrous; leaves less than 3 mm wide; spikelets

3. 0- 4. 5 mm long, almost lanceolate

P. vaginatum

Paspalum dilatatum Poir.

Fig. 155. PI. 144.

Dallis grass, gewone pas-
palum, watergras.

Perennial; rhizomatous (rhi-
zome short and creeping), or tuft-
ed; 300-1800 mm tall. Leaf
blades 90-350(^450) mm long;

6-14 mm wide. Spikelets 3-4 mm
long; 2. 0-2. 5 mm wide. Basal
sheaths glabrous or sometimes hairy; ligule conspicuous,
membranous, 2-8 mm long; inflorescence with central axis
30-200 mm long; racemes 4-9, scattered on axis; spikelets
fringed with white hairs and arranged in four rows on one
side of the rachis.

Flowering October to May. Usually in damp places,
most often in disturbed areas such as roadsides, gardens and
cultivated lands. Common. Naturalized from South
America. Biome: Fynbos, Savanna, Grassland, and Nama-
Karoo. Tropics worldwide. Pasture (widely used as fodder
or leys), or erosion control (stabilization of minedumps), or
weed (common invader). Resembles P. urvillei , which has
more racemes and smaller spikelets.

Description: Webster 1987 (172), Chippindall & Crook
1976 (12), Hitchcock & Chase 1950 (590), Chippindall
1955 (387), Clayton et al. 1970-1982 (608). Illustration:
Chippindall 1955 (pi. 13(11)), Hitchcock & Chase 1950 (fig.
1244). Voucher: Smook 4134. PRECIS code

9901070-00100.

Paspalum distichum L.

(=P. paspalodes (Michx.)

Scribn.).

Couch paspalum, bankrot-
kweek.

Perennial; hydrophyte (root-
ing at the nodes), or rhizomatous,
or stoloniferous (mat-forming);

100-300 mm tall. Leaf blades
20-220 mm long; 3-8 mm wide. Spikelets 2. 5-3. 5 mm
long; 1.3-1. 5 mm wide. Racemes usually two; spikelets
arranged in two rows, glabrous, lanceolate, tips acute; lower
glume usually developed into a small triangular scale or
sometimes absent.

Flowering November to May. Always in or near salt or
fresh water on river banks, in vleis and along pan edges in
muddy soil, sand or black turf. Locally common. Biome;
Fynbos, Savanna, Grassland, and Nama-Karoo. Tropics
worldwide. Weed (pest in lands, difficult to eradicate).
Closely related to P . vaginatum, in which the lower glume
is absent or reduced to a rim. the upper glume glabrous and
the leaves less than 3 mm wide. Conflicting opinions exist
about the nomenclature and status of these two species but
leaf anatomical differences were found by Ellis (1974
Bothalia 1 1 : 235).

Description: Stapf 1898-1900 (371), Chippindall 1955
(389). Illustration: Chippindall 1955 (fig. 331). Voucher:
Jacobsz 4003. Smook & Gibbs Russell 2242. PRECIS code
9901070-00150.

Paspalum notatum Fluegge

Bahia grass, Bahia paspalum.

Perennial; long rhizomatous
and tufted (often decumbent);

100-600 mm tall. Leaf blades
60-240 mm long; 4-10 mm wide.

Spikelets 2. 8-3. 7 mm long;

2. 0-2. 8 mm wide. Rhizome well
developed, almost woody, clad
in overlapping leaf sheaths; racemes 2 (occasionally 3),
25-130 mm long; rachis often zig-zag; spikelets glabrous,
glossy, tips rounded, more than 1.5 times longer than wide
and arranged in two rows.

Flowering November to April. In high rainfall areas on
sandy or clayey soil, often in disturbed places and
cultivation. Infrequent. Naturalized from South America.
Biome: Fynbos and Grassland. Tropical Africa and tropical
America. Pasture (fodder, not extensively cultivated,
improved stains used for fodder for sheep), or erosion con-
trol (binding soil on bench terraces), or weed (tough and
aggressive invader of cultivation and disturbed areas).
Closely related to P. scrobiculatum, which has smaller
spikelets and a flat, almost leaf-like rachis.

Description: Webster 1987 (176), Chippindall & Crook
1976 (13). Hitchcock & Chase 1950 (583), Chippindall
1955 (389), Clayton et al. 1970-1982 (609). Illustration:
Hitchcock & Chase 1950 (fig. 1214). Voucher: Mogg
35370. PRECIS code 9901070-00200.

Paspalum scrobiculatum L.

(=P. commersonii Lam.) 2;

( =P . orbiculare Forst.) 2; (=P.
polystachyum R. Br.) 2.

Creeping paspalum, dronk-
gras.

Perennial; hydrophyte (occasi-
onally), or rhizomatous (short-

247

ly), or stoloniferous (sometimes), or tufted (loosely, erect
or decumbent); 100-700 mm tall. Leaf blades
150-200(-380) mm long; 6— 8(— 1 0) mm wide. Spikelets
2.0-2. 5 mm long; 1.8-2. 4 mm wide. Racemes 1-5,
30— 80(— 150) mm long; rachis linear and almost as wide as
the raceme; spikelets glabrous, ovate, arranged in two rows.

Flowering September to May. Moist, semi-swampy
areas or on fertile well-drained soils, often in disturbed
places and abandoned lands. Locally common. Biome: Fyn-
bos, Savanna, and Grassland. Old world tropics and
subtropics. Food and drink (domesticated as cereal in
India), or pasture (palatable fodder but seedheads subject
to ergot infection), or poisonous (when infected with ergot),
or weed (ruderal in damp areas). Closely related to P.
notatum , which has larger spikelets, a narrower rachis and
a very well developed rhizome.

Description: Stapf 1919 (573), Chippindall & Crook
1976 (14), Stapf 1898-1900 (370), Hitchcock & Chase
1950 (601), Chippindall 1955 (387). Illustration: Chippin-
dall 1955 (fig. 330), Clayton et al. 1970-1982 (fig. 142).
Voucher: Smook 1006. PRECIS code 9901070-00550.

Paspalum urvillei Steud.

structureless soils), or erosion control (sand binder at
coasts), or weed (in irrigation furrows and rice lands).
Closely related to P. distichum , which has a lower glume
developed into a small triangular scale or up to 1/2 the
spikelet length and the upper glume minutely hairy.
Conflicting opinions exist about the nomenclature and
status of these two species, however leaf anatomical
differences were found by Ellis (1974 Bothalia 1 1: 235).

Description: Stapf 1919 (570), Chippindall & Crook
1976 (16), Hitchcock & Chase 1950 (580), Chippindall 1955
(389), Clayton et al. 1970-1982 (609). Illustration: Hitch-
cock & Chase 1950 (fig. 1206). Voucher: Michelmore 162.
PRECIS code 9901070-00700.

Pennisetum Rich.

Amphochaeta Anderss., Beckeropsis Fig. & de Not.,
Catatherophora Steud., Eriochaeta Fig. & De Not,
Gymnotrix P. Beauv., Loydia Delile, Macrochaeta Steud.,
Penicillaria Willd., Pentastachya Steud., Sericura Hassk.

Vasey grass, langbeen pas-
palum.

Perennial; rhizomatous and
tufted (coarsely and erect);

100-2500 mm tall. Leaf blades
250-600 mm long; 4-15 mm
wide. Spikelets 1 .6-2.8 mm long;

1.2-1. 4 mm wide. Basal sheaths
densely hairy; ligule conspicuous, membranous, 2-9 mm
long; inflorescence with central axis 120-300 mm long;
racemes 10-30, closely spaced on axis; spikelets arranged
in four rows, fringed with white hairs to give a woolly
appearance.

Flowering October to April. Near water or in moist
places, along water furrows, roadsides and streambanks on
sandy loam. Common. Naturalized from South America.
Biome: Fynbos, Savanna, and Grassland. Tropics

worldwide. Domestic use (old inflorescences used as whisk
brooms for brushing lint), or pasture (cultivated for hay;
frost resistant, only occasionally susceptable to ergot,
young growth palatable and nutritious, but stalky when
when old). Resembles P. dilatatum, which has larger spike-
lets and fewer racemes.

Description: Webster 1987 (181), Chippindall & Crook
1976 (15), Hitchcock & Chase 1950 (595), Chippindall
1955 (387). Illustration: Chippindall 1955 (pi. 13(1)), Hitch-
cock & Chase 1950 (fig. 1246). Voucher: Liebenberg 8769.
PRECIS code 990 1 070-00600.

Paspalum vaginatum Swartz

Brak paspalum, seashore pas-
palum.

Perennial; hydrophyte (and
then rooting at the nodes below
the water level and branching
above), or rhizomatous (shortly),
or stoloniferous (mat-forming);

300-400(-600) mm tall. Leaf
blades 40-90 mm long; 2-3(-4) mm wide. Spikelets
3.0-^L5 mm long; 0.9-1. 5 mm wide. Internodes short,
branching at most nodes; racemes usually two; spikelets
arranged in two rows, glabrous, lanceolate with tips acute;
lower glume absent or reduced to a rim.

Flowering December to April. Near coasts, in or near
estuaries or rivers, also inland at water edges on sandy soils
but most often in saline water. Locally common. Biome:
Fynbos, Savanna, Grassland, and Desert. Worldwide in
tropics and subtropics. Potential pasture (on some brak and

Annual (rarely), or perennial; long-stoloniferous, or
caespitose, or decumbent. Culms 150-8000 mm high; her-
baceous; branched above, or unbranched above. Ligule a
fringed membrane to a fringe of hairs. Plants with her-
maphrodite florets. The spikelets of sexually distinct forms
on the same plant (peripheral spikelets of the glomerules
may be male-only), or all alike in sexuality. Plants with
hidden cleistogenes (e.g. P. clandestinum, which lacks
‘normal’ inflorescences), or without hidden cleistogenes.

Inflorescence a false spike, with clusters of spikelets on
reduced axes, or paniculate (the spikelets fascicled in false
spikes, in small groups or apparently solitary, but always
surrounded at their bases by reduced-branch bristles );
contracted (into false spikes); espatheate. Spikelet-bearing
axes disarticulating (but the main axis persistent ); falling
entire (the false spikes or spikelet-plus-bristle clusters
falling).

Spikelets with ‘involucres' of ‘bristles’ (these relatively
slender, basally free or scarcely united, by contrast with
Cenchrus). The ‘bristles' relatively slender, not spiny.
Female-fertile spikelets compressed dorsiventrally; falling
with the glumes, or not disarticulating (in cultivated forms).
Glumes two; very unequal (G1 often minute or vestigial);
awnless; very dissimilar, or similar (hyaline or
membranous). Proximal incomplete florets 1 ; paleate, or
epaleate, palea when present fully developed to reduced;
male, or sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes, or decidedly firmer than the glumes; smooth, or
striate; not becoming indurated (membranous to
subcoriaceous); hairy (near the margins), or hairless
(glabrous); having the margins lying flat and exposed on
the palea; with a clear germination flap; 5-7 nerved; entire;
awnless, or mucronate. Palea present; relatively long. Lodi-
cules when present 2; glabrous. Stamens 3. Ovary glabrous.
Fruit small; hilum short; embryo large.

Photosynthetic pathway. C4; NADP-ME (2 species);
XyMS-. PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. About 80
species. In warm regions. Helophytic, mesophytic, and
xerophytic; in shade and in open habitats (savanna,
woodland, weedy ground). Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. Indigenous species ( 8 ), naturalized species (4),
cultivated species (1).

References. 1. Chippindall. 1955. Gr. & Past. 2.
Brunken. 1977. Amer. J. Bot. 64: 161. 3. Clayton &
Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

248

Fig. 156. Pennisetum sphacelatum

1(0). Inflorescence hidden in uppermost leaf sheath ....

P. clandestinum

Inflorescence prominent 2

2(1). Inflorescence 200-500 mm by 30 mm; spikelet
clusters borne on stalk 3-6 mm long . P. glaucum
Inflorescence less than 250 mm long; spikelet clusters

not borne on a stalk 3

3(2). Inflorescence bristles 4-5 times as long as spikelet

4

Inflorescence bristles usually less than 2.5 times as

long as spikelet 6

4(3). Inflorescence a compound panicle, branches filiform
and usually in groups of 2-3; each spikelet

subtended by a single bristle P. unisetum

Inflorescence not a compound panicle, spikelets

subtended by about 30 bristles 5

5(4). Inflorescence bristles white, usually all plumose . .

P. villosum

Inflorescence bristles purple, only inner ones plumose
P. setaceum

6(3). Culms branched, often profusely 7

Culms not branched 9

7(6). Plants forming bamboo-like clumps up to 7500 mm

tall; only inner bristles loosely plumose

P. purpureum

Plants profusely branched, the tufts shorter than 1000

mm; all bristles either plumose or scabrid 8

8(7). Inflorescence lax; bristles plumose

P. foermerianum

Inflorescence dense; bristles not plumose

P. mezianum

9(6). Plants 1000-3000 mm tall; lowest leaf sheath with
numerous short transverse veins; inflorescence
200-300 mm long; spikelet involucre with up to 30

bristles P. glaucocladum

Plants 200-2000 mm tall; lowest leaf sheath without
numerous short transverse veins; inflorescence
50-250 mm long; spikelet involucre with up to 20
bristles 10

10(9). Most inflorescence bristles as long as spikelet,

usually straw-coloured 11

Most inflorescence bristles longer than spikelet,

straw-coloured to purple 12

I 1(10). Plants 800-2500 mm tall; inflorescence 120-250

mm long P. macrourum

Plants 400-900 mm tall; inflorescence 50-150 mm

long P. sphacelatum

1 2(10). Plants 500-2000 mm tall; inflorescence 70-220 mm
long; lower palea well developed; anther lobes

without a minute tuft of hairs at apex

P. natalense

Plants 200-800 mm tall; inflorescence 30-50 mm
long; lower palea absent; anther lobes with a
minute tuft of hairs at apex .... P. thunbergii

NOTE: P. glaucocladum, P macrourum , P. natalense, P.

sphacelatum, and P. thunbergii are all closely
related and may therefore be difficult to key out.

Pennisetum clandestinum Chiov.

Fig. 157.

Kikuyu grass.

Perennial; rhizomatous and
stoloniferous; 30-1200 mm tall.

Leaf blades 50-300 mm long; 3-7
mm wide. Spikelets 10-20 mm
long. Creeps vigorously by rhi-
zomes and stolons; culms closely
sheathed, with abundance of
bright green leaves; inflorescence partly enclosed in
uppermost leaf sheath; filamentous stigmas and the stamens
with long filaments are clearly visible at flowering time.

Flowering August to April. Requires high rainfall. Infre-
quent. Naturalized from the east African highlands. Biome:
Fynbos and Grassland. East Africa and introduced world
wide. Cultivated pasture, or weed, or ornamental (lawns).

Description: Chippindall & Crook 1976 (182), Chippin-
dall 1955 (444). Illustration: Chippindall 1955 (fig. 369).
Voucher: Smook 1165. PRECIS code 9901390-00300.

Pennisetum foermerianum Leeke

Perennial; tufted and rhizo-
matous; 200-600 mm tall. Leaf
blades 80-150 mm long; 2 mm
wide. Spikelets 4. 5-6.0 mm long;
1-2 mm wide. Rhizome stout,
creeping; culms branched usually
a few nodes up from base; inflor-
escence an interrupted panicle;
spikelet clusters open; involucral

249

bristles plumose, usually about 5 mm long.

Flowering December to April. Sandy soil and
mountainous areas. Locally common. Biome: Nama-Karoo.
Endemic. Maybe confused with Cenchrus ciliaris, which
has a dense, bristly false spike.

Description: Muller 1984 (208), Chippindall 1955 (447).
Illustration: Muller 1984 (fig. 103), Chippindall 1955(371).
Voucher: Smook 5128. PRECIS code 9901390-00500.

Pennisetum glaucocladum Stapf & C.E. Hubb.

Riverbank pennisetum.

Perennial; tufted; 1000-3000
mm tall. Leaf blades 300-650
mm long; 5-1.3 mm wide. Spike-
lets 3-6 mm long; 1 mm wide.

Lowest leaf sheath with numer-
ous transverse veins; leaf blades
widely spaced and often held
at right angles to culm; inflorescence 200-300 mm long;
involucral bristles up to 30 and most of them longer than
the spikelets; spikelets similar to P. macrourum.

Flowering January to May. River banks and wet areas.
Infrequent. Biome: Savanna. Tropical Africa. Domestic use
(thatching). This species grows much larger than P.
macrourum and occurs more commonly along river banks.

Description: Chippindall 1955 (441). Illustration:
Chippindall & Crook 1976 (184). Voucher: Tinley 437.
PRECIS code 9901390-00600.

Pennisetum glaucum (L.) R. Br.

( =P . americanum (L.) Leeke
subsp. americanum) 3; ( -P .
albicauda Stapf & C.E. Hubb.)

2; (-P. echinurus (K. Schum.)

Stapf & C.E. Hubb.) 2; (=P.
nigritarum (Schlecht.) Dur. &

Schinz) 2; ( =P . typhoides
(Burm. f.) Stapf & C.E. Hubb.)

2; ( =Setaria lutescens (Wiegel)

F T. Hubb.) 3.

Pearl millet.

Annual; tufted; 2000-3000 mm tall. Leaf blades
300-500 mm long; 20-50 mm wide. Spikelets about 7 mm

long; 5 mm wide. Inflorescence a false spike 200-500 mm
long, 30 mm wide; each spikelet cluster on a hairy stalk
about 5 mm long; involucral bristles shorter to as long as
the spikelets, only the inner bristles plumose.

Flowering January to April. Widely cultivated in semi-
arid tropics, may grow with as little as 250 mm rainfall per
annum. Biome: Savanna and Grassland. Tropical and sub-
tropical Africa. Cultivated pasture.

Description: Chippindall 1955 (447). Illustration: Chip-
pindall 1955 (fig. 372). Voucher: Hardy, Retief and Herman
5315. PRECIS code 9901390-00650.

Pennisetum macrourum Trin.

Beddinggras.

Perennial; tufted; 800-2500
mm tall. Leaf blades 250-600
mm long; 4-1 1 mm wide. Spike-
lets 4-6 mm long; 1 mm wide.

Rhizome creeping, often branch-
ed; inflorescence light green or
straw-coloured, often tinged
with purple, 120-250 mm long; involucres of up to 20
bristles which are mostly as long as the spikelets, with one
bristle longer than the rest; lower glume minute or absent,
upper glume 1/4- 1/3 the length of the spikelet; lower floret
reduced to a lemma; female-fertile (upper) lemma similar
to lower lemma.

Flowering November to May. Near streams or damp
places. Common. Biome: Fynbos, Savanna and Grassland.
Tropical Africa. Clayton 1982 (690) regards P. macrourum
as a polymorphic species including P. natalense , P.
glaucocladum and 22 other tropical species.

Description: Chippindall 1955 (442), Clayton et al.
1970-1982 (689). Voucher: Smook 3167. PRECIS code
9901390-00700.

Pennisetum mezianum Leeke

( =P . stapfianum F. Bol.).

Perennial; tufted and rhizo-
matous; to 600 mm tall. Leaf
blades to 10 mm long; 3 mm
wide. Spikelets 3^1 mm long; 1
mm wide. Rhizome short and
woody; culms profusely branched
and shrubby; inflorescence dense,

10-30 mm long; involucral bristles not plumose.

Flowering March to April. Prefers plains with impeded
drainage. Infrequent. Biome: Savanna. Tropical and sub-
tropical Africa. P. stapfianum is here considered to belong
to this species, as the specimens available in Pretoria show
no morphological differences.

Description: Clayton et al. 1970-1982 (686). Voucher:
Smook 5117. PRECIS code 9901390-00800.

Pennisetum natalense Stapf

Suurbuffelsgras.

Perennial; tufted; 500-2000
mm tall. Leaf blades 100-400
mm long; 3-8 mm wide. Spike-
lets 2. 5-3. 5 mm long; 1 mm wide.

Inflorescence 70-220 mm long;
most involucral bristles twice as
long as spikelets; lower floret
male, lower palea almost as long as lemma.

Flowering February to June. Forms large tufts in water
on river banks and vleis. Common. Biome: Savanna and
Grassland. Similar toP. macrourum , which is a larger plant
and lacks a lower palea.

Description: Chippindall 1955 (440). Illustration: Chip-
pindall 1955 (fig 365). Voucher: Strey 10968. PRECIS code
9901390-00900.

250

Pennisetum purpureum Schumach.

Napier fodder, elephant grass,
mfufu.

Perennial; tufted; 1800-7500
mm tall. Leaf blades 30^100 mm
long; 10-30 mm wide. Spikelets
4. 5-7.0 mm long; 1 mm wide.

Often forms tall, bamboo-like
clumps; involucre of up to 40
bristles, the inner few are loosely plumose towards the base;
lower floret usually male; in other spikelet characters
similar to P. macrourum.

Flowering January to June. Riverine sites, valley floors
and forest margins, with a preference for rich soils. Infre-
quent. Naturalized from tropical Africa. Biome: Savanna
and Grassland. Tropical Africa. Domestic use (fencing
reeds), or pasture (cultivated). Many cultivars and hybrids
occur; a well known example is Bana grass, a cross withP.
glaucum.

Description: Chippindall 1955 (443), Clayton et al.
1970-1982 (677). Illustration: Chippindall 1955 (fig. 368).
Voucher: Smook 1794. PRECIS code 9901390-01 100.

Pennisetum setaceum (Forssk.) Chiov.

Fig. 158. PI. 145.

Fountain grass, pronkgras.

Perennial; tufted; 600-1000
mm tall. Leaf blades 20-40 mm
long; 1-2 mm wide. Spikelets
4. 0-6. 5 mm long; 3 mm wide. In-
florescence purple, 100-250 mm
long; involucral bristles, mostly
about 20 mm long, 4-5 times as

Fig. 158. Pennisetum setaceum

long as the spikelets, the inner bristles plumose.

Flowering November to July. A ruderal on stony slopes
and dry open places. Infrequent. Naturalized from north
Africa. Biome: Fynbos, Savanna, and Nama-Karoo. To
north Africa, Arabia. Garden ornamental.

Description: Chippindall 1955 (447). Illustration:
Chippindall & Crook 1976 (183). Voucher: Retief & Reid
503. PRECIS code 9901390-01200.

Pennisetum sphacelatum (Nees) Dur. & Schinz

(=P. sphacelatum (Nees)

Dur. & Schinz var. sphacelatum)

3; (=P. sphacelatum (Nees) Dur.

& Schinz var. tenuifolium
(Hack.) Stapf) 3.

Bulgras.

Perennial; tufted; 400-900
mm tall. Leaf blades 100-400 mm long; 2-3 mm wide.
Spikelets about 3 mm long; 1 mm wide. Leaves often fili-
form; inflorescence straw-coloured, 50-150 mm long, hairy
for some distance below inflorescence; most involucral
bristles equalling to twice as long as the spikelets.

Flowering November to April. Wet areas, vleis, usually
hillsides, moist or clay soil. Common. Biome: Grassland.
Similar to P macrourum , which is a much larger plant.

Description: Chippindall 1955 (442). Illustration: Chip-
pindall 1955 (fig. 366). Voucher: Smook 4692. PRECIS
code 9901390-01300.

Pennisetum thunbergii Kunth

Thunberg’s pennisetum.

Perennial; tufted; 200-800
mm tall. Leaf blades 100^400
mm long; 4—7 mm wide. Spike-
lets 3 mm long; 1 mm wide. In-
florescence 30-50 mm long, pur-
ple; lower glume absent; upper
glume 1/4-1/5 as long as spikelet;
lemmas mucronate to nearly awned; anther lobes with a
minute tuft of hairs at apex; other spikelet characters as for
P. macrourum.

Flowering October to June. Grows in wet places, river
banks, vleis. Common. Biome: Fynbos and Grassland.
Uplands of African tropics and also occurs in Yemen and
Sri Lanka.

Description: Chippindall 1955 (443), Clayton et al.
1970-1982 (687). Illustration: Chippindall & Crook 1976
(184), Chippindall 1955 (fig. 367). Voucher: Smook 2575.
PRECIS code 9901390-01700.

Pennisetum unisetum (Nees) Benth.

( =Beckeropsis uniseta (Nees)

K. Schum.) 3.

Natal grass, silky grass

Perennial; tufted; 900-2400
mm tall. Leaf blades 200^150
mm long; 5-10 mm wide. Spike-
lets 2. 5-3. 5 mm long; 1 mm wide.

Inflorescence a compound pani-
cle, branches filiform, usually in
groups of 2-5; spikelets subtended by a single bristle 3-4
times as long as spikelet.

Flowering March to June. Near water and in shady
places. Common. Biome: Grassland. Cultivated pasture.
Sometimes placed in a separate genus, Beckeropsis,
because of the single involucral bristle and the reduction of
the inflorescence to a single raceme; this species also
apparently lacks the germination flap characteristic of
Pennisetum.

251

Description: Chippindall 1955 (448). Illustration: Chip-
pindall 1955 (fig. 373). Voucher: Nicholson 1750. PRECIS
code 9901390-01750.

Pennisetum villosum R. Br. ex Fresen.

Feathertop.

Perennial; tufted; 200-900
mm tall. Leaf blades 8-15 mm
long; 3 mm wide. Spikelets 9-14
mm long; 4 mm wide. Inflores-
cence usually white, 40-100 mm
long; involucral bristles nearly all
plumose, some 4-5 times as long
as spikelet, most of them about 30 mm long.

Flowering January to May. Roadsides and disturbed
areas. Infrequent. Naturalized from Ethiopia. Biome: Sa-
vanna and Grassland. North and central Africa. Established
ornamental.

Description: Chippindall 1955 (446). Voucher: Smook
2127. PRECIS code 9901390-01800.

Pentameris P. Beauv.

Perennial; caespitose. Culms 250-2000 mm high;
woody and persistent, or herbaceous (from a woody or
suffrutescent base); branched above, or unbranched above.
Leaf blades linear to linear-lanceolate. Ligule a fringe of
hairs.

Inflorescence paniculate; open, or contracted
(sometimes scanty); non-digitate (branching sometimes
trichotomous); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 13-25 mm long (rarely to 30 mm)\
compressed laterally; disarticulating above the glumes.
Glumes two; more or less equal; much exceeding the
spikelets ; awned (setaceously acuminate), or awnless;
similar (ovate-lanceolate, thin). All florets usually female-
fertile only; distal incomplete florets occasionally present,
merely underdeveloped; proximal incomplete florets
absent.

Female-fertile florets 2 (rarely 1). Lemmas similar in
texture to the glumes to decidedly firmer than the glumes;
hairy (the hairs in longitudinal rows between the veins);
without a germination flap; 7 or 9 (-11) nerved; incised;
deeply cleft\ awned. Awns i; median and lateral (the lateral
lemma lobes each with a 1-7 mm bristle from the inner side,
more or less adnate below). The median awn different in
form from the laterals; from the sinus; geniculate (near the
middle); much longer than the body of the lemma. Palea
present (hairy); relatively long; 2-nerved. Lodicules 2;
fleshy; ciliate, or glabrous. Stamens 3. Ovary hairy (with a
deciduous apical tomentum, of branched hairs). Fruit small
(3 mm); subglobose; hilum long-linear; pericarp free;
embryo small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 7 species. South Africa. Mesophytic; in open
habitats; glycophytic. Cape Province. 7 indigenous species.

References. 1. Schweickerdt. 1938. Feddes Reprium 42:
91. 2. Chippindall. 1955. Gr. & Past. 3. Ellis. 1985a.
Bothalia 15: 561-566. 4. Ellis. 1985b. Bothalia 15:
567-571. 5. Ellis. 1985c. Bothalia 15: 573-578. 6. Ellis.
1985d. Bothalia 15: 579-585. 7. Ellis. 1986. Bothalia 16:
235-241. 8. Barker. 1986. Bothalia 16: 65-69.

Species treatment by N.P. Barker.

252

1(0). Spikelets with one floret .... P. sp. 2 (=Ellis 2546)

Spikelets with two florets 2

2(1). Leaf sheaths with purple to dark brown or black
auricles; lemma lobes truncate, free from lateral

bristle P. thuarii

Leaves without dark auricles; lemma lobes acute or
acuminate, adnate to lateral bristle for half or all
their length 3

3(2). Panicle lax, globose, 170-300 mm long; basal leaf
sheaths 120 mm long or longer, clustered around
and free from the culm base; culm plus

inflorescence usually taller than 1200 mm

P. longiglumis

Panicle contracted, lanceolate, up to 150 mm long;
basal leaf sheaths seldom longer than 120 mm,
partially free or appressed to the culm base; culm
plus inflorescence up to 1200 mm in height ... 4
4(3). Glumes usually with three nerves at the very base;
lodicules ciliate, cilia as long or longer than

lodicule body P. obtusifolia

Glumes 1 -nerved; lodicules glabrous or shortly
ciliate, sometimes with one or two arm-like

extensions 5

5(4). Leaves flexible, pubescent or sometimes glabrous,
rolled, folded or open; glumes 12-16 mm long . .

P. dregeana

Leaves rigid, glabrous, cylindrical, permanently
infolded or tightly rolled with narrow, deep adaxial

groove; glumes 15 mm or longer 6

6(5). Leaf blades straight to falcate, 30-350 mm long,
acicular, permanently infolded, sometimes
pungent; leaf sheaths tightly appressed to culm;

found at all altitudes P. macrocalycina

Leaf blades falcate to curled (especially when dead),
up to 1 10 mm long, tightly rolled, strongly pungent;
leaf sheaths not tightly appressed to culm; found at
high altitudes only P. sp. 1 (=Esterhuysen 11115)

Pentameris dregeana Stapf

Perennial; decumbent to tuft-
ed; 300-1000 mm tall. Leaf
blades to 250 mm long; to 1 .5 mm
wide. Spikelets 12-17 mm long
(excluding awns); to 10 mm wide.

Culms often branched basally;
leaf sheaths pubescent to woolly,
without auricle, sheath mouth
densely bearded; blades open,
folded or rolled, flexible, rarely glabrous, usually densely
pubescent, especially near the base; panicle lanceolate,
loosely contracted, 50-1 10 mm long; spikelets 2-flowered;
glumes 12-16 mm long, 1-nerved, glabrous or pubescent;
lemma lobes acuminate, partially fused to lateral bristle;
lodicules glabrous or ciliate, sometimes with one or two
arm-like extensions.

Flowering September to December. In rock crevices and
coarse sandy soil of the Cape fold mountains. Infrequent to
common (after fire). Biome: Fynbos. Endemic. Domestic
use (used for bedding in mountain huts and caves). A
widespread Fynbos species which is quite variable. Ellis
(1986) found three anatomical forms within this species.

Description: Stapf 1898-1900 (515), Chippindall 1955
(253). Illustration: Chippindall 1955 (fig. 224 (spikelet
only)). Voucher: Compton 13952. PRECIS code

9902080-00100.

Pentameris longiglumis (Nees) Stapf

Perennial; tufted; to 1700 mm
tall. Leaf blades to 550 mm long;
to 4 mm wide. Spikelets 16-25
mm long; to 10 mm wide. Culms
erect, unbranched; basal sheaths
persistent, 120 mm or longer, 10
mm or more wide, loose or free
from culm; leaf sheaths without
auricles; leaf blades glabrous,
rigid, rolled; panicle lax, globose, 170-300 mm long; spike-
lets 2-flowered; glumes 15-25 mm long, 1-nerved,
glabrous; lemma lobes acuminate, adnate to lateral bristle
for most of their length; lodicules glabrous.

Flowering September to December. Moist, rocky slopes.
Rare. Biome: Fynbos. Endemic. There are two

morphologically and geographically distinct varieties,
separable on the size of the floral parts. Ellis (1985b)
considers this species to be anatomically distinct, but allied
to P. macrocalycina.

Description: Stapf 1898-1900 (514), Chippindall 1955
(253). Voucher: Taylor 7231. PRECIS code 9902080-
00200.

Pentameris macrocalycina (Steud.) Schweick.

Fig. 159.

(=P. speciosa Nees) 1.

Perennial; tufted; 400-1100
mm tall. Leaf blades 30-350 mm
long; to 1.5 mm wide. Spikelets
17-25 mm long; 5-10 mm wide.

Culms branching basally: leaf
sheaths tightly appressed to culm,
bearded at the mouth, auricles absent; leaf blades rigid,
erect, glabrous, acicular, sometimes pungent, permanently
folded into a cylinder with deep adaxial groove when seen
in cross section; panicle contracted, lanceolate, 60-120 mm
long; spikelets 2-flowered; glumes 16-24 mm long, 1-
nerved, glabrous; lemma lobes acuminate, adnate to lateral
bristle for most of their length; lodicules glabrous.

Flowering September to December. In rock crevices and
stony or sandy soils. Infrequent to common (after fire).
Biome: Fynbos. Endemic. Plants are vegetatively variable,
as those growing soon after fire are rigid and robust,
whereas plants from older Fynbos are softer. Ellis (1985d)
has found this species to be anatomically quite constant.

Description: Stapf 1898-1900 (515), Chippindall 1955
(252). Illustration: Chippindall 1955 (fig. 223). Voucher:
Esterhuysen 23236. PRECIS code 9902080-00300.

Pentameris obtusifolia (Hochst.) Schweick.

(=P. squarrosa Stapf) 1.

Perennial; decumbent scramb-
ler; seldom more than 500 mm
tall. Leaf blades 50-150 mm
long; to 4 mm wide. Spikelets
20-26 mm long; 5-10 mm wide.

Culms much branched, produced
from a woody base, decumbent,
to 900 mm long, protected by the remains of numerous
overlapping, leaf sheaths from base to growth point; leaf
sheaths appressed to culms, without auricles; leaf blades
short, rigid, rolled, present only at branch tips; panicle
contracted, lanceolate, 70-130 mm long; spikelets 2-
flowered; glumes pale, 18-25 mm long, generally 3-nerved
at the base; lemma lobes almost fully adnate to lateral
bristle; lodicules densely long-ciliate at apex.

253

Flowering January to April. Lower slopes of Hottentots
Holland and Kogelberg mountains. Locally common.
Biome: Fynbos. Endemic. Anatomical studies (Ellis
1985b), seed morphology (Barker 1986) and floral
morphology indicate that this species does not belong in
this genus. However, correct generic placing currently
impossible because acceptable generic limits have not been
established in the subfamily.

Description: Stapf 1898-1900 (536). Voucher: Barker
331. PRECIS code 9902080-00400.

Pentameris thuarii Beauv.

PI. 146.

Perennial; tufted; 350-2000
mm tall. Leaf blades to 500 mm
long; to 10 mm wide. Spikelets
16-22 mm long; 5-8 mm wide.

Culms single or branched; leaf
sheaths with purple to dark brown
or black auricles; leaf blades open
or folded, glabrous to sparsely
pubescent; panicle lax, globose,

70-220 mm long; spikelet 2-flowered; glumes 16.0-21.5
mm long, 1 -nerved, glabrous; lemma lobes truncate, almost
wholly free from lateral bristle; lodicules glabrous.

Flowering September to December. In seeps and along
river banks. Locally common (in damp habitats). Biome:
Fynbos. Endemic. This species has been divided by Nees
into two varieties based on overall plant size, but this
character is too variable to be useful. It is anatomically
distinct from the other species in the genus (Ellis 1985c).

Description: Stapf 1898-1900 (513), Chippindall 1955
(252). Illustration: Chippindall 1955 (fig. 222 (spikelet
only)). Voucher: McDonald 816. PRECIS code

9902080-00500.

Pentameris sp. 1 (=Esterhuysen 11115)

Perennial; tufted; 250^470
mm tall. Leaf blades 60-1 10 mm
long; to 1.5 mm wide. Spikelets
15-21 mm long; to 10 mm wide.

Culms sometimes branched and
somewhat decumbent; leaf
sheaths loosely appressed to
culm, without auricles; leaf
blades tightly rolled, falcate to
curled, especially when dead, strongly pungent; panicle
contracted, 30-83 mm long; spikelets 2-flowered; glumes

I - nerved, generally glabrous, 15.5-20.0 mm long; lemma
lobes acute to acuminate, adnate to lateral bristle for about
half their length; lodicules glabrous.

Flowering September to December. High altitude
mountains. Locally common (Hottentots Holland, Hexrivier
and Riviersonderend mountains). Biome: Fynbos. Endemic.
This species, included in P . obtusifolia by PRE in the past,
is anatomically similar to P . macrocalycina (Ellis 1985d).
It is distinguished from this latter species by the rolled but
not permanently folded, falcate leaf blades, and leaf sheaths
which are loose to free from the culm. This new species
consists of two morphological forms, distinguished by
glume vesture.

Voucher: Esterhuysen 11115. PRECIS 9902080-99999.
Pentameris sp. 2 (=Ellis 2546)

Perennial; decumbent to tuft-
ed; about 400 mm tall. Leaf
blades to 130 mm long; to 1 mm
wide. Spikelets 11-13 mm long;

5-8 mm wide. Culms slender, to
650 mm long; leaf sheaths
without auricles; leaf blades
filiform; panicle 40-60 mm long;
spiklets 1 -flowered; glumes

II- 12 mm long, 1 -nerved, glabrous; lemma lobes

acuminate, almost fully adnate to lateral bristle; lodicules
glabrous or minutely ciliate.

Flowering September to December. Damp south facing
cliffs of Cape fold mountains. Infrequent. Biome: Fynbos.
Endemic. Although only having one floret, this species
belongs in the genus on the basis of seed characters, the
caryopsis being an achene with the characteristic apical
appendages.

Voucher: Ellis 2546. PRECIS code 9902080-99999.

Pentaschistis (Nees) Spach

Achneria Benth., Afrachneria Sprague, Poagrostis
Stapf.

Perennial (usually), or annual (less commonly); usually
caespitose. Culms 100-1500 mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear
to lanceolate (or filiform, often with stalked or saucer-
shaped glands); rolled (usually), or flat. Ligule a fringe of
hairs.

Inflorescence paniculate ( the branches often with
glands ); open, or contracted (sometimes spicate); espathe-
ate. Spikelet-bearing axes persistent.

Fig. 160. Pentaschistis airoides subsp. airoides

254

Spikelets 1-19 mm long; compressed laterally; disartic-
ulating above the glumes. Callus short. Hairy callus
present. Glumes two; more or less equal; about equalling
the spikelets to much exceeding the spikelets ; awnless;
similar (narrow to lanceolate, green or scarious, rarely
hyaline, shining, often with glands). All florets usually
female-fertile only; distal incomplete florets occasionally
present, merely underdeveloped; proximal incomplete
florets absent.

Female-fertile florets 2 (rarely 1 - Poagrostis). Lemmas
similar in texture to the glumes (membranous); incised
(bifid, rarely 3-4-fid)\ hairy, or hairless; without a germin-
ation flap; 5-7 nerved; not deeply cleft ; awnless, or
mucronate, or awned (generally awned from the central
sinus, and with a point or straight awn on each lobe). Awns
when present 1 (rarely), or 3 (usually), or 5 (rarely)',
median, or median and lateral (usually). The median awn
different in form from the laterals (when laterals present,
they are inserted in the sinus and partially fused to the
lateral lemma lobes); from the sinus; usually geniculate;
much longer than the body of the lemma. Palea present; rel-
atively long; 2-nerved. Lodicules 2; fleshy; glabrous (or
with bristles). Stamens 3. Ovary glabrous. Fruit small;
hilum short (but linear-oblong); pericarp free, or fused, or
loosely adherent; embryo large to small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x - 7, 10, and 13. Arundinoideae; Danthonieae.
About 65 species. Africa, Madagascar. Xerophytic, or
mesophytic; in open habitats; glycophytic. Namibia,
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 57 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1969. Kew Bull. 23: 294. 3. Linder & Ellis. 1989. Contr.
Bol. Herb. 12.

Species treatment by H.P. Linder & R.P. Ellis.

Key to species groups

Plants annual Key A

Spikelets without awns Key B

Plants with stalked glands Key C

Plants with villous bases Key D

Plants without stalked glands Key E

Key A Plants annual

1(0). Spikelets 15-18 mm long P. triseta

Spikelets 2. 5-5.0 mm long 2

2(1). Lemmas without awns P. capillaris

Lemmas with awns 3

3(2). Leaves with slender long aristae P. aristifolia

Leaves without aristae 4

4(3). Anthers 0.3 mm long, plants very fine, with slender
leaves P. airoides subsp. airoides

Anthers 0. 5-3.0 mm long, plants more robust

P. patula

Key B. Spikelets without awns

1(0). Leaves borne on erect aerial stems; inflorescence

either a spike or with spikelets reflexed 2

Leaves borne basally; inflorescence always an open

panicle, spikelets not reflexed 3

2(1). Inflorescence a dense spike; plants without glands .

P. ecklonii

Inflorescence a panicle with reflexed spikelets; glume

keels glandular P. reflexa

3(1). Inflorescence nodes glabrous 4

Inflorescence nodes villous 8

4(3). Spikelets usually 1 -flowered, 2. 5-3.0 mm long . . .

P. pusilla

Spikelets always 2-flowered, 3-7 mm long .... 5
5(4). Spikelets 3 mm long; plants mat-forming; leaves stiff,

2-A mm broad, less than 10 mm long

P. microphylla

Spikelets 3. 5-7.0 mm long; plants caespitose; leaves
soft, 1-6 mm broad and more than 10 mm long . 6
6(5). Plants with flat woody bases, with scattered culms

rising through the vegetation; often with

conspicuous glands P. ampla

Plants with compact, caespitose bases; with
inconspicuous linear pedicel glands 7

7(6). Spikelets 3. 5-5.0 mm long; plants often glabrous . .

P. aurea subsp. aurea

Spikelets 6-7 mm long; plants usually softly villous

P. aurea subsp. pilosogluma

8(3). Glumes obtuse, the apex finely brown-puberulous;

plants without glands P. malouinensis

Glumes acute, glabrous; plants with stalked glands

9

9(8). Plants annual; spikelets 3 mm long . . . P. capillaris
Plants perennial; spikelets 3. 5-6.0 mm long .... 10

10(9). Plants caespitose, usually villous; leaves to 1 mm

wide P. setifolia

Plants mat-forming, glabrous; leaves 2-4 mm wide

P. galpinii

Key C. Plants with stalked glands

1 (0). Lemmas without awns 2

Lemmas with awns 8

2(1). Inflorescence a dense spike P. ecklonii

Inflorescence an open panicle 3

3(2). Spikelets reflexed P. reflexa

Spikelets not reflexed 4

4(3). Plants annual P. capillaris

Plants perennial 5

5(4). Plants caespitose; leaves linear, more than 20 mm

long 6

Plants short, mat-forming with creeping rhizomes;

leaves less than 20 mm long 7

6(5). Glumes finely acute to acuminate; plant base a flat

woody disc P. ampla

Glumes obtuse to acute; plant base contracted, not

forming a flat woody disc P. setifolia

7(5). Inflorescence nodes glabrous; inflorescence staying

open after flowering P. microphylla

Inflorescence nodes villous; inflorescence contracting

after flowering P. galpinii

8(1). Lateral awns included within the glumes 9

Lateral awns as long as or exserted from the glumes

13

9(8). Central awn 0-5 mm long 10

Central awn 6-12 mm long 11

10(9). Plants glabrous; from the Drakensberg

P. galpinii

Plants puberulous to villous; from Namaqualand

P. tomentella

1 1(9). Plants annual; nodes with rings of bristles

P. patula

Plants perennial 12

12(1 1). Leaves rigid, setaceous, stiffly erect .... P. lima
Leaves flaccid, generally expanded . P. oreodoxa
13(8). Plants annual or weakly perennial; anthers 0. 3-1.0

mm long 14

Plants perennial; anthers 1.5-5. 0 mm long ... 15
14(13). Plants weakly perennial; inflorescence nodes

villous P. airoides subsp. jugorum

Plants annual; inflorescence nodes glabrous or

puberulous P. airoides subsp. airoides

15(13). Leaves cauline 16

Leaves basal 19

16(15). Spikelets 3. 5^4. 5 mm long; old leaves turning pink;

plants to 30 mm tall P. densifolia

Spikelets 5-7 mm long; old leaves drying grey or

pink; plants more than 30 mm tall 17

17(16). Lateral awns 5-6 mm long; central awn 1 1-14 mm

long P. papillosa

Lateral awns 3 mm long; central awn 8-1 1 mm long
18

255

18(17). Plants cushion-forming; lateral awns as long as the
glumes; anthers 2. 3-2. 8 mm long . . P. aspera
Plants loosely caespitose; lateral awns exserted
from the glumes; anthers 1. 8-2.0 mm long . . .

P. barbata

19(15). Spikelets evenly scattered in an expanded
inflorescence, with the pedicels longer than the

spikelets 20

Spikelets clustered in an expanded inflorescence,
with the pedicels as long as or shorter than the

spikelets 22

20( 19). Spikelets 6-7 mm long; awn 9-1 3 mm long .. 21
Spikelets 4. 5-5.0 mm long; awn 7 mm long ....

P. longipes

21(20). Plants from Natal to Kenya; glands minute, only on

the keels of glumes P. natalensis

Plants from the southwestern Cape; glands variable

P. rupestris

22(19). Glands sunken, scattered on the backs of the U-

shaped leaves P. glandulosa

Glands stalked, usually restricted to veins and leaf

margins 23

23(22). Lemma length 1.5-2. 9 mm 24

Lemma length 3-4 mm 27

24(23). Spikelets 3-5 mm long; awns 6-7 mm long ....

P. pallida

Spikelets 5-7 mm long; awns 7-12 mm long . 25
25(24). Leaves rigid, less than 3 mm wide, usually not

glandular along the margins P. pallida

Leaves flaccid, more than 3 mm wide, usually

densely glandular along the margins 26

26(25). Lateral awns exserted from the glumes; basal

sheaths white; lemmas 2. 8-3.0 mm long

P. barbata

Lateral awns as long as the glumes; basal sheaths
soon decaying; lemmas 2. 0-2. 8 mm long ....

P. veneta

27(23). Inflorescence contracted; glumes hyaline or white;

a coastal dune or limestone plant . . . P. pallida
Inflorescence open; glumes usually greenish,

brownish or purplish; not on dunes 28

28(27). Glumes acute; restricted to the Drakensberg ....

P. oreodoxa

Glumes acuminate; from southern and western

Cape Province 29

29(28). Leaves usually rolled, less than 2 mm wide

P. cirrhulosa

Leaves always expanded, 2-6 mm wide 30

30(29). Spikelets 7-8 mm long; plants glabrous or with fine

hairs P. rupestris

Spikelets 5. 0-6. 5 mm long; plants coarsely hairy
with cushion-based hairs P. veneta

Key D. Plants with villous bases

1(0). Awns 15-25 mm long 2

Awns 5-12 mm long 6

2(1). Old leaf blades expanded and recurved; blades

sometimes rigid and pungent P. pungens

Old leaf blades rolled and shriveled; blades never

pungent 3

3(2). Glumes 12-20 mm long, straw-coloured to brown;

awns 20-25 mm long 4

Glumes 8-12 mm long, silvery; awns 10-20 mm long

5

4(3). Plants with horizontal rhizomes; lemmas 5. 5-7.0 mm

long P. aristidoides

Plants without rhizomes; lemmas 3—4 mm long . . .

P. velutina

5(3). Leaf blades glabrous; base of plant stoloniferous . .

P. argentea

Leaf blades hairy; base of plant compact

P. viscidula

6(1). Leaves rolled, 0.5- 1.5 mm wide 7

Leaves expanded, 1-6 mm wide 8

7(6). Plants without glands; spikelets 10-12 mm long . . .

P. pyrophila

Plants with glandular inflorescences; spikelets 6-7

mm long P. lima

8(6). Plants forming stout tussocks; leaves to 6 mm wide;

spikelets 7-8 mm long P, rupestris

Plants loosely caespitose; leaves 1-4 mm wide;
spikelets 5. 0-6. 5 mm long P. pallida

Key E. Plants without stalked glands

1(0).

2(1).

3(2).

4(3).

5(4).

6(5).

7(2).

8(7).

Plants annual P. aristifolia

Plants perennial 2

Lemmas without awns 3

Lemmas with awns 7

Inflorescence often spike-like; glumes apically blunt,

brown-puberulous P. malouinensis

Inflorescence a panicle; glumes acute, glabrous or

scaberulous 4

Spikelets usually 1 -flowered, 2. 5-3.0 mm long . . .

P. pusilla

Spikelets 2-flowered, 4-7 mm long 5

Plants with flat woody bases P. ampla

Plants with compact caespitose bases 6

Spikelets 3. 5-5.0 mm long; plants often glabrous . ..

P. aurea subsp. aurea

Spikelets 6-7 mm long; plants usually softly villous

P. aurea subsp. pilosogluma

Inflorescence linear or densely contracted 8

Inflorescence an open, hemispherical panicle ... 19
Spikelets 3-6 mm long, if more than 6 mm long then
the leaves are finely villous (P. calcicola var.

hirsuta) 9

Spikelets 7-15 mm long; leaves usually glabrous . .

11

9(8). Lemmas with 5-9 awns P. heptamera

Lemmas with 3 awns 10

10(9). Awn 10-12 mm long; from South Africa

P. calcicola subsp. hirsuta

Awn 6 mm long; from South or northeast Africa

P. pallida

1 1(8). Sheaths, or at least the sheath mouth, hairy

P. eriostoma

Sheaths villous or glabrous 12

12(11). Leaf margins thickened; spikelets ivory-coloured

P. curvifolia

Leaf margins not thickened; spikelets various . 13
13(12). Leaves densely villous on inside down whole length

- if broken the strands clearly visible

P. basutorum

Leaves not densely hairy on the whole length of the

inside 14

14(13). Base of plant swollen and densely hairy; awns

15-20 mm long P. argentea

Base of plant not swollen, more or less glabrous;

awns 8—1 5 mm long 15

15( 14). Inflorescence with 10-100 spikelets 16

Inflorescence with 100-300 spikelets 18

16(15). Inflorescence linear, 10-20 mm wide; lemma lobes
half as long as the lemma, ca. 2 mm long; from

the Drakensberg P. praecox

Inflorescence linear to contracted, 15-30 mm wide;
lemma lobes less than half as long as the lemma,
ca. 1 mm long; from the southern and western

Cape Province 17

17(16). Spikelets 10-12 mm long P. pyrophila

Spikelets 7-8 mm long P. rigidissima

18(15). Lateral awns 4-5 mm long, as long as the glumes;

sheath mouth glabrous P. tortuosa

Lateral awns 5. 5-6.0 mm long, longer than the
glumes; sheath mouth villous . . . P. eriostoma

19(7). Leaves rolled, setaceous 20

Leaves expanded 32

20(19). Lateral awns 0.1-3. 0 mm long 21

Lateral awns 3.5-15.0 mm long 26

21(20). Lateral awns 0. 1-1.0 mm long; central awn 3-4 mm

long P. holciformis

Lateral awns 1-3 mm long; central awn 5-12 mm
long 22

256

22(21). Leaves rigid, pungent; plants forming spiny tufts

P. rigidissima

Leaves flaccid, not pungent; plants soft 23

23(22). Lemmas more than 3 mm long; leaves stiffly erect,
glabrous, usually with a sheath of burnt-off basal

fibres P. tysonii

Lemmas to 3 mm long; leaves curly, usually
villous, rarely with a sheath of burnt-off basal

fibres 24

24(23). Inflorescence with 50-100 spikelets; from the

eastern Transvaal P. chippindalliae

Inflorescence with 5-50 spikelets; from the western

Cape Province 25

25(24). Central awn 5-6 mm long; plants with a strong base

P. montana

Central awn 7-1 0 mm long; plants with weak bases

P. alticola

26(20). Glumes often with cushion-based hairs; bases of
plants weak, with decaying brown leaf remnants

P. rosea subsp. rosea

Glumes without cushion-based hairs; bases of
plants woody, usually with white sheaths . . 27

27(26). Inflorescence with fewer than 30 spikelets

P. colorata

Inflorescence with more than 50 spikelets .... 28

28(27). Awns 15-25 mm long 29

Awns 5-1 1 mm long 30

29(28). Awns 15-20 mm long; spikelets 7-10 mm long .

P. viscidula

Awns 20-25 mm long; spikelets 12-15 mm long

P. velutina

30(28). Lateral awns included in glumes P. tysonii

Lateral awns exserted from glumes 31

31(30). Plants to 300 mm tall; sheath bases brown;

inflorescence nodes glabrous

P. calcicola var. calcicola

Plants to 600 mm tall, sheath bases white;

inflorescence nodes villous P. exserta

32(19). Leaves cauline, all leaves along the stem the same

size 33

Leaves basal or radical 35

33(32). Inflorescence nodes glabrous; plants rhizomatous;

from coastal sands P. scandens

Inflorescence nodes villous; plants without

rhizomes; from mountains 34

34(33). Sheath mouth and lemmas glabrous; anthers 1.2

mm long P. caulescens

Sheath mouth and lemmas villous; anthers 4.0-4. 5

mm long P. acinosa

35(32). Awns spreading, not geniculate P. capensis

Awns erect, geniculate 36

36(35). Glumes usually with cushion-based hairs; bases
weak; culms often geniculate; glumes purplish

P. rosea subsp. purpurascens

Glumes never with cushion-based hairs; plants not
with the above combination of characters . . 37

37(36). Glumes 5-9 mm long P. elegans

Glumes 10-20 mm long 38

38(37). Old leaves expanded, curling; young leaves often

pungent P. pungens

Old leaves rolled; young leaves never pungent . 39
39(38). Awns 20-25 mm long; plants with strong creeping

rhizomes P. aristidoides

Awns 10-17 mm long; plants caespitose 40

40(39). Lateral awns 4-8 mm long; anthers 2. 7-3.0 mm

long; basal leaves brown and rolled

P. pseudopallescens

Lateral awns 3 mm long; anthers 3. 5-4.0 mm long;
basal leaves flat P. pallescens

Pentaschistis acinosa Stapf

Perennial; loosely tufted (or
even cushion- forming); 150-300
mm tall. Leaf blades to 40 mm
long; to 4 mm wide. Spikelets
9-10 mm long. Plants without
glands; leaves spreading on erect
stems; lemmas awned.

Flowering (anthesis) October
to January. Restricted to sand-
stone rock ledges. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (495), Chippindall 1955
(261), Linder & Ellis 1990 (99). Illustration: Chippindall
1955 (fig. 230). Voucher: Esterhuysen 35125 (BOL).
PRECIS code 9902050-00100.

Pentaschistis airoides (Nees) Stapf subsp. airoides

Fig. 160.

( =P . patula (Nees) Stapf var.
glabrata Stapf) 3.

Annual; 60-350 mm tall. Leaf
blades to 30 mm long; to 2 mm
wide. Spikelets 2. 5-3. 5 mm long.

Plants with stalked glands; lem-
mas awned, 1 .5-1.8 mm long; an-
thers less than 0.5 mm long.

Flowering (anthesis) August to December. On ‘richer’
soils, and absent from sandstone derived soils. Common.
Biome: Fynbos, Nama-Karoo, and Succulent Karoo.
Endemic.

Description: Stapf 1898-1900 (510, 511), Chippindall
1 955 (242, 269), Linder & Ellis 1990 (48). Voucher: Linder
4289 (BOL). PRECIS code 9902050-00200.

Pentaschistis airoides (Nees) Stapf subsp. jugorum

(Stapf) Linder

(=P. jugorum Stapf) 3.

Biennial; tufted (nearly cush-
ion-forming); 60-350 mm tall.

Leaf blades 40-60 mm long; 1-3
mm wide. Spikelets 3. 5-5.0 mm
long. Plants with stalked glands;
lemmas awned; anthers 0. 5-1.0
mm long.

Flowering (anthesis) February. On shallow soils and dis-
turbed areas in alpine and high montane areas. Common.
Biome: Afromontane. Endemic.

Description: Stapf 1898-1900 (504), Chippindall 1955
(264), Linder & Ellis 1990 (48). Voucher: Linder 4840
(BOL). PRECIS code 9902050-00250.

Pentaschistis alticola Linder

Biennial; cushion-forming,
with weak bases; 100-300 mm
tall. Leaf blades 30-80 mm long;

0.2-0. 5 mm wide. Spikelets 4-6
mm long. Plants without glands;
lemmas with an awn 7-10 mm
long.

Flowering (anthesis) Novem-
ber to January. Flowering after
fire, on rocky upper slopes of sandstone mountains. Locally
common (on mountain slopes). Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (79). Illustration:
Linder & Ellis 1990 (fig. 10). Voucher: Esterhuysen 28359
(BOL). PRECIS code 9902050-00260.

257

Pentaschistis ampla (Nees) McClean

Perennial; tufted (tussocks
weak, from a woody base);

400-700 mm tall. Leaf blades to
300 mm long; 1-6 mm wide.

Spikelets 3. 3^1.6 mm long. Pedi-
cels usually with obscure linear
glands, rarely with obscure sunk-
en glands; lemmas muticous.

Flowering (anthesis) Decem-
ber to March. At low to mid-altitudes on sandstone derived
soils, often found on rock ledges. Common. Biome: Fynbos
and Succulent Karoo. Endemic.

Description: Chippindall 1955 (266), Linder & Ellis
1990 (59). Illustration: Chippindall 1955 (fig. 236), Linder
(fig. 2). Voucher: Esterhuysen 22769 (BOL). PRECIS code
9902050-00300.

Pentaschistis aspera (Thunb.) Stapf

Perennial; cushion-forming;

300-600 mm tall. Leaf blades
40-100 mm long; 3-6 mm wide.

Spikelets 5-7 mm long. Culms
branched; leaf margins with
prominent stalked glands; lateral
lemma awns as long as the
glumes; central lemma awn 8-10
mm long.

Flowering (anthesis) September to December. In light
disturbances on stony slopes, on both granitic and quartzitic
soils. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (500), Chippindall 1955
(262), Linder & Ellis 1990 (32). Illustration: Chippindall
1955 (fig. 23 1 ). Voucher: Bolus 3340 (BOL). PRECIS code
9902050-01100.

Pentaschistis argentea Stapf

( =P . involuta sensu Adamson
and Chippindall) 3.

Perennial; tufted (geophytic
with a swollen villous under-
ground base, often with stolons);

300-800 mm tall. Leaf blades to
250 mm long; to 2 mm wide.

Spikelets 9-12 mm long.

Plants without glands or with glands linear, restricted to the
pedicels; lemmas awned.

Flowering (anthesis) October and November. On dry
mountain slopes and foothills in sandstone derived soils.
Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (487), Chippindall 1955
(258), Linder & Ellis 1990 (68). Voucher: Linder 4351
(BOL). PRECIS code 9902050-00650.

Pentaschistis aurea (Steud.) McClean subsp. aurea

Perennial; tufted (bases com-
pact); 300^-50 mm tall. Leaf
blades to 300 mm long; 1-5 mm
wide. Spikelets 3. 5-5.0 mm long.

Pedicels with obscure linear
glands; lemmas muticous.

Flowering (anthesis) January
to March. Usually in marshy
areas on sandstone derived soils,
usually at lower altitudes. Common. Biome: Fynbos.
Endemic.

Description: Chippindall 1955 (267), Linder & Ellis
1990 (76). Illustration: Chippindall 1955 (238). Voucher:
Esterhuysen 33107 (BOL). PRECIS code 9902050-01200.

Pentaschistis aurea (Steud.) McClean subsp.
pilosogluma (McClean) Linder

Pentaschistis aristidoides (Thunb.) Stapf

Perennial; rhizomatous (rhi-
zomes stout, villous; shoots
single or several in groups of
3-5); 500-1000 mm tall. Leaf
blades to 300 mm long; 5-10 mm
wide. Spikelets 12-20 mm long.

Leaves mostly borne directly on
the rhizomes; pedicels with ob-
scure linear glands; lemmas
awned.

Flowering (anthesis) September to November. On rocky
sandstone slopes, occasionly on sandy flats. Common.
Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (485), Chippindall 1955
(257), Linder & Ellis 1990 (65). Illustration: Chippindall
1955 (fig. 225.6). Voucher: Esterhuysen 32766 (BOL).
PRECIS code 9902050-00900.

(=P. pilosogluma
McClean) 3.

Perennial; tufted (forming
large floppy tussocks); 600-700
mm tall. Leaf blades to 350 mm
long; to 5 mm wide. Spikelets 6-7
mm long. Plants sometimes with
obscure linear glands; lemmas
muticous.

Flowering (anthesis) December to February. Along
streams and seepages on cave sandstone and basaltic soils.
Infrequent. Biome: Afromontane. Endemic.

Description: Chippindall 1955 (266), Linder & Ellis
1990 (76). Illustration: Chippindall 1955 (fig. 235).
Voucher: Linder 4858 (BOL). PRECIS code

9902050-01250.

Pentaschistis barbata (Nees) Linder subsp. barbata

Pentaschistis aristifolia Schweick.

Annual; about 250 mm tall.

Leaf blades to 60 mm long; to 4
mm wide. Spikelets 2. 5-3.0 mm
long. Plants without glands; leaf
apices with long slender aristae;
lemmas awned.

Flowering (anthesis) Septem-
ber and October. On heavier soils
associated with the Karoo sed-
iments. Common. Biome: Nama-Karoo. Endemic.

Description: Chippindall 1955 (270), Linder & Ellis
1990 (49). Voucher: Linder 4279 (BOL). PRECIS code
9902050-01000.

(=P. angulata sensu
Adamson, non Nees) 3; (=P.
leucopogon Stapf) 3.

Perennial; weakly tufted
(almost forming cushions); 300-
600 mm tall. Leaf blades to 200
mm long; to 12 mm wide. Spike-
lets 5-6 mm long. Leaf mar-
gins and pedicels usually with stalked glands; lemmas awn-
ed, the lateral awns exserted from the glumes.

Flowering (anthesis) September to November. Coastal
sands in slightly disturbed areas. Common (west coast sand
flats). Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (500), Chippindall 1955
(262), Linder & Ellis 1990 (31). Voucher: Esterhuysen
31336 (BOL). PRECIS code 9902050-01320.

258

Pentaschistis barbata (Nees) Linder subsp. orientalis

Linder

Perennial; tufted; 300-600
mm tall. Leaf blades to 300 mm
long; to 9 mm wide. Spikelets
8-10 mm long. Leaf margins with
stalked glands; lemmas awned, 5
mm long.

Flowering (anthesis) Febru-
ary. On coastal dunes. Conserva-
tion status not known. Infrequent. Biome: Fynbos.
Endemic.

Description: Linder & Ellis 1990 (31). Voucher: Van der
Merwe 2168 (STE). PRECIS code 9902050-01370.

Pentaschistis basutorum Stapf

Perennial; tufted; 500-700
mm tall. Leaf blades to 600 mm
long; to 0.5 mm wide. Spikelets
7-10 mm long. Plants without
glands; leaves tough, villous
inside; lemmas awned.

Flowering (anthesis) Decem-
ber and January. On shallow soil
over sandstone. Locally common
(western slopes of Drakensberg). Biome: Afromontane.
Endemic.

Description: Chippindall 1955 (260), Linder & Ellis
1990 (94). Voucher: Dieterlen 1 162 (BOL). PRECIS code
9902050-01400.

Pentaschistis calcicola Linder var. calcicola

Perennial; tufted (tussocks
very neat); 200-300 mm tall. Leaf
blades 30-100 mm long; 0.3-0. 5
mm wide. Spikelets 5-7 mm long.

Plants without glands; leaves in a
tight basal tussock; lemmas
awned.

Flowering (anthesis) October.

Restricted to limestone pave-
ments. Conservation status not known. Infrequent. Biome:
Fynbos. Endemic. Sometimes confused with P patuliflora
Rendle), a synonym of P. cirrhulosa.

Description: Linder & Ellis 1990 (83). Illustration:
Linder & Ellis 1990 (fig. 12). Voucher: Du Toil 1960
(BOL). PRECIS code 9902050-01520.

Pentaschistis calcicola Linder var. hirsuta Linder

Perennial; tufted; 200-300
mm tall. Leaf blades to 30 mm
long; to 0.5 mm wide (puberu-
lous). Spikelets 6-7 mm long.

Plants without glands; lemmas
awned.

Flowering (anthesis) Septem-
ber. Restricted to limestone
pavements. Conservation status
not known. Infrequent. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (83). Voucher: Linder
4366 (BOL). PRECIS code 9902050-01550.

Pentaschistis capensis (Nees) Stapf

Perennial; tangled; 200-350
mm tall. Leaf blades to 120 mm
long; to 4 mm wide. Spikelets 6-9
mm long. Glands absent; lemmas
with spreading awns, awns not
geniculate.

Flowering (anthesis) Decem-
ber and January. Restricted to
rocky streams where it grows in

water, often over waterfalls. Locally common. Biome: Fyn-
bos. Endemic.

Description: Stapf 1898-1900 (494), Chippindall 1955
(261), Linder & Ellis 1990 (103). Illustration: Chippindall
1955 (fig. 225.7). Voucher: Esterhuysen 33060 (BOL).
PRECIS code 9902050-01600.

Pentaschistis capillaris (Thunb.) McClean

Annual; 80-400 mm tall. Leaf
blades to 50 mm long; to 5 mm
wide. Spikelets 3 mm long. Plants
with stalked glands; lemmas mu-
ticous.

Flowering (anthesis) Septem-
ber and October. In coastal sands.

Infrequent. Biome: Fynbos and
Succulent Karoo. Endemic.

Description: Chippindall 1955 (268), Linder & Ellis
1990 (48). Illustration: Chippindall 1955 (fig. 241).
Voucher: Pillans 7938 (BOL). PRECIS code

9902050-01700.

Pentaschistis caulescens Linder

Perennial; tangled; 150-300
mm tall. Leaf blades 30-40 mm
long; 1-2 mm wide. Spikelets
8-12 mm long. Glands absent;
leaves spreading from aerial
stems; lemmas awned, glabrous.

Flowering (anthesis) Septem-
ber to October. On shale bands on
dry stony slopes. Conservation
status not known. Abundance not known. Biome: Fynbos.
Endemic.

Description: Linder & Ellis 1990 (99). Illustration:
Linder & Ellis 1990 (fig. 17). Voucher: Esterhuysen 26349
(BOL). PRECIS code 9902050-01720.

Pentaschistis chippindalliae Linder

Perennial; tufted; 300-500
mm tall. Leaf blades to 200 mm
long; to 0.5 mm wide. Spikelets
4. 5-7. 5 mm long. Plants without
glands; inflorescence open and
fine; lemmas awned.

Flowering (anthesis) February
to March. Restricted to sour
grassland in highlying ground in
the eastern Transvaal, probably restricted to quartzites. Lo-
cally common. Biome: Afromontane. Endemic.

Description: Linder & Ellis 1990 (92). Illustration:
Linder & Ellis 1990 (fig. 15). Voucher: Linder 471 1 (BOL).
PRECIS code 9902050-01740.

Pentaschistis cirrhulosa (Nees) Linder

( -P . angustifolia (Nees)

Stapf var. cirrhulosa (Nees)

Stapf) 3; ( -P . hurchellii Stapf)

3; (-P. patuliflora Rendle) 3.

Perennial; tufted; 150-300
mm tall. Leaf blades 30-70 mm
long; 0. 5-3.0 mm wide (usually
rolled, purplish). Spikelets 5-9
mm long. Stalked glands present on leaf sheaths, pedicels
and glumes; lemmas awned.

Flowering (anthesis) October. On sandstone gravels at
lower altitudes. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900(501, 503), Linder & Ellis
1990 (42). Voucher: Esterhuysen 17189 (BOL). PRECIS
code 9902050-01760.

259

Pentaschistis colorata (Steud.) Stapf

( =P . colorata (Steud.) Stapf
var. polytricha Stapf) 3.

Perennial; tufted (or cushion-
forming, possibly tangled, habit
of living plants needs study);

300-600 mm tall. Leaf blades
150-300 mm long; 0.3-1. 0 mm
wide (usually curly). Spikelets
8-13 mm long. Plants without glands; inflorescence gener-
ally with fewer than 20 spikelets; lemmas awned.

Flowering (anthesis) September to December. With a
wide ecological range, usually occurs on stony slopes in
sandstone derived soils. Common. Biome: Fynbos.
Endemic.

Description: Stapf 1898-1900 (491), Chippindall 1955
(260), Linder & Ellis 1990 (77). Voucher: Esterhuysen
19017 (BOL). PRECIS code 9902050-01800.

Pentaschistis curvifolia (Schrad.) Stapf

Fig. 161. PI. 147.

Perennial; tufted; 400-500
mm tall. Leaf blades to 300 mm
long; to 4 mm wide (margins
thickened). Spikelets 8-12 mm
long. Plants without glands; infl-
orescence compact, with the pedi-
cels obscured by the ivory-col-
oured spikelets; lemmas awned.

Flowering (anthesis) October
and November. Widespread over wide altitude range, usual-
ly in Fynbos on sandstone derived soils. Common. Biome:
Fynbos. Endemic.

Description: Stapf 1898-1900 (491), Chippindall 1955
(258), Linder & Ellis 1990 (96). Illustration: Chippindall
1955 (fig. 226). Voucher: Linder 4793 (BOL). PRECIS
code 9902050-02000.

Pentaschistis densifolia (Nees) Stapf

(-P. densifolia (Nees) Stapf
var. intricata Stapf) 3.

Softly herbaceous perennial;
cushion-forming; 90-250 mm
tall. Leaf blades to 40 mm long;
to 1.5 mm wide (old blades dry-
ing pink). Spikelets 3. 5-4. 5 mm
long. Leaf margins with stalked
glands; lemmas awned, awn 3-6 mm long.

Flowering (anthesis) December to January. On ledges
and in crevices at mid-altitudes on mountains, often grow-
ing in moss-beds. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (506), Chippindall 1955
(263), Linder & Ellis 1990 (41). Voucher: Esterhuysen
22595 (BOL). PRECIS code 9902050-02100.

Pentaschistis ecklonii (Nees) McClean

( =P . bachmannii
McClean) 3.

Perennial; tufted; 200-300
mm tall. Leaf blades to 60 mm
long; to 2 mm wide. Spikelets 3-4
mm long. Plants with stalked
glands; inflorescence spikelike;
lemmas muticous.

Flowering (anthesis) January to March. Local on ‘richer’
soils derived from shales or sand in the lowlands. Infre-
quent. Biome: Fynbos. Endemic.

Description: Chippindall 1955 (267), Linder & Ellis
1990 (52). Illustration: Chippindall 1955 (fig. 239).
Voucher: Esterhuysen 24035 (BOL). PRECIS code
9902050-02250.

Fig. 161. Pentaschistis curvifolia

260

Pentaschistis elegans (Nees) Stapf

Pentaschistis giandulosa (Schrad.) Linder

Perennial; tufted; 200-300
mm tall. Leaf blades 20-30 mm
long; to 1 mm wide. Spikelets 7-9
mm long. Plants without glands;
lemmas sparsely villous at the
apex, with the central awn 15 mm
long.

Flowering (anthesis) Septem-
ber. In sand on coastal flats. Rare.

Biome; Fynbos. Endemic.

Description: Stapf 1898-1900 (496), Chippindall 1955
(261), Linder & Ellis 1990 (90). Voucher: Henderson 1820
(BOL). PRECIS code 9902050-02300.

Pentaschistis eriostoma (Nees) Stapf

(=P. angustifolia (Nees)

Stapf var. micrathera (Nees)

Stapf) 3.

Perennial; tufted; 100-350
mm tall. Leaf blades 80-300 mm
long; 1-3 mm wide. Spikelets
4. 0-5. 5 mm long. Leaf blades
with sunken glands; lemmas
awned.

Flowering (anthesis) October. ‘Richer' granite-derived
soils. Common. Biome: Fynbos and Savanna. Endemic.

Description: Stapf 1898-1900 (503), Chippindall 1955
(264), Linder & Ellis 1990 (60). Voucher: Linder 4807
(BOL). PRECIS code 9902050-02850.

(=P. juncifolia Stapf) 3.

Perennial; tufted; 300-900
mm tall. Leaf blades to 400 mm
long; to 1.5 mm wide. Spikelets
8-12 mm long. Plants without
glands; basal leaf sheaths gener-
ally with a dense woolly cover-
ing, or sometimes only the sheath
apex with wool; lemmas awned.

Flowering (anthesis) September to November. Many
habitats. Common. Biome: Fynbos and Succulent Karoo.

Description: Stapf 1898-1900 (489), Chippindall 1955
(260), Linder & Ellis 1990 (106). Illustration: Chippindall
1955 (fig. 227). Voucher: Esterhuysen 27416 (BOL).
PRECIS code 9902050-02400.

Pentaschistis heptamera (Nees) Stapf

Perennial; single or several
shoots; 200-300 mm tall. Leaf
blades to 80 mm long; to 0.5 mm
wide. Spikelets 5-6 mm long.

Plants without glands; lemmas
awned, with 4-6 lateral awns.

Flowering (anthesis) Novem-
ber to December. Restricted to
coastal sands. Conservation status
not known. Infrequent. Biome: Savanna. Endemic.

Description: Stapf 1898-1900 (504), Chippindall 1955
(262), Linder & Ellis 1990 (106). Illustration: Chippindall
1955 (fig. 225.8). Voucher: Fourcade 1810 (BOL). PRECIS
code 9902050-02900.

Pentaschistis exserta Linder

Perennial; tufted (plant base
with horizontal stolons); about
600 mm tall. Leaf blades to 300
mm long; to 0.5 mm wide. Spike-
lets 7. 5-8. 5 mm long. Glands ab-
sent; lemmas awned, lateral awns
exserted from the glumes.

Flowering (anthesis) January.

Local in seeps and along streams in the montane belt. Lo-
cally common. Biome: Afromontane. Endemic. Mostly
under P. tysonii aff.

Description: Linder & Ellis 1990 (92). Voucher: Ellis
5723 (PRE). PRECIS code 9902050-02650.

Pentaschistis galpinii (Stapf) McClean

PI. 148.

Perennial; cushion-forming,
the cushions low and rounded, or
sometimes the plant with stolons
and mat-forming; 150-300 mm
tall. Leaf blades 60-180 mm
long; 2-5 mm wide (basally
aggregated). Spikelets 4-6 mm
long. Plants with stalked glands;
inflorescence usually contracted,
with the basal pedicel segments as long as the spikelets;
lemmas awned or muticous.

Flowering (anthesis) January. Alpine grassland, usually
in bare patches on wet basalt, alt. 2-3000 m. Common.
Biome: Afromontane. Endemic.

Description; Chippindall 1955 (234), Linder & Ellis
1990 (51). Voucher: Linder 4844 (BOL). PRECIS code
9902050-02800.

Pentaschistis holciformis (Nees) Linder

Perennial; tufted; 400-600
mm tall. Leaf blades 150-200
mm long; to 0.5 mm wide. Spike-
lets 6-7 mm long. Plants without
glands; lemmas awned. lateral
awns 0.3 mm long.

Flowering (anthesis) March.

On sandstone derived soils in the
mountains, usually in black soils,
often on firebreaks. Locally common (after fires). Biome:
Fynbos. Endemic.

Description: Stapf 1898-1900 (536), Linder & Ellis
1990 (91). Voucher: Esterhuysen 33498 (BOL). PRECIS
code 9902050-03050.

Pentaschistis lima (Nees) Stapf

Perennial; tufted; about 450
mm tall. Leaf blades to 300 mm
long; to 0.5 mm wide (tightly roll-
ed). Spikelets 6-7 mm long. Pedi-
cels with stalked glands; lemmas
awned.

Flowering (anthesis) Novem-
ber to December. On granitic
soils. Conservation status not
known. Abundance not known. Biome: Nama-Karoo.
Endemic.

Description: Stapf 1898-1900 (498), Linder & Ellis
1990 (44). Voucher: Adamson 1475 (BOL). PRECIS code
9902050-03250.

261

Pentaschistis longipes Stapf

Perennial; tufted; 250-700
mm tall. Leaf blades to 150 mm
long; to 4 mm wide. Spikelets

4. 5- 5.0 mm long. Margins and
glumes with stalked glands; lem-
mas awned, central awn 7 mm
long.

Coastal sands. Conservation
status not known. Abundance not
known. Endemic.

Description: Stapf 1898-1900 (509), Chippindall 1955
(264), Linder & Ellis 1990 (35). Voucher: Liebenberg 3968
(PRE). PRECIS code 9902050-03400.

Pentaschistis malouinensis (Steud.) Clayton

Fig. 162.

( =Achneria capensis (Steud.)

Dur. & Schinz) 2; (=P. steudelii
McClean) 3.

Perennial; tufted; 150-300
mm tall. Leaf blades to 150 mm
long; to 0.5 mm wide. Spikelets

3. 5- 4. 5 mm long. Plants without
glands; glume apices brown.

Fig. 162. Pentaschistis malouinensis

shortly puberulous and rounded; lemmas muticous.

Flowering (anthesis) November to January. Widespread
in a range of habitats, from sparse vegetation to rock ledges,
over a wide altitudinal range, often found in disturbed sites.
Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (459), Chippindall 1955
(267), Linder & Ellis 1990 (87). Illustration: Chippindall
1955 (fig. 240). Voucher: Esterhuysen 19399 (BOL).
PRECIS code 9902050-03500.

Pentaschistis microphylia (Nees) McClean

Perennial; cushion-forming or
sometimes mat-forming; about
300 mm tall. Leaf blades to 50
mm long; to 3 mm wide. Spike-
lets 3 mm long. Plants with stalk-
ed glands, especially on the pedi-
cels; leaves broad and stiff, in
basal rosettes; lemmas muticous.

Flowering (anthesis) Decem-
ber. Arid montane grassland in the Stormberg, in shallow
soils over bedrock at ca. 2000 m. Locally common. Biome:
Afromontane. Endemic.

Description: Chippindall 1955 (266), Linder & Ellis
1990 (51). Illustration: Chippindall 1955 (fig. 237).
Voucher: Flanagan 1668 (BOL). PRECIS code

9902050-03600.

Pentaschistis montana Linder

Perennial; tufted (mat-form-
ing); 150-200 mm tall. Leaf blad-
es to 40 mm long; to 0.5 mm
wide. Spikelets 4. 5-5.0 mm long.

Plants without glands; leaves
basal; lemmas awned, central awn
5-6 mm long.

Flowering (anthesis) Novem-
ber. Stony, arid upper slopes in
sandstone mountains. Common. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (83). Illustration:
Linder & Ellis 1990 (fig. 13). Voucher: Esterhuysen 35723
(BOL). PRECIS code 9902050-03650.

Pentaschistis natalensis Stapf

Perennial; loosely tufted
(forming diffuse tussocks); 400-
800 mm tall. Leaf blades to 300
mm long; 1-5 mm wide. Spike-
lets 6-7 mm long. Glume keels
with small rounded glands; lem-
mas awned.

Flowering (anthesis) Novem-
ber to February. In sour grassland
or near forest margins in the montane belt. Infrequent.
Biome: Afromontane. To southern Tanzania, and

Madagascar.

Description: Stapf 1898-1900 (493), Chippindall 1955
(265), Linder & Ellis 1990 (56). Illustration: Chippindall
1955 (fig. 225.3). Voucher: Du Toit 1174 (BOL). PRECIS
code 9902050-03700.

Pentaschistis oreodoxa Schweick.

Perennial; tufted (tends to-
wards cushion formation); 200-
500 mm tall. Leaf blades to 300
mm long; to 4 mm wide. Spike-
lets 4-6 mm long. Plants with
stalked glands; lemmas awned,
lobes acute.

Flowering (anthesis) January.

In sour grassland over a wide
altitudinal range. Common. Biome: Afromontane.

Endemic.

Description: Chippindall 1955 (265), Linder & Ellis
1990 (57). Voucher: Killick 1300 (BOL). PRECIS code
9902050-03900.

262

Pentaschistis pallescens (Schrad.) Stapf

(=P. sylvatica Adamson) 3.

Perennial; tufted; 600-1200
mm tall. Leaf blades to 600 mm
long; to 8 mm wide. Spikelets
10-12 mm long. Plants without
glands or with linear glands
restricted to pedicels; leaf blades
much darker above than below;
lemmas awned.

Flowering (anthesis) November and December. On the
lower slopes of sandstone mountains, usually found after
fire. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (487), Chippindall 1955
(258), Linder & Ellis 1990 (74). Illustration: Chippindall
1955 (fig. 225.1). Voucher: Esterhuysen 17614 (BOL).
PRECIS code 9902050-04000.

Pentaschistis pallida (Thunb.) Linder

Description: Stapf 1898-1900 (510), Chippindall 1955
(268, 270), Linder & Ellis 1990 (45). Illustration: Linder
& Ellis 1990 (fig. 1). Voucher: Crook 1018 (BOL). PRECIS
code 9902050-04300.

Pentaschistis praecox Linder

Perennial; tufted; 300-600
mm tall. Leaf blades to 300 mm
long; to 0.5 mm wide. Spikelets
8-11 mm long. Plants without
glands; glumes acuminate, golden
brown; lemmas awned.

Flowering (anthesis) Septem-
ber. In sour grassland in the
montane belt. Infrequent. Biome:

Afromontane. Endemic.

Description: Linder & Ellis 1 990 (95). Voucher: Gordon
Gray 8000 (NU). PRECIS code 9902050-04350.

( -P . angustifolia (Nees)

Stapf) 3; (=P. angustifolia var.
albescens StapD 3; ( -P .
filiformis (Nees) StapD 3; ( =P .
heterochaeta StapD 3; ( =P .
imperfecta StapD 3; (=P.
thunbergii sensu StapD 3.

Perennial; tufted; 150-400
mm tall. Leaf blades 120-200 mm long; 1-5 mm wide.
Spikelets 3-5 mm long. Plants usually with stalked glands;
inflorescence with numerous spikelets; lemmas awned, lat-
eral awns exserted from the glumes.

Flowering (anthesis) October. Widespread in places
with slight to heavy distrubance. Common. Biome: Fynbos
and Succulent Karoo. Endemic. This species has seven
intergrading morphological forms.

Description: Stapf 1898-1900 (502, 503, 505, 507, 508),
Chippindall 1955 (264), Linder & Ellis 1990 (36). Voucher:
Esterhuysen 19270 (BOL). PRECIS code 9902050-04150.

Pentaschistis pseudopallescens Linder

Weakly perennial; tufted;

400-800 mm tall. Leaf blades to
300 mm long; to 6 mm wide.

Spikelets 10-12 mm long. Inner
surface of the leaves villous, old
leaves curling; leaf margins and
pedicels with obscure linear
glands; lemmas awned. lateral
awns as long as the glumes.

Flowering (anthesis) November and December. Along
seeps and streams in sand at mid-altitude in the Cape sand-
stone mountains, after fire. Infrequent. Biome: Fynbos.
Endemic.

Description: Linder & Ellis 1990 (72). Illustration:
Linder & Ellis 1990 (fig. 9). Voucher: Linder 4483 (BOL).
PRECIS code 9902050-04420.

Pentaschistis pungens Linder

Pentaschistis papillosa (Steud.) Linder

(-P. subulifolia StapD 3;

(=P. zeyheri StapD 3.

Perennial; tangled; 100-400
mm tall. Leaf blades 35-100 mm
long; 3-6 mm wide. Spikelets 5-7
mm long. Plants with spreading
culms; pedicels with stalked
glands; lemmas awned, lateral
awns usually exserted from the glumes.

Flowering (anthesis) October and November. At low al-
titudes on sandstones. Common. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (497, 499), Chippindall
1955 (262), Linder & Ellis 1990 (32). Voucher: Leighton
201 1 (BOL). PRECIS code 9902050-04250.

Pentaschistis patula (Nees) Stapf

(-P. euadenia StapD 3; ( =P .
patula (Nees) Stapf var. acuta
StapD 3.

Annual; 150-300 mm tall.

Leaf blades to 50 mm long; to 3
mm wide. Spikelets 3. 5-5.0 mm
long. Plants with stalked glands;
lemmas awned, the lateral awns
included in the glumes.

Flowering (anthesis) September to October. Local in
sandy soils at the arid margins of the Fynbos and in Nama-
qualand. Common. Biome: Fynbos and Succulent Karoo.
Endemic.

Biennial; tufted; 200-500 mm
tall. Leaf blades to 120 mm long;
to 4 mm wide. Spikelets 11-15
mm long. Glands absent; old
leaves flat and recurved; lemmas
awned, central awn 17-20 mm
long.

Flowering (anthesis) Septem-
ber. At higher altitudes, usually in
damp sand, after fire. Common. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (97). Illustration:
Linder & Ellis 1990 (fig. 16). Voucher: Esterhuysen 13030
(BOL). PRECIS code 9902050-04460.

Pentaschistis pusilla (Nees) Linder

PI. 149.

(-Poagrostis pusilla (Nees)

StapD 3.

Perennial (soft, pale green
plants); cushion-forming; about
120 mm tall. Leaf blades to 25
mm long; to 1.5 mm wide. Spike-
lets 2. 5-3.0 mm long. Glands ab-
sent; spikelets usually single-
flowered; lemmas muticous.

Local in damp cool habitats in rock crevices, on ledges,
and especially along streams and waterfalls. Common.
Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (760), Chippindall 1955
(272), Linder 1990 (89). Illustration: Chippindall 1955 (fig.
244). Voucher: Esterhuysen 26945 (BOL). PRECIS code
9902050-04520.

263

Pentaschistis pyrophila Linder

Perennial; tufted; 200-600
mm tall. Leaf blades 60-200 mm
long; 0.5-1.5 mm wide (apices
sometimes pungent). Spikelets
10-12 mm long. Plants without
glands, usually with burnt-off leaf
sheaths; lemmas awned.

Flowering (anthesis) Novem-
ber to January. At higher altitudes
on stony slopes on sandstones. Common. Biome: Fynbos.
Endemic.

Description: Linder & Ellis 1990 (81). Illustration;
Linder & Ellis 1990 (fig. 1 1). Voucher: Esterhuysen 28598
(BOL). PRECIS code 9902050-04550.

Pentaschistis reflexa Linder

Biennial; tufted or tangled;

100-350 mm tall. Leaf blades to
30 mm long; to 1.5 mm wide.

Spikelets 3-4 mm long. Plants
with stalked glands; spikelets
reflexed at anthesis; lemmas mu-
ticous.

Flowering (anthesis) October
to December. Local at lower alti-
tudes in arid Fynbos. Conservation status not known. Infre-
quent. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (53). Illustration:
Linder & Ellis 1990 (fig. 7). Voucher: Esterhuysen 17931
(BOL). PRECIS code 9902050-04570.

Pentaschistis rigidissima Pilg. ex Linder

Perennial; cushion-forming,
sometimes forming ‘vegetable
hedgehogs’; 100-300 mm tall.

Leaf blades 40-200 mm long;

0. 5-1.0 mm wide (apices some-
times pungent). Spikelets 6-8
mm long. Plants without glands;
panicle slender, almost spikelike;
spikelets greenish; lemmas
awned.

Flowering (anthesis) September to December. Rock
crevices at mid and upper altitudes in the mountains, and
in arid habitats. Common. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (85). Illustration:
Linder & Ellis 1990 (fig. 14). Voucher: Esterhuyen 12437
(BOL). PRECIS code 9902050-04600.

Pentaschistis rosea Linder subsp. purpurascens Linder

Biennial; cushion-forming;

150 — 400 mm tall. Leaf blades
20-100 mm long; 0. 5-3.0 mm
wide. Spikelets 8-12 mm long.

Leaves cauline; leaf margins and
pedicels with obscure linear
glands; glumes usually purplish
with tufts of cushion-hairs; lem-
mas awned.

Flowering (anthesis) October to December. At higher al-
titudes on sandy flats and stony slopes after fire. Common.
Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990. Voucher: Linder 4403
(BOL). PRECIS code 9902050-04650.

Pentaschistis rosea Linder subsp. rosea

Biennial; tufted; 150-400 mm
tall. Leaf blades 50-100 mm
long; to 1 mm wide. Spikelets
11-12 mm long. Leaves linear,
basal; leaf margins and pedicels
with obscure linear glands;
glumes often with tufts of cushion-
hairs; lemmas awned.

Flowering (anthesis) October.

After fire on deep sandy soils. Locally dominant. Biome:
Fynbos. Endemic.

Description: Linder & Ellis 1990 (71). Voucher:
Esterhuysen 21885 (BOL). PRECIS code 9902050-04670.

Pentaschistis rupestris (Nees) Stapf

Perennial; tufted; 600-1000
mm tall. Leaf blades to 400 mm
long; to 6 mm wide. Spikelets 7-8
mm long. Plants generally with
stalked glands on the leaves and
especially on the pedicels; inflor-
escences usually open, hemis-
pherical; lemmas awned.

Flowering (anthesis) October.

Cedarberg, with a wide habitat range, but restricted to sand-
stone derived soils. Locally common. Biome: Fynbos.
Endemic. Previously confused with P. veneta Linder.

Description: Linder & Ellis 1990 (34). Voucher: Linder
4468 (BOL). PRECIS code 9902050-04700.

Pentaschistis scandens Linder

Perennial; tangled, shoots
spreading through vegetation;

300-500 mm tall. Leaf blades to
15 mm long; to 0.5 mm wide.

Spikelets 10-1 1 mm long. Plants
without glands; leaves flat and
spreading from long, scandent,
aerial stems; lemmas awned.

Flowering (anthesis) August.

In sandy soils on the Bredasdorp plains. Conservation status
not known. Infrequent. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (101). Illustration:
Linder& Ellis 1990 (fig. 18). Voucher: Linder 4766 (BOL).
PRECIS code 9902050-04750.

Pentaschistis setifolia (Thunb.) McClean

Perennial; tufted; 150^400
mm tall. Leaf blades to 300 mm
long; to 0.8 mm wide (curly).

Spikelets 3. 5-5.0 mm long.

Plants with stalked or sunken
glands; lemmas muticous, often
dark-coloured.

Flowering (anthesis) Decem-
ber and January. Sour grasslands
over a wide altitudinal range. Dominant. Biome: Afromont-
ane. Endemic. Delimitation from P oreodoxa , P.
glandulosa and P. ampla is difficult.

Description: Chippindall 1955 (266), Linder & Ellis
1990 (58). Illustration: Chippindall 1955 (fig. 233).
Voucher: Linder 4860 (BOL). PRECIS code

9902050-04800.

264

Pentaschistis tomentella Stapf

( =P . brachyanthera Stapf) 3.

Perennial; tufted (or cushion-
forming); 100-300 mm tall. Leaf
blades to 50 mm long; to 3 mm
wide. Spikelets 4—5 mm long.

Leaf sheaths with distinctive
rows of stalked glands; inflores-
cence compact; lemmas awned,
the lateral awns included in the glumes.

Flowering (anthesis) September. Widespread in the
higher and cooler parts of Namaqualand. Common. Biome:
Nama-Karoo. Endemic.

Description: Stapf 1898-1900 (502, 507), Chippindall
1955 (268), Linder & Ellis 1990 (43). Illustration: Chippin-
dall 1955 (fig. 22.5). Voucher: Taylor 1166 (BOL).
PRECIS code 9902050-05600.

Pentaschistis tortuosa (Trin.) Stapf

(=P. nutans (Nees) Stapf) 3.

Perennial; tufted (tussock
tight); 600-1000 mm tall. Leaf
blades to 500 mm long; to 4 mm
wide (linear). Spikelets 7-1 1 mm
long. Plants without glands; infl-
orescence slender, often some-
what tangled and with the apex
drooping; lemmas awned.

Flowering (anthesis) October to December. In damp
places on mountains and foothills, usually associated with
sandstone soils. Common. Biome: mature Fynbos.

Endemic.

Description: Stapf 1898-1900 (488), Chippindall 1955
(258, 259), Linder & Ellis 1 990 (78). Voucher: Esterhuysen
34035 (BOL). PRECIS code 9902050-05700.

Pentaschistis triseta (Thunb.) Stapf

Annual; 200-600 mm tall.

Leaf blades to 80 mm long; to 4
mm wide. Spikelets 15-18 mm
long. Leaf margins and pedicels
with obscure linear glands; lem-
mas awned.

Flowering (anthesis) Septem-
ber and October. On sandy soils
below 600 m. Locally common
(after fire). Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (495), Chippindall 1955
(261), Linder & Ellis 1990 (69). Illustration: Chippindall
1955 (fig. 229). Voucher: Linder 4297 (BOL). PRECIS
code 9902050-05800.

Pentaschistis tysonii Stapf

Fig. 163.

(-P. fibrosa) Stapf 3.

Perennial; tufted (plants often
with a basal sheath of fibres);

300-500 mm tall. Leaf blades to
300 mm long; 0. 5-2.0 mm wide
(rolled). Spikelets 7-9 mm long.

Plants without glands; leaves
basal; lemmas awned, lateral
awns included in the glumes.

Flowering (anthesis) November. Sour grassland on
mountain slopes. Common. Biome: Afromontane. Endemic.

Description: Stapf 1898-1900 (493), Chippindall 1955
(261), Linder & Ellis 1990 (90). Voucher: Linder 4833
(BOL). PRECIS code 9902050-05900.

265

Pentaschistis velutina Linder

Perennial; tufted (base
somewhat swollen, densely vil-
lous and without a rhizome);

300-600 mm tall. Leaf blades to
180 mm long; to 1 mm wide.

Spikelets 12-15 mm long. Pedi-
cels with obscure linear glands;
lemmas awned.

Flowering (anthesis) October
and November. On gravelly plateaus and shale bands in
mountains. Infrequent. Biome: Fynbos. Endemic.

Description: Linder & Ellis 1990 (66). Illustration:
Linder & Ellis 1990 (fig. 8). Voucher: Linder 4791 (BOL).
PRECIS code 9902050-05930.

Pentaschistis veneta Linder

Perennial; tufted; 200-400
mm tall. Leaf blades to 100 mm
long; to 4 mm wide. Spikelets
5.0-6. 5 mm long. Leaf margins
with stalked glands; lemmas awn-
ed, lateral awns as long as the
glumes.

Flowering (anthesis) Decem-
ber to January. In black sand in
seeps, at mid to upper altitudes in the Cape sandstone
mountains. Common. Biome: Fynbos. Endemic. In many
herbaria under/5, rupestris (Nees) Stapf.

Description: Stapf 1898-1900 (498, as P. rupestris),
Chippindall 1955 (263, as/5, rupestris), Linder & Ellis 1990
(29). Voucher: Esterhuysen 15125 (BOL). PRECIS code
9902050-05960.

Pentaschistis viscidula (Nees) Stapf

Perennial; geophytic with
swollen, villous bases, stolons ab-
sent, shoots single or several;

200-500 mm tall. Leaf blades to
100 mm long; to 0.8 mm wide.

Spikelets 7-10 mm long. Pedicels
sometimes with obscure linear
glands; lemmas awned, lateral
awns 3 mm long.

Flowering (anthesis) October and November. After fire
on sandstone derived soils in the Cape mountains. Com-
mon. Biome: Fynbos. Endemic.

Description: Stapf 1898-1900 (486), Chippindall 1955
(258), Linder & Ellis 1990 (68). Voucher: Esterhuysen
29946 (BOL). PRECIS code 9902050-06000.

Periballia Trin.

Molineria Pari., Molineriella Rouy. Sometimes included
in Deschampsia P. Beauv.

Annual', caespitose (or culms solitary). Culms 30-250
mm high; herbaceous. Leaf blades linear; flat, or rolled
(convolute). Ligule an unfringed membrane .

Inflorescence paniculate ; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets 1.75-2 mm long; compressed laterally; disar-
ticulating above the glumes; with an elongated rachilla
internode between the florets. Rachilla terminated by a
female-fertile floret. Glumes two; more or less equal;
markedly shorter than the spikelets', awnless; similar
(membranous). All florets female-fertile; proximal incom-
plete florets absent.

Female-fertile florets 2. Lemmas less firm than the
glumes (hyaline), or similar in texture to the glumes;
incised (irregularly toothed); 3-7 nerved; awnless. Palea

present. Lodicules 2; membranous; glabrous. Stamens 3.
Ovary glabrous. Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 4 and 7. Pooideae; Poodae; Aveneae. 3 species.
Mediterranean. Xerophytic; in open habitats (dry sandy
places). Cape Province. 1 naturalized species.

References. I. Chippindall. 1955. Gr. & Past. 2. Tutin.
1980. FI. Europ. (N.B. - This species treated under
Molineriella in FI. Europ., without synonymy).

Species treatment by T.M. Sokutu.

Periballia minuta (L.) Asch. & Graebn.

Annual; loosely tufted; 30-
140 mm tall. Leaf blades 7-30
mm long; to 1.5 mm wide. Spike-
lets 1-2 mm long. Glumes shorter
than lemmas; lemmas obtuse or
truncate, awnless; palea not keel-
ed, rounded at the back and
shorter than the lemmas.

Flowering August to Septem-
ber. Moist shallow soil. Infrequent to locally common. Nat-
uralized from the Mediterranean. Biome: Fynbos. Southern
Europe. Weed. May be confused with Air a cupaniana , but
distinguished by an elongated internode between the florets,
a fairly open panicle, glumes that are shorter than the
lemmas, and the absence of awns. Collected once in
Simonstown in 1943.

Description: Chippindall 1955 (86). Voucher: Salter
8766. PRECIS code 9901870-00100.

Perotis Aiton

Xystidium Trin.

Annual, or perennial (rarely); caespitose. Culms
120-1000 mm high; herbaceous. Ligule an unfringed
membrane to a fringed membrane.

Inflorescence a single spike, or a single raceme (a
narrow ‘ bottlebrush’ , bearded by the long glume awns)',
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary (often reflexing when mature); subses-
sile to pedicellate', 1.2-5. 5 mm long; compressed laterally;
falling with the glumes. Glumes two; more or less equal;
long relative to the adjacent lemmas (considerably
exceeding them); awned', similar (narrow, membranous to
cartilaginous, tipped by long capillary awns). All florets
female-fertile; proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline)', 1 nerved; entire; awnless. Palea present;
conspicuous but relatively short (but almost equalling the
lemma). Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small to medium sized (almost as long as
the glumes); hilum short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 10
species. Africa, India, Ceylon, eastern Asia, Australia.
Mesophytic, or xerophytic; in open habitats (savanna and
grassland, often ruderal); glycophytic. Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal and Cape
Province. 3 indigenous species.

References. 1. Clayton. 1971. Kew Bull. 25: 250. 2.
Clayton et al. 1974. FTEA.

Species treatment by G.E. Gibbs Russell.

266

1(0). Base of spikelet rounded or flat, not elongated into a
stipe; awns usually purple; widespread distribution

P. patens

Base of spikelet elongated into a narrow tapering
stipe; awns usually green; northern Namibia and

Botswana 2

2(1). Spikelets 3. 5-5. 5 mm long (including stipe); awns

13-25 mm long P. vaginata

Spikelets 2. 5-3.0 mm long (including stipe); awns
20^4-0 mm long P. leptopus

Perotis leptopus Pilg.

Delicate annual; loosely tuft-
ed; 250-600 mm tall. Leaf blades
10-40 mm long; 2-5 mm wide.

Spikelets 2. 5-3.0 mm long.

Spikelet base stipitate; awns
20-40 mm long, green (rarely
purple-tinged).

Flowering February to March.

Open places in savanna wood-
land. Conservation status not known. Biome: Savanna.
North to Tanzania.

Description: Clayton et al. 1970-1982 (395). Voucher:
Joubert & Du Toil 8. PRECIS code 9902800-00100.

Perotis patens Gand.

Bottlebrush grass; purple
spike grass.

Short-lived perennial, or an-
nual; loosely tufted; 200-600 mm
tall. Leaf blades 10-70 mm long;

3-12 mm wide. Spikelets 1 .2-2.7
mm long. Spikelet base flat or
rounded, not elongated into a
stipe; glumes with purple awns 9-17 mm long.

Flowering throughout the year. Dry, poor or sandy soils,
in bare ground and disturbed places. Common. Biome: Sa-
vanna and Grassland. Also in tropical Africa and
Madagascar. Weed (frequently ruderal).

Description: Chippindall 1955 (109), Clayton et al.
1970-1982 (394). Illustration: Chippindall 1955 (fig. 82).
Voucher: Killick & Strey 2458. PRECIS code

9902800-00200.

Perotis vaginata Hack.

Annual; loosely tufted; 120-
400 mm tall. Leaf blades 10—40
mm long; 1. 5-6.0 mm wide.

Spikelets 3. 5-5. 5 mm long.

Culms robust; spikelet base stipi-
tate; awns 1 3—25 mm long, green.

Flowering February to April.

Open savanna, in sandy soil. Con-
servation status not known.

Biome: Savanna. North to Zaire and Tanzania.

Description: Clayton et al. 1970-1982 (397). Voucher:
De Winter & Wiss 4342. PRECIS code 9902800-00300.

Phacelurus Griseb.

Pseudophacelurus (Steud.) A. Camus.

Perennial. Culms 200-600 mm high; herbaceous
(robust, tough); unbranched above. Leaf blades linear; flat,
or folded (or rarely terete). Ligule an unfringed membrane.
Plants bisexual, with bisexual spikelets. The spikelets of
sexually distinct forms on the same plant ; overtly hetero-
morphic (pedicellate spikelets usually smaller or vestigial),
or homomorphic.

Inflorescence of spike-like main branches (usually
terminal, of flattened ‘racemes’ (rarely solitary, or on
elongated axis))', digitate or subdigitate (usually)', espathe-
ate; not comprising ‘partial inflorescences’ and foliar
organs. Spikelet-bearing axes ‘racemes’ (spicate); clustered
(on a common axis, rarely solitary); with substantial
rachides (clavate or inflated); disarticulating at the joints.
‘Articles’ with a basal callus-knob.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the

Fig. 164. PI. 150.

267

Fig. 165. Phacelurus franksae

rachis (not articulated, by contrast with Pseudovossia ). The
sessile spikelets hermaphrodite. The pedicellate spikelets
hermaphrodite (rarely), or male-only, or sterile. Female-
fertile spikelets compressed dorsiventrally (dorsally flat,
convex or rarely concave); falling with the glumes. Hairy
callus absent ( callus truncate, minute ). Glumes two; more
or less equal; awnless; very dissimilar (leathery to
membranous: G1 2-keeled, flat, G2 cymbiform). Proximal
incomplete florets 1; male, or sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire ; awnless. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Rottboelliinae. 7 species. Africa to Indo-China and
Japan. Helophytic to mesophytic; in shade, or in open
habitats (woodland and grassland, in moist places);
glycophytic. Natal. 1 indigenous species.

References. 1. Clayton. 1978. Kew Bull. 33: 175.

Species treatment by G.E. Gibbs Russell.

Phacelurus franksae (J.M. Wood) Clayton

Fig. 165. PI. 151.

(=Ischaemum franskae J.M.

Wood) 1.

Perennial; tufted; 200-600
mm tall. Leaf blades setaceous or
to 1 mm wide. Spikelets (sessile)

6-8 mm long. Lower glumes with
short stiff tubercle-based hairs on
the prominent nerves; pedicels
and rachis internodes swollen.

Flowering October to January. Mountain grassland, in
burned veld, alt. 1700-2600 m. Rare. Biome: Afromontane.
The spikelets are superficially similar to those of
Andropogon brazzae.

pindall 1955 (fig. 399). Voucher: Killick 1070. PRECIS
code 9900180-00100.

Phalaris L.

Baldingera Gaertn., Meyer & Scherb., Digraphis Trim,
Endallex Raf., Phalaridantha St-Lager, Phalaroides Wolf,
Typhoides Moench.

Annual, or perennial; long-rhizomatous, or caespitose,
or decumbent (some species reedlike). Culms 100-2000
mm high; herbaceous; unbranched above. Leaf blades linear
to linear-lanceolate; flat. Ligule an unfringed membrane.
Plants bisexual, with bisexual spikelets. The spikelets of
sexually distinct forms on the same plant (rarely), or all
alike in sexuality.

Inflorescence paniculate', open (rarely), or contracted;
espatheate . Spikelet-bearing axes persistent.

Female-fertile spikelets 3. 5-9. 5 mm long; compressed
laterally (strongly)', disarticulating above the glumes, or
falling with the glumes, or not disarticulating. Glumes two;
more or less equal', about equalling the spikelets to much
exceeding the spikelets; awnless; similar (papery).
Proximal incomplete florets usually 1 or 2.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes; entire; 5 nerved; awnless. Palea present; rela-
tively long; keel-less. Lodicules 2; membranous; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum long-linear;
embryo large (up to a third of the grain), or small.

Cytology, classification, distribution. Chromosome base
number, x - 6 and 7. Pooideae; Poodae; Aveneae. 16
species. North temperate, South America. Helophytic, or
mesophytic; in open habitats (in weedy places, damp soils
and swamps). Transvaal, Orange Free State, Natal, Lesotho,
and Cape Province. 6 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Anderson. 1961. Iowa State J. Sci. 36: 1.

Species treatment by M. Koekemoer.

1(0). Fertile spikelet surrounded by six sterile spikelets,
often reduced to clavate knobs; spikelets falling in
groups of seven; fertile floret glabrous or with only
a few hairs at the base of the sterile florets; sterile
florets reduced (0. 1-0.2 mm long) . P. paradoxa
Fertile spikelets without surrounding sterile spikelets;
spikelets falling individually or in variable groups
with more than one fertile spikelet; fertile floret
densely or sparsely pubescent; sterile florets
usually at least 1/3 as long as fertile florets .... 2
2(1). Sterile floret one, well developed or reduced, rarely

268

with also a much reduced second floret shorter than

0.5 mm 3

Sterile florets two, more or less equal 4

3(2). Plants annual; sterile floret either reduced to 0.3 mm
or 1.0-1. 8 mm long; glume wings usually toothed

or erose P. minor

Plants perennial; sterile floret well developed, 1 .0-2.2
mm long, rarely with a second floret less than 0.5

mm long; glume wings usually entire

P. aquatica

4(2). Glumes broadly winged, wing broadening upwards;
sterile florets 2. 5-4. 5 mm long . . P. canariensis
Glumes wingless or very narrowly winged, wing of
even width throughout; sterile florets 0.7-2. 3 mm

long 5

5(4). Glumes abruptly tapering at the tips, winged; panicle
narrowly cylindrical, 6-12 mm wide; spikelets
tightly appressed to the central axis; sterile florets

0.7-1 .5 mm long P. angusta

Glumes gradually tapering to the tips, wingless;
panicle broadly cylindrical, often tapering to the tip
and interrupted at the base, 10-30 mm wide;
spikelets loosely appressed to the central axis;
sterile florets 1.2-2. 3 mm long . P. arundinacea

Fig. 166. Phalaris aquatica

Phalaris angusta Nees ex Trin.

Annual; tufted; to 1500 mm
tall. Leaf blades 50-300 mm
long; 3-12 mm wide. Spikelets
2. 9-5. 5 mm long. Panicle 60-150
mm long, 6-12 mm wide,
narrowly cylindric; glumes very
narrowly winged, tapering
abruptly to the tip; female-fertile
floret densely pubescent, sterile
florets two, 0.7-1. 5 mm long.

Flowering September to December. An adventive in
cultivated and fallow lands. Infrequent. Naturalized from
South America. Biome: Fynbos and Savanna. North and
South America. Weed.

Description: Anderson 1961 (61), Chippindall 1955
(90). Voucher: Salter 9054. PRECIS code 9901630-00100.

Fig. 166. PI. 152.

Phalaris aquatica L.

(-P. nodosa L.) 2; (=P.
tuberosa L.) 2.

Towoomba Canary grass.

Perennial; loosely tufted (lat-
eral culms geniculately ascend-
ing); 500-1500 mm tall. Leaf
blades 180-350 mm long; 2-15
mm wide. Spikelets 4. 5-7. 5 mm

long. Panicle 15-1 10 mm long, 10-25 mm wide, occasion-
ally interrupted at the base; glumes broadly winged, margin
entire; female-fertile floret densely pubescent; sterile floret
usually one, 0.2-2. 2 mm long (rarely with a second sterile
floret, which is then shorter than 0.5 mm).

Flowering November to April. Wet ground by streams
and channels or as a weed in moist fields. Locally common.
Naturalized from the Mediterranean region. Biome: Fynbos
and Grassland. Mediterranean region eastwards to Iraq.
Introduced and cultivated in India, Africa, Australia and
North America. Planted winter pasture.

Description: Anderson 1961 (43), Bor 1985 (1773),
Chippindall 1955 (90), Clayton et al. 1970-1982 (97).
Illustration: Chippindall 1955 (fig. 60). Voucher: Oliver
6207. PRECIS code 9901630-00200.

Phalaris arundinacea L.

Reed Canary grass.

Perennial; rhizomatous (rhi-
zomes scaly, creeping); 600-
1500 mm tall. Leaf blades 50-
200 mm long; 5-15 mm wide.

Spikelets 3. 5-7. 5 mm long. Pani-
cle 70-160 mm long, 10-30 mm
wide, often interrupted in the
lower part; glumes wingless, gradually tapering to the tip;
female-fertile floret sparsely pubescent; sterile florets two,
1 .2-2.3 mm long.

Flowering November to April. Marshes, river banks,
swamp margins and damp hollows in upland areas. Locally
common. Naturalized from northern U.S.A. Biome: Grass-
land. Introduced worldwide. The var . picta L. has striped
leaves and is planted in gardens as an ornamental.

Description: Anderson 1961 (37), Stapf 1898-1900
(683), Hitchcock & Chase 1950 (534), Chippindall 1955
(89), Clayton et al. 1970-1982 (95). Illustration: Chippin-
dall 1955 (fig. 59), Clayton et al. 1970-1982 (fig. 32),
Hitchcock & Chase 1950 (fig. 1128). Voucher: Devenish
1387. PRECIS code 9901630-00300.

269

Phalaris canariensis L.

Common Canary grass.

Annual; tufted (culms usually
fascicled, erect or geniculately
ascending); 300-600 mm tall.

Leaf blades 100-260 mm long;

3-12 mm wide. Spikelets 7-8 mm
long. Panicle 1 5 — 40 mm long,

10-15 mm wide; glumes promi-
nently winged, wing broadening upwards; female-fertile
floret pilose; sterile florets two, more or less equal, 2. 5-4. 5
mm long, broad and somewhat chaffy.

Flowering October to December. Ruderals of disturbed
areas such as waste places, road verges, cultivated lands and
pastures. Locally common. Naturalized, possibly a native
of northwest Africa and the Canary Islands. Biome: Fynbos
and Grassland. Cultivated in many countries. Commercially
cultivated seed for cage birds, weed. Very similar to P.
minor , which has a single sterile floret and a much narrower
wing on the glume.

Description: Anderson 1961 (57), Bor 1985 (1771),
Chippindall & Crook 1976 228), Hitchcock & Chase 1950
(531), Chippindall 1955 (90). Illustration: Hitchcock &
Chase 1950 (fig. 1118). Voucher: Dryfhout 701. PRECIS
code 9901630-00500.

Phalaris minor Retz.

Small Canary grass.

Annual; loosely tufted (culms
erect or geniculately ascending);

(10-)200-1000 mm tall. Leaf
blades 50-250 mm long; 5-10
mm wide. Spikelets 4-6 mm long.

Panicle 20-50 mm long, 10-15
mm wide; glumes evenly winged,
keel denticulate-undulate; female-fertile florets pilose;
sterile floret one, 1.0-1. 8 mm long (occasionally 0.2-0. 3
mm long).

Flowering September to January. Disturbed areas such
as cultivated and fallow lands, roadsides and waste places,
often in damp situations. Locally common. Naturalized
from the Mediterranean. Biome: Fynbos, Grassland, Nama-
Karoo, and Succulent Karoo. Introduced weed in most
temperate regions and in the tropics. Ruderal weed. Very
similar to P. canariensis , which has two sterile florets and
broader glume wings.

Description: Anderson 1961 (31), Bor 1985 (1772),
Chippindall & Crook (228), Stapf 1898-1900 (682), Hitch-
cock & Chase 1950 (532), Chippindall 1955 (90).
Illustration: Chippindall 1955 (fig. 61), Hitchcock & Chase
1950 (fig. 1121). Voucher: Relief & Reid 483. PRECIS
code 9901630-00600.

Phalaris paradoxa L.

(-P. paradoxa L. var.
praemorsa Coss. & Dur.) 2.

Annual; tufted (culms fascic-
led, erect or geniculately ascend-
ing); 300-600(-1000) mm tall.

Leaf blades 50-300 mm long; 2-6
mm wide. Spikelets 5. 5-8. 2 mm
long. Panicle 20-70 mm long,

10-25 mm wide, often enclosed in an inflated leaf sheath;
spikelets borne and falling in units of 6-7 with one female-
fertile spikelet surrounded by 5-6 sterile spikelets, often
reduced to clavate knobs; female-fertile spikelets with
reduced sterile florets.

Flowering August to November. In moist, often poorly
drained soils near ponds or irrigation channels, also in
cultivated and fallow lands. Rare. Naturalized from the

Mediterranean. Introduced and naturalized in many
temperate regions worldwide.

Description: Anderson 1961 (22), Bor ( 1772), Hitchcock
& Chase 1950 (530), Chippindall 1955 (89). Voucher:
P.C.V. du Toil 1867. PRECIS code 9901630-00700.

Phragmites Adans.

Czernya Presl, Miphragtes Nieuwland, Oxyanthe Steud.,
Trichoon Roth, Xenochloa Roem. & Schult.

Perennial; long-rhizomatous and long-stoloniferous (
reeds, often forming dense stands). Culms 600-4000 mm
high (-10 000 mm); woody and persistent to herbaceous
(often somewhat persistent); branched above (especially
when main culm damaged), or unbranched above. Sheath
margins free. Leaf blades linear-lanceolate to lanceolate;
flat, or rolled (convolute). Ligule a fringe of hairs.

Inflorescence paniculate; open (200-600 mm long,
plumose, the fertile lemmas surrounded by long white silky
hairs); espatheate. Spikelet-bearing axes persistent.

Spikelets not in distinct ‘long-and-short’ combinations;
9-16 mm long; compressed laterally; disarticulating above
the glumes (at least above the LI). Glumes two; very
unequal; markedly shorter than the spikelets; awnless;
similar (membranous). Incomplete florets both distal and
proximal to the female-fertile florets. Distal incomplete
florets merely underdeveloped. Proximal incomplete florets
1, paleate, male. The proximal lemmas awnless.

Female-fertile florets (2-)3-10. Lemmas similar in
texture to the glumes (membranous); hairless; 1-3 nerved;
entire; awnless, or awned (narrow-attenuate, muticous to
aristulate). Awns (if lemmas aristulate) 1; median; apical;
non-geniculate; much shorter than the body of the lemma.
Palea present; conspicuous but relatively short; 2-nerved.
Lodicules 2; fleshy; ciliate, or glabrous. Stamens 3 (or two
in the lower floret). Ovary glabrous. Fruit small; hilum
short; pericarp fused; embryo large.

Photosynthetic pathway. C3; XyMS-r.

Cytology, classification, distribution. Chromosome base
number, x = 12. Arundinoideae; Arundineae. 3 species.
Cosmopolitan. Helophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 2 indigenous species.

References. 1. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf blades with tips attenuate, flexuous; leaves
deciduous at base of blade, leaving sheaths behind
on culm; upper glume 6-9 mm long . P. australis
Leaf blades with tips sharp and pungent; leaves
deciduous at base of sheath, old culms therefore
bare; upper glume 3-5 mm long . P. mauritianus

Phragmites australis (Cav.) Steud.

Fig. 167. PI. 153.

( =P . communis Trin.) 1.

Perennial; long rhizomatous;

600^1000 mm tall. Leaf blades to
350 mm long; to 35 mm wide.

Spikelets 10-18 mm long.

Robust, culms solitary, not tiller-
ing; leaves cauline, deciduous at
base of blade; leaves with long
tapering ligule with fringing hairs; hairs equaling or longer
than membranous base; inflorescences compact, 120^100
mm long; upper glume 5-9 mm long; lemmas glabrous.

270

Fig. 167. Phragmites australis

Flowering December to June. Riverbeds and wet places.
Common, or locally dominant (in riverbeds). Biome: Fyn-
bos, Savanna, Grassland, Nama-Karoo, and Desert.
Cosmopolitan. Domestic use (basketry).

Description: Chippindall 1955 (228). Illustration: Chip-
pindall 1955 (fig. 202). Voucher: Burtt Davy H610.
PRECIS code 9902140-00100.

Phragmites mauritianus Kunth

Perennial; long-rhizomatous, to
5000 mm tall. Leaf blades to 300
mm long; to 30 mm wide. Spike-
lets 7-15 mm long. Robust, culms
tillering from lower nodes; leaves
cauline, deciduous at base of
sheath; blades, with sharp, rigid
tips; ligule with fringing hairs
equaling or longer than mem-
branous base; inflorescence broad and lax, 200-400 mm
long; upper glume 3-5 mm long; lemmas glabrous.

Flowering January to June. River beds. Common, or lo-
cally dominant (in riverbeds). Biome: Savanna, Grassland,
and Desert. Tropical Africa. Domestic use (basketry).

Description: Chippindall 1955 (229). Voucher: Galpin
13534. PRECIS code 9902140-00200.

Poa L.

Arctopoa (Griseb.) Probat., Neuropoa Clayton, Oreopoa
Grand., Paneion Lunell, Parodiochloa C.E. Hubb.,
Poagrostis Raf.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 40-1500 mm
high; herbaceous; unbranched above. Sheath margins
joined, or free. Leaf blades linear, or linear-lanceolate
(often ending in a boat-shaped tip); nearly always narrow,
flat, or folded (or canaliculate), or rolled (involute or
convolute). Ligule an unfringed membrane, or a fringed
membrane (rarely).

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets not secund', 2-11 mm long; compressed
laterally; disarticulating above the glumes. Glumes two;
more or less equal ; markedly shorter than the spikelets; de-
cidedly shorter than the adjacent lemmas ; awnless; similar
(membranous). Lower glume 1 nerved, or 3 nerved. Upper
glume 3 nerved (usually). All florets female-fertile, or distal
incomplete florets also present, merely underdeveloped;
proximal incomplete florets absent.

Female-fertile florets 2-I0(-15 ) (very rarely only one).
Lemmas similar in texture to the glumes; 5 nerved (usually),
or 7-11 nerved (rarely: e.g. in the Australian Neuropoa );
entire ; pointed ; awnless (except in the southern South
American P. flabellata. which has a 2mm terminal awn).
Palea present; relatively long, or conspicuous but relatively
short, or very reduced. Lodicules 2; membranous; nearly
always glabrous (occasionally ciliolate). Stamens 3. Ovary
glabrous. Fruit small; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. About 500
species. Cosmopolitan. Helophytic (rarely), or mesophytic
(mostly), or xerophytic (rarely); in shade and in open
habitats (typically in grasslands and meadows); mostly
glycophytic, or maritime-arenicolous (a few, e.g. P.
macrantha, P. confinis ). Namibia, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho and Cape Province.
Indigenous species (3), naturalized species (3).

References. 1. Chippindall. 1955. Gr. & Past. 2. Author.
1980. FI. Europ. 3. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

271

1(0). Spikelets viviparous, parts much distorted, often
enlarged and elongated; culms bulbous at the base

P. bulbosa

Spikelets not viviparous, parts normal; culms not

bulbous at the base 2

2(1). Panicle linear to subspiciform, branches solitary,
appressed to the central axis, usually more than
their own length apart; branches with spikelets over

the whole length P. leptoclada

Panicle ovate to pyramidal, branches solitary or in
fascicles of up to 6, spreading slightly or
horizontally from the central axis, less than their
own length apart; branches with spikelets in the

upper 1/2 3

3(2). Plants annual or biennial; leaves flaccid; anthers

shorter than 1 mm P. annua

Plants perennial; leaves firm; anthers longer than 1

mm 4

4(3). Lemmas glabrous or sparsely ciliate at the base;
panicle with lowest branches solitary or paired;
plants rhizomatous, rhizomes oblique; basal
sheaths splitting into fibres P. binata

Lemmas woolly at the base; panicle with lowest
branches whorled; plants rhizomatous or
stoloniferous; basal sheaths usually not fibrous . 5
5(4). Plants rhizomatous; rhizomes stout, smooth, creeping;
ligules truncate, to 2 mm long; lemma pilose on

keel and marginal veins P. pratensis

Plants stoloniferous; stolons leafy and slender; ligules
ovate or oblong-acute, 4-6 mm long; lemma pilose
on keel only P. trivialis

Poa annua L.

Annual bluegrass.

Annual (sometimes biennial);
loosely or compactly tufted
(culms usually geniculate at the
base); 25-300 mm tall. Leaf
blades 20— 50(— 1 40) mm long;

1-5 mm wide. Spikelets 4-6 mm
long. Leaf blades flaccid; panicle
roughly pyramidal, 10-120 mm long; branches solitary or
paired, spreading horizontally or almost so at maturity;
spikelets aggregated in upper 1/2 of branches, 3-5-
flowered; anthers 0. 6-0.8 mm long.

Flowering throughout the year (usually in the rainy
season of a particular region). Damp places on roadsides,
gardens and waste land or other disturbed areas. Common.
Naturalized from Europe. Biome: Fynbos, Savanna, Grass-
land, and Desert. Europe and Mediterranean region
eastwards to India and central Asia, introduced worldwide.
Weed (in moist disturbed places). Other Poa species in our
area are perennial and have longer anthers.

Description: Bor 1985 (1745), Linder (46), Stapf
1898-1900 (715), Hitchcock & Chase 1950 (105), Chippin-
dall 1955 (53), Clayton et al. 1970-1982 (49). Illustration:
Chippindall 1955 (fig. 22), Hitchcock & Chase 1950 (fig.
167). Voucher: Smook 3576. PRECIS code 9904070-
GO 100.

Poa binata Nees

( -P . atherstonei Stapf) 3;

{=P. heterogama Hack.) 3.

Perennial; rhizomatous (rhi-
zome oblique), or tufted;

150-600 mm tall. Leaf blades
30-200 mm long; 1-5 mm wide.

Spikelets 4-6 mm long. Old leaf
sheaths split into fibres; panicle
ovate to pyramidal, 50-150 mm long, branches solitary or
binate, less than their own length apart; spikelets
aggregated on the upper 1/2 of the branches, 3-5-flowered;
lemmas glabrous at the base.

Flowering September to May. Along mountains and
escarpment in moist areas. Common. Biome: Grassland and
Nama-Karoo. Northwards into Zimbabwe. Very closely
related to P. pratensis and P. trivialis , which have lemmas
woolly at the base.

Description: Linder (50), Stapf 1898-1900 (714), Chip-
pindall 1955 (53). Illustration: Chippindall 1955 (fig. 23).
Voucher: Stirton 5421. PRECIS code 9904070-00400.

Poa bulbosa L.

(=P. vivipara (L.) Willd.) 3.

Bulbous bluegrass.

Perennial; tufted; 150-300
(-500) mm tall. Leaf blades
20-90(-150) mm long, filiform;

1-2 mm wide. Spikelets 4-6 mm
long. Roots fibrous; culms

Fig. 168. PI. 154.

272

bulbous at the base, covered with scarious remains of old
sheaths; most leaves basal, much longer than those along
the culm; ligule 4-6 mm long; spikelets 3-6-flowered,
nearly always viviparous with distorted and enlarged floret
parts; lemmas sparsely scabrid on the keels, 3-5(-7) mm
long, developing into a leaf in the older florets; anthers
1 .0-1 .5 mm long.

Flowering July to November. On gravelly well-drained
soils in damp situations such as streamsides and around
seasonal pans. Infrequent to locally common. Biome:
Nama-Karoo and Succulent Karoo. Possibly endemic and
introduced to Europe and western Asia. The viviparous
spikelets easily distinguish this species from other Poa
species in our area.

Description: Bor 1985 (1741), Linder (47), Stapf
1898-1900 (712), Hitchcock & Chase 1950 (123), Chippin-
dall 1955 (53). Illustration: Chippindall 1955 (fig. 21),
Hitchcock & Chase 1950 (fig. 213). Voucher: C.M. van
Wyk 1407. PRECIS code 9904070-00450.

Poa leptoclada A. Rich.

Perennial; straggling or com-
pactly tufted; 200-600 mm tall.

Leaf blades 20-120 mm long,
filiform; 0.5— 4.0 mm wide.

Spikelets 3.0-4.5(-6.0) mm long.

Lower leaf sheaths sometimes
fibrous; panicle linear to subspi-
ciform, 50-190 mm long, branch-
es solitary, appressed to central
axis, usually more than their own length apart; spikelets
2-5-flowered, borne throughout the length of the branches;
anthers 0. 5-1.0 mm long.

Flowering around July. Wet places in the Drakensberg
range. Extremely rare. Biome: Grassland. Northwards on
the tropical African mountains to Ethiopia and Cameroun.
The panicle is very different from other Poa species, which
have branches less than their own length apart and spikelets
in the upper half.

Description: Linder (52), Clayton et al. 1970-1982 (47).
Voucher: Hilliard & Burtt 17708 (NU). PRECIS code
9904070-00550.

Poa pratensis L.

{-P. bidentata Stapf) 3.

Kentucky bluegrass , meadow
grass.

Perennial; loosely to compact-
ly tufted, or rhizomatous (rhi-
zome long and wiry); 250-600
(-800) mm tall. Leaf blades
60-250 mm long; 2-5 mm wide. Spikelets 3. 0-5. 5 mm
long. Ligules truncate, to 2 mm long; panicle ovate, 50-200
mm long, lowest branches whorled; spikelets 2-5-flowered,
aggregated on the upper part of branches; lemma keel and
marginal veins pilose; anthers 1. 5-2.0 mm long.

Flowering September to January (and April). Moist
shady areas, usually in mountains. Locally common. Nat-
uralized from Europe. Biome: Fynbos and Grassland. From
Europe and Mediterranean region eastwards to central
Asia, introduced elsewhere. Valuable pasture (cultivated to
a limited extent), or ornamental (as lawns, but needs fertile
soil and plenty of moisture). Closely related to P. binata,
which has lemmas glabrous or sparsely ciliate on the keel
and the lowest panicle branches solitary or paired, and to
P. trivialis , which is stoloniferous, has ligules longer and
lemmas pilose on the keels only.

Description: Bor 1985 (1744), Linder (49), Hitchcock &
Chase 1950 (112), Chippindall 1955 (51). Illustration:
Hitchcock & Chase 1950 (fig. 181). Voucher: Devenish
1 158. PRECIS code 9904070-00600.

Poa trivialis L.

Roughstalk bluegrass.

Perennial; loosely tufted
(spreading from a decumbent
base), or rhizomatous (stolons
creeping and leafy); 200-900 mm
tall. Leaf blades 50-150 mm
long; 1-5 mm wide. Spikelets 4-5
mm long. Ligule ovate or oblong-
acute, 4-6 mm long; panicle ovate or pyramidal, 75-200
mm long, lowest branches whorled; spikelets 3-5-flowered,
aggregated on upper part of branches; lemma pilose on keel
only; anthers about 1.5 mm long.

Flowering December and March. Moist disturbed
places. Rare. Naturalized from Europe. Biome: Grassland.
Temperate areas worldwide. Closely related to P. binata,
which has lemmas glabrous or sparsely ciliate and lowest
panicle branches solitary or binate, and to P . pratensis,
which is rhizomatous, has shorter ligules and lemmas that
are pilose on the keels and marginal veins.

Description: Bor 1985 (1743), Linder (48), Stapf
1898-1900 (714), Hitchcock & Chase 1950 (116).
Illustration: Hitchcock & Chase 1950 (fig. 190). Voucher:
Meredith PRE34056. PRECIS code 9904070-00650.

Pogonarthria Stapf

Annual, or perennial; caespitose. Culms 130-2500 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear; flat, or rolled (convolute). Ligule a
fringed membrane, or a fringe of hairs.

Inflorescence of spike-like main branches (a raceme of
numerous, up-curved, spike-like branches)', non-digitate
(the branches tending to whorls); espatheate. Spikelet-
bearing axes disarticulating', falling entire (the racemes
falling after the spikelets have broken up).

Spikelets solitary; biseriate; 3. 3-7. 8 mm long; com-
pressed laterally; disarticulating above the glumes; not dis-
articulating between the florets, or disarticulating between
the florets (disarticulating between the lemmas, or the
glumes and lemmas falling irregularly to leave the paleas
on the persistent rachilla). Glumes two; very unequal (Gl
about 2/3 of G2)\ markedly shorter than the spikelets;
awnless; similar (rigidly membranous). Incomplete florets
distal to the female-fertile florets, merely underdeveloped,
awnless; proximal incomplete florets absent.

Female-fertile florets 2-8 ( decreasing in size upwards).
Lemmas similar in texture to the glumes; without a germin-
ation flap; 3 nerved; entire; awnless (but sometimes
subaristate). Palea present; relatively long. Lodicules 2;
fleshy (but narrow); glabrous. Stamens 3. Ovary glabrous.
Fruit small (0.5-1 mm long); ellipsoid; hilum short; peri-
carp fused; embryo large (about 1/2 grain length).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven to even (more even
in P. fleckii than in P. squarrosa). PCR sheath extensions
present, or absent. Maximum number of extension cells
when present 1. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 4 species. Tropical and southern
Africa. Mesophytic to xerophytic; in open habitats (savanna
grasslands, often in shallow or sandy soils or in disturbed
places); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 3 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1974. FTEA.

Species treatment by M. Koekemoer.

273

1(0). Plants perennial, sometimes with a short rhizome;
racemes ascending or spreading, often falcately
curved upwards; spikelets 4-10-flowered; lower
glume 0.8-1. 5 mm long; upper glume 1.6-2. 3 mm
long P. squarrosa

Fig. 169 . Pogonarthria squarrosa

Plants annual, tufted; racemes spreading up to 90
degrees from the main axis, seldom falcately
curved; spikelets 4-6-flowered; lower glume

1.2-3. 2 mm long; upper glume 2.4-3. 2 mm long

2

2(1). Plants densely tufted, decumbent, not robust, culms
and leaves hairy; spikelets 4-5-flowered; lower
glume 1.2-2. 3 mm long; upper glume 2.4-3. 2 mm

long; known from Namibia and elsewhere

P. fleckii

Plants loosely tufted, erect, robust; culms and leaves
glabrous; spikelets 5-6-flowered; lower glume
2. 1-3.2 mm long; upper glume 3. 2^1.8 mm long;
known only from Namibia P. leiarthra

Pogonarthria fleckii (Hack.) Hack.

Annual; densely tufted;

130^120 mm tall. Leaf blades
60-180 mm long; 3-6 mm wide.

Spikelets 5-10 mm long.

Vegetative parts covered with
bulbous-based hairs; spikelets

4- 5-flowered; lower glume

1. 2-2.4 mm long; upper glume
2.4-3. 2 mm long.

Flowering March to May. Sandy or well-drained soil in
open bush or mopane veld, often in disturbed areas. Com-
mon. Biome: Savanna and Desert. Zimbabwe.

Description: Launert 1970 (160:153), Chippindall 1955
(185). Voucher: Seydel 4290. PRECIS code 9903340-
00100.

Pogonarthria leiarthra Hack.

Annual; loosely tufted;

250-800 mm tall. Leaf blades
70-200 mm long; 3—4 mm wide.

Spikelets 5-8 mm long. Culms
and leaves glabrous; spikelets

5- 6-flowered; lower glume

2. 1- 3.2 mm long; upper glume

3.2 - 4.8 mm long.

Flowering February to March.

Red sand. Rare. Biome: Savanna. This species looks like a
more robust and glabrous form of P. fleckii.

Description: Launert 1970 ( 160: 153), Hackel 1912 Mitt,
bot. Mus. Univ. Zurich. Jahr. 57, Chippindall 1955 (185).
Voucher: Schoenfelder S.575. PRECIS code 9903340-
00200.

Pogonarthria squarrosa (Roem. & Schult.) Pilg.

Herringbone grass, sekelgras.

Perennial; tufted (or with a
short rhizome); 270-1400 mm
tall. Leaf blades 40-330 mm
long; 2. 0-5. 5 mm wide. Spikelets

3. 3- 7. 8 mm long. Leaf sheaths
glabrous, racemes usually falcate-
ly curved upwards; spikelets
4- 1 0-flowered; lower glume 0.8- 1 .5 mm long; upper glume
0.8-1 .5 mm long.

Flowering November to May. Open veld or under trees,
in light sandy soil, often in disturbed places. Common.
Biome: Savanna, Grassland, and Nama-Karoo. Tropical
Africa. Occasional weed.

Description: Chippindall & Crook 1976 (175), Launert
1970 (160:154), Chippindall 1955 (185), Clayton et al.
1970-1982 (267). Illustration: Chippindall 1955 (fig. 159),
Clayton et al. 1970-1982 (fig. 73). Voucher: Smook 2639.
PRECIS code 9903340-00300.

Fig. 169. PI. 155.

274

Polevansia De Winter

Perennial; long-rhizomatous and long-stoloniferous
(mat-forming, with long decumbent stems). Culms 40-450
mm high; herbaceous; unbranched above. Leaf blades linear
to linear-lanceolate; flat. Ligule a fringed membrane
(minutely fimbriate).

Inflorescence of spike-like main branches (of oppressed
racemes, 20-30 mm long)', contracted; espatheate. Spikelet-
bearing axes ‘racemes' ; persistent.

Spikelets solitary; pedicellate ; 3. 5-4. 5 mm long; com-
pressed dorsiventrally; disarticulating above the glumes.
Glumes two; very unequal; long relative to the adjacent
lemmas (i.e., the upper glumes); awnless; very dissimilar
(G1 hyaline-membranous, nerveless, obtuse, G2 lanceolate,
firmly membranous, 1 nerved). All florets female-fertile;
proximal incomplete florets absent.

Female-fertile florets l . Lemmas similar in texture to the
glumes (cf. G2); without a germination flap; 3 nerved;
entire; shortly mucronate. Palea present; relatively long
(narrowly elliptic, nearly equalling the lemma). Lodicules
2; fleshy; glabrous. Stamens 3. Ovary glabrous.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
absent. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. South Africa. Mesophytic;
in open habitats (mountain grassland); glycophytic. Orange
Free State, Lesotho, and Cape Province. 1 indigenous
species.

Fig. 170. Polevansia rigida

References. 1. De Winter. 1966. Bothalia 9: 130.

Species treatment by M. Koekemoer.

Polevansia rigida De Winter

Fig. 170. PI. 156.

Mat-forming perennial; sto-
loniferous; 100-410 mm tall.

Leaf blades 10-30 mm long;

1 .5-2 mm wide. Spikelets 3.5— 4.5
mm long. Leaf sheaths loose and
overlapping; spikelets dorsally
compressed, callus short, obtuse;
glumes persistent, unequal, with
a single thick central nerve;
lemma subcoriaceous, acute and awnless.

Flowering February to May. On lands wom out by
traditional pastoralism, or on rocky outcrops, often near
water; at altitudes higher than 1250 m. Locally common.
Biome: Grassland. Very closely related to Willkommia,
which has a longer, pungent callus and the lemma thinly
membranous, acute or obtuse, mucronate.

Description: De Winter 1966 (130). Voucher: Killick
1983. PRECIS code 9903101-00100.

Polypogon Desf.

Chaetotropis Kunth, Nowodworskya Presl., Raspailia
Presl., Santia Savi.

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 20-1200 mm high; herbaceous. Leaf blades linear
to linear-lanceolate; flat (usually). Ligule an unfringed
membrane.

Inflorescence paniculate', contracted', espatheate.
Spikelet-bearing axes persistent.

Spikelets 1.5-3 mm long; compressed laterally
(somewhat);/a//z>7g with the glumes . Rachilla terminated by
a female-fertile floret. Glumes present; two; relatively

Fig. 171. Polypogon monspeliensis

275

large; more or less equal; long relative to the adjacent
lemmas (exceeding the floret); awned (usually, apically), or
awnless; similar (papery). All florets female-fertile\
proximal incomplete florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); 5 nerved; entire to incised (truncate,
finely toothed via excurrent nerves); awnless, or awned
(usually). Awns 1\ median; from the sinus, or dorsal; non-
geniculate; much shorter than the body of the lemma to
about as long as the body of the lemma. Palea present; rela-
tively long, or conspicuous but relatively short
0 Chaetotropis ). Lodicules 2; membranous; glabrous.
Stamens 3. Ovary glabrous. Fruit small; fusiform
0 Chaetotropis ), or ellipsoid; hilum short; embryo large
(rarely), or small.

Cytology, classification, distribution. Chromosome base
number, x - 1. Pcoideae; Poodae; Aveneae. 18 species.
Mediterranean, southwest Asia. Helophytic to mesophytic;
maritime-arenicolous, or halophytic, or glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, and Cape Province. Indigenous species
(2), naturalized species (2).

Intergeneric hybrids with Agrostis ( X Agropogon P.
Fount.).

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA. 3. Tutin. 1980. FI. Europ.

Species treatment by G.E. Gibbs Russell.

1(0). Glumes with long, conspicuous awns that stand out

from the panicle 2

Glumes awnless or rarely with short inconspicuous

awns 3

2(1). Awns of glumes 10-25 mm long; awns of lemmas

5-10 mm long P. strictus

Awns of glumes 4— 8(— 1 0) mm long; lemmas awnless
or with a short inconspicuous awn to 2.5 mm long

P. monspeliensis

3(1). Plant 150-600 mm tall; panicle somewhat open,
branches ascending; spikelets 1. 5-2.0 mm long;

lemmas awnless P. viridis

Plant 30-200 mm tall; panicle narrowly cylindrical,
branches appressed; spikelets 1.0- 1.4 mm long;

lemmas with a short fine awn from tip

P. griquensis

Polypogon griquensis (Stapf) Gibbs Russell ined.

( =Agrostis griquensis Stapf);

( =P . minutiflorus Pilg.).

Annual; tufted; 30-200 mm
tall. Leaf blades to 60 mm long;

1.0- 1. 5 mm wide. Spikelets

1.0- 1. 4 mm long. Panicle

cylindrical, branches appressed;
lemmas with a short fine awn
from tip.

Flowering October. Wet places. Rare. Endemic.
Transferred from Agrostis because the entire floret plus a
short stipe together form the disseminule.

Description; Stapf Kew Bull. 1897 (290), Stapf 1898-
1900 (546), Chippindall 1955 (98). Voucher: Acocks 2466.
PRECIS code 9902440-00150.

Polypogon monspeliensis (L.) Desf.

Fig. 171. PI. 157.

Annual; 60-500 mm tall. Leaf
blades 50-200 mm long; 2-8 mm
wide. Spikelets 2-3 mm long
(excluding awns). Glume awns
spreading, to 7 mm long; lemmas
awnless or with a short awn to 2.5
mm.

Flowering September to April.

Damp and disturbed places, often

in brackish soils. Common. Naturalized from Europe and
Asia. Biome: Fynbos, Savanna, Grassland, Nama-Karoo,
Succulent Karoo, Desert, and Forest. Widely naturalized.
Weed. One of the very few species recorded from all major
biomes.

Description: Chippindall 1955 (102), Clayton et al.
1970-1982 (100). Illustration: Chippindall 1955 (fig. 74),
Clayton et al. 1970-1982 (fig. 33). Voucher: Codd 636.
PRECIS code 9902440-00400.

276

Polypogon strictus Nees

Annual; tufted; 70-700 mm
tall. Leaf blades to 200 mm long;

1-5 mm wide. Glumes with awns
10-25 mm long; lemmas with
awns 5-10 mm long.

Flowering October to April.

Wet places, usually in coastal
areas. Locally common.

Endemic.

Description: Chippindall 1955 (102). Voucher:

Compton 2613. PRECIS code 9902440-00600.

Polypogon viridis (Gouan.) Breistr.

Fig. 172.

( =Agrosti_s semiverticillata
(Forssk.) C. Christ.) 4; (=P .
semiverticillatus (Forssk.)

Hyl.) 4.

Annual; 150-600 mm tall.

Leaf blades to 150 mm long; to 7
mm wide. Spikelets 1. 5-2.0 mm
long. Panicle open, branches
ascending; lemmas awnless.

Flowering September to April. Wet places, especially
riverbanks. Locally common. Naturalized from southern
Europe. Biome: Fynbos, Savanna, Grassland, Nama-Karoo,
and Desert.

Description: Tutin 1980 (5:236). Voucher: Seydel 822.
PRECIS code 9902440-00700.

Prionanthium Desv.

Chondrolaena Nees, Prionachne Nees.

Inflorescence a single spike, or a single raceme ( spike -
like)-, contracted (30-80 mm long, the axis curved beside
each spikelet); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs, or in triplets; biseriate; not
in distinct ‘long-and-short’ combinations; 3-6 mm long;
compressed laterally; disarticulating above the glumes.
Glumes two; more or less equal; about equalling the
spikelets to much exceeding the spikelets; awnless; similar
(navicular, rigid, leathery with with membranous margins
enfolding the floret). All florets female-fertile only; or
distal incomplete florets also present, merely underdevel-
oped, awnless; proximal incomplete florets absent.

Female-fertile florets 2. Lemmas less firm than the
glumes; hairy, or hairless; without a germination flap; 3
nerved ; entire; awnless. Palea present (sub-linear); rela-
tively long; 2-nerved. Lodicules 2; fleshy. Stamens 3.
Ovary glabrous.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 2-3 species. South Africa. Helophytic (in
seasonally wet places); in open habitats; glycophytic. Cape
Province. 3 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2.
Davidse. 1988. Bothalia 18: 143-153.

Species treatment by N.P. Barker.

1(0). Lemmas pubescent P. dentatum

Lemmas glabrous 2

2(1). Spikelets solitary; rachis triquetrous; glands on keel
of glumes sessile or slightly stalked (rarely absent)

P. pholiuroides

Spikelets paired (may be solitary near base and apex
of inflorescence); rachis cylindrical; glands on keel
of glumes conspicuously stalked .... P. ecklonii

Annual; caespitose. Culms 40-430 mm high; herbaceous
(slender); branched above, or unbranched above. Leaf
blades linear (or filiform); flat, or rolled. Ligule a fringe of
hairs.

Prionanthium dentatum (L.f.) Henr.

(=Prionanthium rigidum
Desv.) 1.

Annual; tufted; 30-430 mm
tall. Leaf blades 15-105 mm
long; 0. 5-3.0 mm wide. Spikelets
3. 2-5. 2 mm long; to 1.2 mm
wide. Panicle spike-like, 5-75
mm long; rachis cylindrical;
spikelets densely aggregated, laterally arranged, not
obviously paired; glumes with prominent stalked glands on
keel; lemmas pubescent; paleas pubescent between keels.

Flowering September. Nieuwoudtville area in Western
Mountain Karoo. Rare. Biome: Succulent Karoo. Endemic.
Collected again in 1975, over 200 years after it was last
collected by Thunberg, by whom the taxon was first
described.

Description: Davidse 1988 (151), Stapf 1898-1900
(455), Chippindall 1955 (271). Voucher: Davidse 33396.
PRECIS code 9901800-00050.

Prionanthium ecklonii (Nees) Stapf

Annual; tufted; 190-370 mm
tall. Leaf blades 40-160 mm
long; 0.5-1. 5 mm wide. Spikelets
4.4-6. 1 mm long; to 1.5 mm
wide. Panicle spike-like, incon-
spicuously secund, 15-95 mm
long; rachis cylindrical; spikelets
arranged alternately in pairs, but
usually solitary near base and
apex; glumes with prominent, conspicuously stalked glands
on the keel; lemmas glabrous; paleas glabrous.

277

Flowering September to October. Low altitudes in
Coastal Renosterveld. Rare. Biome: Fynbos. Endemic.
Description: Davidse 1988 (151), Stapf 1898-1900

(455) , Chippindall 1955 (271). Illustration: Davidse 1988
(fig. 2). Voucher: Ecklon & Zeyher s.n. PRECIS code
9901800-00100.

Prionanthium pholiuroides Stapf

Fig. 173. PI. 158.

Annual; tufted; 40-250 mm
tall. Leaf blades 15-70 mm long;

0.5-1. 5 mm wide. Spikelets
3. 1-7.0 mm long; to 1.5 mm
wide. Inflorescence a secund, 2-
ranked spike, 15-60 mm long;
rachis triquetrous; spikelets ar-
ranged alternately, single (rarely
in pairs); keels of glumes with
sessile glands; lemmas glabrous; paleas glabrous.

Flowering October to December. Seasonally wet,
shallow depressions. Rare. Biome: Fynbos. Endemic.
Description: Davidse 1988 (151), Stapf 1898-1900

(456) , Chippindall 1955 (271). Illustration: Chippindall
1955 (fig. 243). Voucher: Anderson 8. PRECIS code
9901800-00200.

Prosphytochloa Schweick.

Perennial; long-rhizomatous (rhizomes horizontal, with
cataphylls). Culms 10000 mm high (or more); herbaceous;
scandent (by retrorse hairs on the leaf blade margins)',
branched above. Leaf blades linear-lanceolate to lanceolate;
flat. Ligule an unfringed membrane.

Inflorescence paniculate (terminating main culm and
laterals); espatheate. Spikelet-bearing axes persistent.

Spikelets solitary; 6-9 mm long; compressed laterally
(slightly); disarticulating above the glumes (i.e. above the
rudimentary glumes). Glumes present, or absent; two (these
reduced to a bilobed to entire hyaline cup); minute; more
or less equal; awnless. Proximal incomplete florets 2;
epaleate; sterile (subulate, edged with minute hyaline
spines, variable in size).

Female-fertile florets 1. Lemmas entire; awnless; 5
nerved. Palea present (similar to the lemma, which clasps
it); relatively long; with several nerves (3). Lodicules 2;
membranous (above, but fleshy below); glabrous. Stamens
6. Ovary glabrous. Fruit medium sized (5 to 6 mm long,
brown); fusiform; hilum long-linear; embryo small.

Transverse section of leaf blade. Mesophyll with arm
cells; with fusoids (i.e. with lateral sheath extensions), or
without fusoids (if these not so interpreted). Midrib vascu-
larization complex (there being a small bundle adaxial to
the main one).

Cytology, classification, distribution. Chromosome base
number, x - 12. Bambusoideae; Oryzodae; Oryzeae. 1
species. South Africa. Helophytic; in shade; glycophytic.
Transvaal, Swaziland, Natal, and Cape Province. 1 indige-
nous species.

References. 1 . Schweickerdt. 1961 . Der Zuchter 31 : 194.

Species treatment by G.E. Gibbs Russell.

Prosphytochloa prehensilis (Nees) Schweick.

Fig. 174. PI. 159.

(- Potamophila prehensilis
(Nees) Benth.) 1 .

Perennial; climber; to 10000
mm tall. Leaf blades to 150 mm
long; 5-10 mm wide (scabrid).

Spikelets 6-9 mm long.

Inflorescence a loose panicle;
spikelets with solitary female-

fertile florets subtended by minute cuplike glumes and two
subulate sterile lemmas.

Flowering November to April. Moist forests, where it
climbs in dense masses. Infrequent. Biome: Forest.
Endemic.

Description: Chippindall 1955 (33). Illustration: Chip-
pindall 1955 (fig. 3). Voucher: Codd 8411. PRECIS code
9901561-00100.

Pseudechinolaena Stapf

Loxostachys Peter.

Annual', decumbent. Culms 100-600 mm high; herba-
ceous; branched abo ve. Leaf blades lanceolate (acuminate);
flat; pseudopetiolate', with readily visible transverse veins.
Ligule a fringed membrane . The spikelets of sexually
distinct forms on the same plant (some variously
incomplete), or all alike in sexuality.

Inflorescence of spike-like main branches (spiciform
racemes ); espatheate. Spikelet-bearing axes persistent.

Spikelets in pairs, or solitary (via suppression of one of
the pair); consistently in ‘long-and-short’ combinations, or
not in distinct ‘long-and-short’ combinations. Female-
fertile spikelets 4.6 mm long; adaxial', compressed laterally;
falling with the glumes. Glumes two; more or less equal (or
G1 shorter); awnless; very dissimilar (first smooth, upper
gibbous with translucent intercostal glands, and these often

278

with hooked spines). Proximal incomplete florets 1 ; paleate,
palea fully developed; male, or sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (papery);
smooth; not becoming indurated; hairless (sometimes with
hooks); having the margins tucked in onto the palea; with
a clear germination flap; 3-5 nerved; entire; awnless. Palea
present. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small (1.5 mm), ellipsoid; hilum short;
embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 6 species. 5 in Madagascar, 1 pantropical.
Mesophytic; in shade (forest); glycophytic. Transvaal,
Natal and Cape Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Pseudechinolaena polystachya (Kunth) Stapf

Fig. 175. PI. 160.

Soft slender perennial, or an-
nual (often mat-forming); to 400
mm tall. Leaf blades 10-80 mm
long; to 14 mm wide. Spikelets
3. 5-5.0 mm long (reduced spike-
lets often present). Culms
prostrate; spikelets burr-like;
upper glume usually with stiff
hooked hairs.

Flowering August to September and December to April.
In forest shade. Locally common. Biome: Forest.

Throughout tropics. Similar in habit to other forest grasses
in Panicum and Oplismenus , which all lack the hooklike
hairs on the upper glumes that aid in dispersing the mature
spikelets of this genus.

Description; Chippindall 1955 (365), Clayton et al.
1970-1982 (545). Illustration: Chippindall 1955 (fig. 314).
Voucher: Schweickerdt 1442. PRECIS code 9901010-
00100.

Pseudopentameris Conert

Sometimes included in Danthonia sensu lato.

Perennial; caespitose. Culms 300-1200 mm high; herba-
ceous; branched above, or unbranched above. Leaves auric-
ulate (from the base of the blade). Leaf blades linear; flat,
or rolled. Ligule a fringe of hairs.

Inflorescence paniculate (40-250 mm long); contracted
(but central axis visible); espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary; 35-55 mm long ; compressed laterally;
disarticulating above the glumes. Glumes present; two; rel-
atively large (35-55 mm long); more or less equal; about
equalling the spikelets to much exceeding the spikelets;
awnless; similar (lanceolate, membranous). Incomplete
florets distal to the female-fertile florets, merely underde-
veloped; proximal incomplete florets absent.

Female-fertile florets 2. or 3 . Lemmas decidedly firmer
than the glumes (leathery); hairy (villous); without a ger-
mination flap; 9 nerved; incised; awned. Awns 3; median
and lateral. The median awn different in form from the
laterals; from the sinus; geniculate; much longer than the
body of the lemma. Palea present (glabrous, by contrast
with Pentameris ); relatively long (exceeding the lemma
lobes); 2-nerved. Lodicules fleshy; ciliate (or at least
ciliolate), or glabrous. Stamens 3. Ovary glabrous. Fruit
medium sized (about 6 mm long); hilum long-linear (more
than half the grain length); pericarp fused.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 2 species. South Africa. Mesophytic; in open
habitats (mountain Fynbos); glycophytic. Cape Province. 2
indigenous species.

References. 1. Conert. 1971. Mitt. Bot. Stsamml.
Munch. 10: 304. 2. Ellis. 1985. Bothalia 15: 561.

Species treatment by N.P. Barker.

1(0). Lemmas pubescent over entire surface

P. brachyphylla

Lemmas basally glabrous, pubescent apically,
including the lemma lobes P. macrantha

Pseudopentameris brachyphylla (Stapf) Conert

Fig. 176.

(=Danthonia brachyphylla
Stapf) 1.

Perennial; tufted; 300-900
mm tall. Leaf blades to 150 mm
long; to 4 mm wide. Spikelets
27-37 mm long (excluding
awns); 10 mm wide (excluding
awns). Leaves clustered basally,
obviously distichous, open and flat or folded; dead leaves
tightly curled; lemma body 5. 5-6. 5 mm long, lemma lobes
10-19 mm long, including bristles into which the lobes
attenuate; lemma backs completely pubescent; central awn
geniculate, 19-30 mm long.

Flowering August to December. Rocky, gravelly or

279

Fig. 176. Pseudopentameris brachyphylla

sandy lower slopes of Cape fold mountains. Locally com-
mon (hills behind Hermanus and Betty’s Bay). Biome: Fyn-
bos. Endemic. Studied anatomically by Ellis (1985), and
found to be very similar to P. macrantha.

Description: Stapf 1898-1900 (520), Chippindall 1955
(250). Illustration: Chippindall 1955 (fig. 221). Voucher:
Zeyher 1825b. PRECIS code 9902081-00100.

Pseudopentameris macrantha (Schrad.) Conert

PI. 161.

( =Danthonia macrantha
Schrad.) 1.

Perennial; tufted; 800-1200
mm tall. Leaf blades to 500 mm
long; to 4 mm wide. Spikelets
30-45(-60) mm long (excluding
awns); 15 mm wide. Leaves
basally clustered, open or in-
volute in cross section; dead leaves falcate but not tightly

curled; lemma body 5.5-10.0 mm long, lemma lobes 10-23
mm long, including bristles into which the lobes attenuate;
lemma backs glabrous in lower half, pubescent in upper half
including lemma lobes; central awn geniculate, 15-50 mm
long.

Flowering August to December. Rocky, stony or sandy
slopes in both TMS and limestone geologies. Locally com-
mon (on the lower slopes of Table Mountain). Biome: Fyn-
bos. Endemic. Ellis (1985) considers this species to be
anatomically almost identical to P. brachyphylla.

Description: Stapf 1898-1900 (519), Chippindall 1955
(251). Illustration: Conert 1971 plate 1. Voucher: Sandwith
73. PRECIS code 9902081-00200.

Puccinellia Pari.

Atropis (Trin.) Griseb.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 40-1000 mm
high; herbaceous. Sheath margins free (but rarely closed to
almost one third their length). Leaf blades linear; flat, or
folded, or rolled. Ligule an unfringed membrane .

Inflorescence paniculate ; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets not secund', 2-13 mm long; compressed
laterally, disarticulating above the glumes. Rachilla pro-
longed beyond the uppermost female-fertile floret. Glumes
two; very unequal; markedly shorter than the spikelets; de-
cidedly shorter than the adjacent lemmas', awnless; non-
carinate', similar. Incomplete florets distal to the female-
fertile florets; proximal incomplete florets absent.

Female-fertile florets 2-10. Lemmas similar in texture
to the glumes; 5 nerved; entire (or erose, often ciliolate);
awnless. Palea present; relatively long. Lodicules 2;
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short', embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. About 80 species.
North temperate. Helophytic, or mesophytic; usually
halophytic. Namibia, Orange Free State, and Cape
Province. Indigenous species (3), naturalized species (1).

Intergeneric hybrids with Phippsia — X Pucciphippsia
Tsvelev.

References. 1. Chippindall. 1955. Gr. & Past. 2. Linder.
Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Panicle linear, 5-15 mm in diameter, branches very
slender, appressed or very slightly spreading;
spikelets usually their own length apart but not
overlapping for more than half their length .... 2
Panicle ellipsoid or pyramidal, more than 15 mm in
diameter, branches slender or stout, at least the
lower ones spreading, horizontal or reflexed;
spikelets dense, overlapping for more than half

their length 3

2( 1 ). Culms 1 -noded; lowest leaf sheaths shiny, longer than
50 mm, overlapping, enclosing the culms and lower
part of the panicle; lemmas 2. 2-2. 8 mm long . . .

P. angusta

Culms 2-noded; lowest leaf sheaths dull, shorter than
50 mm, usually not overlapping, culms and panicle
usually not enclosed; lemmas 2.0-2. 2 mm long .

P. acroxantha

3(1). Panicle pyramidal or elongate; branches slender, often
deflexed, naked in the lower half .... P. distans
Panicle ellipsoid, rather compact; branches stout,
stiff, not more than 90 degrees from the main axis,

bearing spikelets nearly to the base

P. fasciculata

280

Puccinellia acroxantha Smith & C.E. Hubb.

PI. 162.

Perennial; loosely tufted;

100-600 mm tall. Leaf blades
50-200 mm long; 1-2 mm wide.

Spikelets 3-5 mm long; to 1.5
mm wide. Culms 2-noded; lowest
leaf sheath dull, shorter than 50
mm, usually not overlapping and
enclosing culm and lower part of
panicle; panicle linear, branches
very slender, contracted; lemmas 2. 0-2. 5 mm long.

Flowering January. On Karoo-turf soil of varying
salinity, in depressions periodically flushed with fresh

water. Rare. Biome: Grassland. Endemic. The status of this
species is very uncertain, it might well be a local form of
P. distans. Further research is needed.

Description: Smith & Hubbard 1929 Kew Bull. (86),
Linder (39), Chippindall 1955 (50). Voucher: Smith 5415.
PRECIS code 9904150-00100.

Puccinellia angusta (Nees) Smith & C.E. Hubb.

Perennial; densely tufted;

300-600 mm tall. Leaf blades
75-100(-300) mm long; 1.0-1. 5
(-2.5) mm wide. Spikelets
4. 0-5. 5 mm long; 1.0-1 .5 mm
wide. Culms 1-noded; lowest leaf
sheaths shining, longer than 50
mm, overlapping and enclosing
the culms and lower part of the
panicle; panicle linear, branches appressed, slender;
lemmas 2. 2-2. 8 mm long.

Flowering August to October. In disturbed areas on
strongly saline moist soils. Infrequent to locally common.
Biome: Fynbos, Grassland and Desert. Endemic. Said to be-
a good winter pasture. The voucher and type specimen was
collected in abnormally high saline soil where few other
plants survived. One wonders if the vegetative differences
that distinguish it from P. acroxantha were induced by the
abnormal habitat.

Description: Smith & Hubbard 1929 Kew Bull. (85),
Linder (38), Chippindall 1955 (50). Voucher: Smith 4385.
PRECIS code 9904150-00200.

Puccinellia distans (L.) Pari.

Perennial; tufted; 250-650
mm tall. Leaf blades 70-180 mm
long; 2-4 mm wide. Spikelets 4—8
mm long; 1-2 mm wide. Panicle
pyramidal or elongate, branches
(at least some) naked in the lower
half, spreading, often horizontal
or deflexed: lemmas 1. 5-4.0 mm
long.

Flowering April, June, July, and October. In wet often
very saline habitats along rivers, irrigation canals and
furrows. Infrequent. Naturalized from Europe. Biome: Fyn-
bos, Nama-Karoo, and Succulent Karoo. Cosmopolitan in
temperate regions. Weed. Distinguished by its pyramidal
panicle with spikelets only in the upper half of the branches.

Description: Hughes & Halliday 1980 FI. Europ.
(5:168), Linder (37), Hitchcock & Chase 1950 (81).
Voucher: Smook 3383. PRECIS code 9904150-00250.

Puccinellia fasciculata (Torr.) Bickn.

Fig. 177.

Perennial; tufted; 200-400
mm tall. Leaf blades 80-200 mm
long; 2-5 mm wide. Spikelets 4-7
mm long; 1-2 mm wide. Panicle
ellipsoid, rather compact,
branches stout, bearing spikelets
nearly to the base; lemmas
1 .5-2.5 mm long.

Flowering September to Jan-
uary. In wet, saline habitats like salt marshes, often in
disturbed areas. Infrequent. Naturalized from Europe.
Biome: Fynbos, Nama-Karoo, and Succulent Karoo.
Europe. Distinguished by its compact panicle and stiff
branches.

Description: Hughes & Halliday 1980 FI. Europ.
(5:168), Linder (36), Hitchcock & Chase 1950 (80), Chip-
pindall 1955 (50). Illustration: Chippindall 1955 (fig. 19),
Hitchcock & Chase 1950 (fig. 112). Voucher: Adamson
2989. PRECIS code 9904150-00300.

281

Rendlia Chiov.

Sometimes included in Microchloa R. Br.

Perennial; densely caespitose (from a cushion of old,
fibrous leaf sheaths). Culms 50-350 mm high; herbaceous;
unbranched above. Leaf blades linear; to 0.7 mm wide ;
folded (at the base, the adaxial surfaces adnate). Ligule a
fringed membrane (the ‘ membrane ’ unusually firm). The
spikelets of sexually distinct forms on the same plant (the
uppermost 2-3 spikelets reduced), or all alike in sexuality.

Inflorescence a single spike (to 50 mm long — rarely a
pair of spikes)\ espatheate. Spikelet-bearing axes persistent.

Fig. 178. Rendlia altera

Female-fertile spikelets solitary; alternately biseriate;
4-5.5 mm long; with the G1 compressed obliquely, the G2
compressed dorsiventrally; disarticulating above the
persistent Gl, the G2 falling with and enveloping the
florets; not disarticulating between the florets. Rachilla
prolonged beyond the uppermost female-fertile floret.
Glumes two; more or less equal; much exceeding the
spikelets; awnless; very dissimilar (firmly membranous, the
Gl laterally compressed, the G2 dorsally compressed). In-
complete florets distal to the female-fertile florets, one per
spikelet, male or sterile, banana-shaped, hairless, the lemma
shorter and thinner than the L 1 , the palea reduced or absent;
awnless. Proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes; without a germination flap; 3 nerved; incised;
awnless. Palea present; relatively long (slightly exceeding
the lemma). Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous (suppressed in the upper floret). Fruit small
(2 mm); ellipsoid; hilum short; embryo small (seemingly,
judged from immature material).

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 1 species. Eastern tropical and
southern Africa. Mesophytic; in open habitats (shallow
soils in grasslands); glycophytic. Transvaal, Orange Free
State, Natal, and Cape Province. 1 indigenous species.

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Rendlia altera (Rendle) Chiov.

Fig. 178. PI. 163.

( =R . nelsonii (Stapf)

Chiov.) 1.

Mahem’s crest, kleinrolblaar.

Perennial; tufted (cushion-
like); 200-400 mm tall. Leaf
blades 30-250 mm long; less than
1.5 mm wide. Spikelets 4. 0-5. 5
mm long. Leaf bases persistent and becoming fibrous with
age; spike a solitary ‘toothbrush’, 20-50 mm long; glumes
twice as long as the florets.

Flowering September to May. Shallow humiferous or
well-drained sandy soils. Locally common. Biome: Grass-
land. Tropical Africa. Clayton & Renvoize (1986) put this
genus in synonymy with Microchloa. As recognized here,
Rendlia has a larger inflorescence and spikelets with two
florets.

Description: Chippindall 1955 (193), Clayton et al.
1970-1982 (331 ). Illustration: Chippindall 1955 (fig. 169),
Clayton et al. 1970-1982 (fig. 93). Voucher: Du Toit 2502.
PRECIS code 9902941-00100.

Rhytachne Desv.

Lepturopsis Steud.

Annual, or perennial; caespitose. Culms 250-1200 mm
high; herbaceous; unbranched above (few noded). Leaf
blades linear; flat, or folded, or rolled, or acicular. Ligule
an unfringed membrane (short). Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant ; overtly heteromorphic (the pedicellate
spikelets variously reduced).

Inflorescence a single raceme (of single ‘racemes’
terminating the culms, these cylindrical and culm-like until
the embedded spikelets open)', spatheate, or espatheate; not
comprising ‘partial inflorescences’ and foliar organs.

282

Spikelet-bearing axes spike-like\ solitary; with substantial
rachides; disarticulating at the joints. ‘Articles’ with a basal
callus-knob.

Spikelets solitary (accompanied by a scale-tipped
pedicel, when the pedicellate spikelet is suppressed), or in
pairs; consistently in ‘long-and-short’ combinations; these
pedicellate/sessile. Pedicels free of the rachis. The sessile
spikelets hermaphrodite. The pedicellate spikelets male-
only, or sterile (variously reduced, sometimes suppressed).

Female-fertile spikelets 2-8 mm long; compressed dorsi-
ventrally; falling with the glumes. Glumes two; more or less
equal; awned (G1 and/or G2, sometimes), or awnless; very
dissimilar (G 1 leathery, convex, often transversely
rugulose, G2 membranous or hyaline, with or without a
terminal subule). Proximal incomplete florets /; paleate,
palea reduced; male, or sterile (rarely).

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present; relatively
long, or conspicuous but relatively short, or very reduced.
Lodicules 2; fleshy. Stamens 3. Ovary glabrous.

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Rottboelliinae. 12
species. Tropical and southern Africa, Madagascar, tropical
South America. Helophytic (pans and riversides), or meso-
phytic (grasslands); in open habitats; glycophytic. Namibia,
Natal, and Cape Province. 3 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf blades setaceous; sessile spikelets 3-5 mm long,

lower glume strongly transversely rugose

R. rottboellioides

Leaf blades expanded; sessile spikelets over 5 mm
long, lower glume not transversely rugose .... 2
2(1). Leaf blades 2—4 mm wide; lower glume smooth
below; pedicel and rachis with a few hairs; pedicel
not broad and flattened, smooth, with a green line
beside each edge; pedicellate spikelets 3 mm long;

Namibia R. robusta

Leaf blades 6-10 mm wide; lower glume with strong
longitudinal nerves; pedicel and rachis lacking
hairs; pedicel broad and flattened, with many faint
nerves, entirely green; pedicellate spikelets reduced
to a scale less than 1 mm long; Natal . R. latifolia

Rhytachne latifolia Clayton

Perennial; tufted; 400-1000
mm tall. Leaf blades 150-500
mm long; 6-10 mm wide. Spike-
lets (sessile) 5.5-8 mm long (ped-
icellate reduced to a scale less
than 1 mm long). Rachis and
pedicels glabrous, pedicels broad
and flattened, with many faint
nerves, entirely green; lower
glume with strong longitudinal nerves.

Flowering January to March. Shaded streamsides and
woodland pans. Rare. North to Tanzania. Not well
separated from R. robusta, and possibly conspecific with it.

Description: Clayton et al. 1970-1982 (845). Voucher:
Tinley 895. PRECIS code 9900340-00050.

Rhytachne robusta Stapf

Fig. 179.

Perennial; tufted; about 1200
mm tall. Leaf blades to 400 mm
long; 2 -4 mm wide. Spikelets
(sessile) 5-6 mm long (pedicel-
late reduced in size, to 3 mm
long). Rachis and pedicels with
sparse hairs, pedicels not broad
and flattened, with a green line
beside each edge; lower glume of
sessile spikelets smooth below, with nerves on upper end
only.

Flowering January. Grassveld. Rare, but possibly locally
common. Biome: Savanna. Southern tropical Africa.

Description: Stapf 191V (82). Voucher: Killick &
Leistner 3287. PRECIS code 9900340-00100.

283

Rhytachne rottboellioides Desv.

PI. 164.

Slender perennial; densely
tufted; 250-1000 mm tall. Leaf
blades to 300 mm long;
setaceous. Spikelets (sessile) 3-5
mm long (pedicellate reduced to
an arista). Plant reddish or
purplish brown; lower glume
strongly transversely rugose.

Flowering November to Feb-
ruary. Vleis and swampy ground. Rare. Tropical Africa,
Madagascar and Brazil. The plant resembles Schizachyrium
sanguineum , which grows in the open veld.

Description: Chippindall 1955 (519), Clayton et al.
1970-1982 (843). Illustration: Chippindall 1955 (fig. 415),
Clayton et al. 1970-1982 (fig. 198). Voucher; Fluntley 779.
PRECIS code 9900340-00200.

Rottboellia L.f.

Stegosia Lour.

Annual-, caespitose. Culms 300-3000 mm high; herba-
ceous; branched above. Leaf blades broad-, flat. Ligule an
unfringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant-, overtly heteromorphic.

Inflorescence a single raceme, or paniculate ( with
terete, spike-like ‘racemes' , terminating the culms and
branches, or axillary, solitary or in fascicles)-, spatheate;
a complex of ‘partial inflorescences’ and intervening foliar
organs. Spikelet-bearing axes spike-like (cylindrical, with
embedded spikelets); solitary and clustered (fascicled); with
substantial rachides; disarticulating at the joints. ‘Articles’
with a basal callus-knob.

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; these ‘pedicellate’/sessile. Pedicels of the
‘pedicellate’ spikelets discernible, but fused with the rachis.
The sessile spikelets hermaphrodite. The ‘pedicellate’
spikelets male-only, or sterile, striate, compressed,
herbaceous. Female-fertile spikelets compressed dorsiven-
trally (trigonous); falling with the glumes (and with the
joint, the pedicellate spikelets falling separately). Glumes
two; more or less equal; awnless; very dissimilar (lower
flat-backed, 2-keeled above, upper naviculate, winged).
Proximal incomplete florets I; paleate, palea fully
developed; male.

Female-fertile florets 1. Lemmas less firm than the
glumes; entire; awnless. Palea present; relatively long. Lod-
icules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Rottboelliinae. 4 species. Tropical and
subtropical Africa, Asia. Helophytic to mesophytic; in
shade, or in open habitats (woodland, swamps, often in
disturbed ground or a weed of cultivated ground);
glycophytic. Namibia, Botswana, Transvaal, Swaziland,
and Natal. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Rottboellia cochinchinensis (Lour.) Clayton

Fig. 180. PI. 165.

( =R . exaltata L.f) 2.

Guineafowl grass, kokoma
grass, tarentaalgras.

Annual (often robust); 300-
3000 mm tall. Leaf blades to 600
mm long; 1 0-30 mm wide. Spike-
lets (sessile and pedicellate)

4-7 mm long. Racemes cylindrical, spikelets sunken; basal
sheaths with stiff irritating hairs.

Flowering December to June. Wet places, often on black
turf soil, and in disturbed places. Infrequent. Biome: Savan-
na. Throughout Old World tropics, introduced to America.
Weed (ruderal).

Description: Chippindall 1955 (520), Clayton et al.
1970-1982 (853). Illustration: Chippindall 1955 (fig. 416),
Clayton et al. 1970-1982 (fig. 203). Voucher: Ward 2118.
PRECIS code 9900310-00050.

Fig. 180. Rottboellia cochinchinensis

284

Sacciolepis Nash

Rhampholepis Stapf.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 100-2000 mm
high; herbaceous; branched above. Leaf blades linear to
linear-lanceolate; flat, or rolled (convolute). Ligule an
unfringed membrane to a fringed membrane . Plants
bisexual, with bisexual spike lets.

Inflorescence paniculate ; open (rarely), or contracted
(usually spicate); espatheate. Spikelet-bearing axes
persistent.

Spikelets 0.8-5. 2 mm long; compressed laterally to not
noticeably compressed (gibbous, often oblique)', falling
with the glumes. Glumes two; very unequal; awnless; very
dissimilar, or similar (both membranous or hyaline, but
upper often inflated and gibbous). Upper glume distinctly
saccate. Proximal incomplete florets 1 ; paleate, orepaleate,
palea when present fully developed to reduced; male
(rarely), or sterile.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes (papery to subcrustaceous); smooth; becoming
indurated, or not becoming indurated; hairless (glossy);
having the margins tucked in onto the palea; with a clear
germination flap; 3 nerved, or 5 nerved (obscurely so);
entire; awnless. Palea present; relatively long. Lodicules 2;
fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit small;
hilum short; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae. 30
species. Tropical and subtropical. Hydrophytic to helo-
phytic; in open habitats (in or near water or in wet places);
glycophytic. Namibia, Botswana, Transvaal, Swaziland,
Natal and Cape Province. 8 indigenous species

References. 1. Simon. 1972. Kew Bull. 27: 387. 2.
Clayton & Renvoize. 1982. FTEA.

Species treatment by M. Koekemoer.

1(0). Inflorescence a loosely contracted panicle; spikelets
distinctly pedicelled; upper glume more than 5
times the length of the lower glume . S. curvata
Inflorescence a false spike; spikelets subsessile; upper
glume less than 3 times the length of the lower

glume 2

2(1). Spikelets up to 2.7 mm long 3

Spikelets more than 2.7 mm long 5

3(2). Plants annual; spikelets 1.3-1. 9 mm long

S. huillensis

Plants perennial; spikelets 1.5-2. 2 mm long 4

4(3). Leaf blades subterete to convolute; spikelets more or
less loosely arranged on the central axis, pubescent;
lower leaf sheaths rarely with cross-veins; plants up

to 900 mm tall S. chevalieri

Leaf blades flattened or folded; spikelets more or less
tightly arranged on the central axis, glabrous or
rarely pubescent; lower leaf sheaths with cross-
veins; plants up to 1500 mm tall .... S. typhura
5(2). Leaf sheaths with prominent auricles 1.5-2. 5 mm
long; spikelets laterally compressed; plants
perennial or slender annuals; culms not spongy, less

than 5 mm in diameter 6

Leaf sheaths without auricles or auricles minute when
present; spikelets slightly dorsally compressed;
plants perennial, robust, aquatic; culms spongy at

the base, 5-18 mm in diameter 7

6(5). Plants annual, prostrate, with prominent aerial roots
from the lower nodes; culms 8-10-noded; spikelets
2.7-3. 1 mm long, with the lower glume 1. 0-1.5 mm

long and the lower palea 1 .2—1 .8 mm long

S. indica

Plants perennial, erect, with a short oblique rhizome;
culms 4-6-noded; spikelets 3.0-^L4 mm long, with

Fig. 181. Sacciolepis typhura

285

the lower glume 1. 8-2.9 mm long and the lower

palea 1.8-2. 8 mm long S. rigens

7(5). Spikelets 2.7-4. 1 mm long, obtuse to subacute, light
green to brown; lower palea 1.5-1. 8 mm long;

upper floret 2. 2-3. 2 mm long S. africana

Spikelets 3. 9-4. 5 mm long, acute to acuminate, light
green to yellowish; lower palea 2. 0-2. 2 mm long,
upper floret 3. 2-3. 5 mm long .... S. interrupta

Sacciolepis africana C.E. Hubb. & Snowden

Perennial; rhizomatous (culms
thick, spongy, often decumbent
and rooting at the nodes);

300-1800 mm tall. Leaf blades
50-400 mm long; 3-15 mm wide.

Spikelets 2.5 — 4. 1 mm long. Leaf
sheaths with cross-veins; panicle
dense, spikelike, 40-300 mm
long; spikelets obtuse to
subacute, dorsally compressed; lower glume 1/4-1/3 the
spikelet length; upper floret 2. 2-3. 2 mm long.

Llowering Lebruary to May. Standing in water in
swampy or seasonally flooded areas and along river banks.
Infrequent. Biome: Savanna. Throughout tropical Africa.
Closely related to S. interrupta, which has larger spikelets
with acute or acuminate tips.

Description: Chippindall 1955 (351), Clayton et al.
1970-1982 (455). Illustration: Clayton et al. 1970-1982
(fig. 120). Voucher: De Winter & Marais 4528. PRECIS
code 9901240-00100.

Sacciolepis chevalieri Stapf

Perennial; shortly rhizomatous
and tufted; 200-900 mm tall. Leaf
blades 50-200 mm long; usually
rolled or folded, 1-3 mm wide.

Spikelets 1.5-2. 2 mm long. Leaf
sheaths with cross-veins absent or
inconspicuous; panicle spikelike,
scanty, sometimes interrupted,

20-160 mm long; spikelets
laterally compressed; lower glume 2/3 and upper glume 4/5
the spikelet length.

Llowering October to March. In wet soils, usually black
turf, in swamps, vleis or along streams. Infrequent. Biome:
Savanna and Grassland. Throughout tropical Africa and in
Madagascar. Intergrades into S. typhura, which is a larger
plant with thicker, usually spongy culms, prominent cross-
• veins on the leaf sheaths and denser,longer panicles.

Description: Stapf 1920 (754), Clayton et al. 1970-1982
(459). Voucher: Reid 426. PRECIS code 9901240-00300.

Sacciolepis curvata (L.) Chase

Lorest hood grass, kappiegras.

Shortlived perennial, or an-
nual; tufted (prostrate at the base,
trailing, rooting at the lower
nodes); 200-900 mm tall. Leaf
blades 20-100 mm long; 3-7 mm
wide. Spikelets 2. 5-3. 5 mm long.

Leaf blades mostly cauline,
narrowly lanceolate, soft and thin; panicle loosely
contracted; spikelets distinctly pedicellate and conspicu-
ously asymmetrical; upper glume more than five times
longer than lower glume.

Llowering mainly October to April. In damp shady
places along rivers or streams and in forest undergrowth.
Occasionally in woodlands and mopaneveld. Infrequent to
J locally common. Biome: Savanna. Tropical east Africa to

India. Natural pasture (but too slender to be productive).
Other Sacciolepis species in our area all have spikelike pan-
icles.

Description: Stapf 1920 (766), Chippindall & Crook
1976 (239), Chippindall 1955 (357), Clayton et al.
1970-1982 (455). Illustration: Chippindall 1955 (fig. 307).
Voucher: Smook 5717. PRECIS code 9901240-00400.

Sacciolepis huillensis (Rendle) Stapf

Annual swamp grass.

Short-lived perennial, or an-
nual; loosely tufted and hydro-
phyte; 100-250(-500) mm tall.

Leaf blades 15-120 mm long; 1-5
mm wide. Spikelets 1.3-1. 8 mm
long. Inflorescence spikelike;
spikelets shortly pedicellate,
laterally compressed; lower glume 1/3-2/3 the spikelet
length; upper glume more or less equalling the spikelet.

Llowering March to June. At high altitudes in sandy
soils at water edges, sometimes partly submerged in water.
Rare. Biome: Savanna. Southern tropical Africa.

Distinguished from S. chevalieri and S. typhura by its
annual habit and smaller spikelets.

Description: Stapf 1920 (755), Chippindall & Crook
1976 (238), Chippindall 1955 (358), Clayton et al.
1970-1982 (458). Voucher: Johnstone 356. PRECIS code
9901240-00700.

Sacciolepis indica (L.) A. Chase

(=S. auriculata Stapf) 2.

Annual; tufted (with few basal
leaves; culms slender, decumbent
or ascending, often with aerial
roots); 100-1000 mm tall. Leaf
blades 20-200 mm long; 1-7 mm
wide. Spikelets 2.7-3. 1 mm long.

Culms solid, 1.0-2. 5 mm in
diameter; leaf sheaths with auricles 1.5-2. 5 mm long; pani-
cle 10-130 mm long; spikelets laterally compressed; lower
glume 1.0-1. 5 mm long, 1/2 the spikelet length; upper
glume equalling the spikelet.

Llowering November to April. In sandy soils at
streamsides and in marshy places. Rare. Biome: Savanna.
Old world tropics. Closely related to S. rigens, which is a
perennial from northern Namibia with a short oblique rhi-
zome and larger spikelets.

Description: Hitchcock & Chase 1950 (688), Clayton et
al. 1970-1982 (458). Voucher: Schackleton 472. PRECIS
code 9901240-00720.

Sacciolepis interrupta (Willd.) Stapf

Perennial; hydrophyte and
rhizomatous (culms thick, spongy
and rooting at lower nodes);

300-1500 mm tall. Leaf blades
50-300 mm long; 3-17 mm wide.

Spikelets 3. 5^1. 5 mm long. Pani-
cle spikelike, 50-300 mm long,
not very dense; spikelets dorsally
compressed, acute to acuminate;
upper floret 3. 2-3. 5 mm long.

Llowering July and March. In shallow water and
swampy areas. Rare. Naturalized from India or Asia.
Biome: Savanna. India to southeast Asia. Introduced into
tropical Africa. Closely related to S. africana, which has
smaller spikelets with obtuse to subacute tips.

Description: Stapf 1920 (757), Clayton et al. 1970-1982
(456). Voucher: Cresswell 19. PRECIS code 9901240-
00730.

286

Sacciolepis rigens (Mez) A. Chev.

Perennial; loosely tufted and
rhizomatous (rhizome short and
oblique); 600-2000 mm tall. Leaf
blades 100^100 mm long; 2-6
mm wide. Spikelets 3. 0-4. 5 mm
long. Culms solid, 1 .0-2.5 mm in
diameter; leaf sheaths with
auricles 1.5-2. 5 mm long; panicle
60-200 mm long; spikelets
laterally compressed; lower glume 1 .8-2.9 mm long, 1/2 the
spikelet length; upper glume equalling the spikelet.

Flowering around January. Sandy moist soil along rivers
or streams. Rare. Biome: Savanna. Unevenly but widely
distributed throughout tropical Africa. Closely related toS.
indica, which is annual with smaller spikelets and known
in southern Africa only from Transkei.

Description: Clayton et al. 1970-1982 (460). Voucher:
De Winter & Wiss 4310. PRECIS code 9901240-00750.

Sacciolepis typhura (Stapf) Stapf

(=S. cinereo-vestita (Pilg.)

C.E. Hubb.) 2; ( =S . glaucescens
Stapf) 2.

Purple hood grass.

Robust, erect perennial; hy-
drophyte and rhizomatous (rhi-
zome creeping and branched,
culms usually spongy at the base); 500-1500 mm tall. Leaf
blades 100-350 mm long; 2-10 mm wide. Spikelets 1 .7-2.5
mm long. Leaf sheaths papery with prominent cross-veins;
panicle spikelike, 100-300 mm long, 4-7 mm wide, dense;
spikelets laterally compressed; lower glume more or less
1/2 the length of the upper glume.

Flowering December to May. Wet soils in seasonal
swamps, marshy places or floodplains; often submerged.
Locally common. Biome: Savanna and Grassland.

Throughout tropical Africa. Intergrades into S. chevalieri ,
which is a smaller plant that lacks spongy culms and has
spikelets more loosely arranged and leaf blades 1-3 mm
wide.

Description: Chippindall & Crook 1976 (240), Chippin-
dall 1955 (358), Clayton et al. 1970-1982 (460).
Illustration: Chippindall 1955 (fig. 308). Voucher: Smith
2682. PRECIS code 9901240-00800.

Fig 181. PI. 166.

Sartidia De Winter

Perennial; caespitose (densely so). Culms 800-2000 mm
high ; herbaceous; unbranched above. Leaf blades linear;
rolled. Ligule a fringed membrane, or a fringe of hairs.

Inflorescence paniculate (erect, narrow, often
interrupted ); open; espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary; 12-30 mm long ; not noticeably com-
pressed; disarticulating above the glumes. Glumes two;
more or less equal; long relative to the adjacent lemmas;
awned, or awnless; non-carinate ( rounded on the back)',
similar (narrow, nerves evanescent). All florets female-
fertile only; proximal incomplete florets absent.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes; hairless (glabrous or scabrid); without a ger-
mination flap; 3 nerved; awned (cf. Aristida). Awns triple
or trifid, commonly with a basal column (or at least with
the three spreading awns twisted together basally)', apical;
non-geniculate; hairless (glabrous or scabrid); about as long

as the body of the lemma to much longer than the body of
the lemma. Palea present; conspicuous but relatively short
(small, scale-like); 2-nerved. Lodicules 2; membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit medium sized
(8-10 mm); fusiform; hilum long-linear; pericarp fused;
embryo small (no more than 1/4 grain length).

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x = 11. Arundinoideae; Aristideae. 4 species.
South Africa. Mesophytic; glycophytic. Namibia and
Transvaal. 3 indigenous species.

References. 1. De Winter. 1965. Bothalia 8: 381.

Species treatment by G.E. Gibbs Russell.

1(0). Lemma and awns 30-40 mm long; lateral awns about
1/2 size of median awn; callus minutely bifid . . .

S. sp. (=Muller 2174)

Lemma and awns more than 50 mm long; lateral awns
about same length as median awn; callus not bifid

2

2(1). Lemmas and awns 50-60 mm long; callus obtuse . .

S. jucunda

Lemma and awns 90-120 mm long; callus acute . .
S. angolensis

Fig. 182. Sartidia angolensis

287

Sartidia angolensis (C.E. Hubb.) De Winter

Fig. 182. PI. 167.

( =Aristida angolensis C.E.

Hubb.) 1.

Perennial; rhizomatous and
tufted (erect); 1000-2000 mm
tall. Leaf blades to 350 mm long;

2—4 mm wide (narrowed below).

Spikelets 90-120 mm long (in-
cluding awns). Lateral awns
about same length as median awns; callus acute.

Flowering February to July. Grasslands on Kalahari
sand, often in depressions. Rare. Biome: Savanna. To
Angola and Zambia.

Description: De Winter 1965 (384). Illustration: De
Winter 1965 (385). Voucher: De Winter 2779. PRECIS
code 9902622-00100.

Sartidia jucunda (Schweick.) De Winter

( =Aristida jucunda
Schweick.) 1.

Tufted perennial; densely tuft-
ed and rhizomatous; to 1000 mm
tall. Leaf blades to 450 mm long;

3-4 mm wide. Spikelets 50-60
mm long (including awns). Old
leaves reddish-brown; lateral awn
about the same length as median awn; callus obtuse.

Flowering April to May. Rocky hillsides, altitudes
1300-2000 m. Rare. Biome: Savanna.

Description: De Winter 1965 (384), Chippindall 1955
(307). Illustration: De Winter 1965 (382), Chippindall 1955
(fig. 272). Voucher: Codd 8686. PRECIS code

9902622-00200.

Sartidia sp. (= Muller 2174)

Perennial; rhizomatous and
tufted (erect); to 800 mm tall.

Leaf blades to 400 mm long; 2-4
mm wide. Spikelets 30^10 mm
long. Lateral awns about 1/2 as
long as median awns; callus
minutely bifid.

Flowering March to June.

Hillslopes, probably restricted to
serpentine soil. Rare. Endemic.

Voucher: Muller 2174. PRECIS code 9902622-99999.

Schismus P. Beauv.

Electro Panz Hemisacris Steud.

Annual, or perennial (infrequently); caespitose (rarely),
or decumbent (low). Culms 30-400 mm high; herbaceous;
unbranched above. Leaf blades linear to linear-lanceolate;
0.5-2. 5 mm wide\ flat, or rolled (convolute). Ligule a fringe
of hairs.

Inflorescence paniculate ; contracted; espatheate.

Spikelet-bearing axes persistent.

Spikelets solitary; 4—8 mm long; compressed laterally
(slightly); disarticulating above the glumes, or falling with
the glumes (rarely); with conventional internode spacings.
Glumes two; more or less equal; markedly shorter than the
spikelets to about equalling the spikelets; awnless; similar
(herbaceous-membranous). Lower glume 5-7 nerved. In-
complete florets distal to the female-fertile florets, merely
underdeveloped; proximal incomplete florets absent.

Female-fertile florets 5—10. Lemmas similar in texture
to the glumes (herbaceous, the lobes and margins hyaline);
hairy; 7-9 nerved; incised (to merely emarginate); awnless,

or mucronate (from the sinus), or awned. Awns, when
present 1 , from the sinus; non-geniculate; much shorter than
the body of the lemma to about as long as the body of the
lemma. Palea present; relatively long; 2-nerved. Lodicules
2; fleshy; ciliate, or glabrous. Stamens 3. Ovary glabrous.
Hilum short; pericarp fused; embryo large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, x - 6. Arundinoideae; Danthonieae. 5 species.
Africa, Mediterranean to northwest India. Xerophytic; in
open habitats. Namibia, Botswana, Orange Free State,
Lesotho, and Cape Province. 4 indigenous species.

References. 1. Conert. & Tuerpe. 1974. Abhant. Senck.
Naturf. Gesell. 32: 532.

Species treatment by N.P. Barker.

1(0). Central awn or mucro of lemmas 1.0-1. 5 mm long;
lemma backs glabrous, lower third of margin

fringed with hairs S. pleuropogon

Central awn or mucro of lemma shorter than 1 mm
or absent; lemma backs and/or margins pubescent

2

2(1). Plants annual; spikelets to 1.5 mm wide, lanceolate,
often pale green;hairs on lemmas often club-shaped

S. barbatus

Plants perennial; spikelets 1. 5-4.0 mm wide, ovate,
often with a purple colouration; hairs on lemmas

never club-shaped 3

3(2). Lemmas densely pubescent, hairs 1.0-1. 5 mm long,
with a short (less than 1 mm long) mucro arising

from the sinus between the lemma lobes

S. inermis

Lemmas sparsely pubescent, hairs short (less than 1
mm long), often arranged only as a row of hairs
across the back, mucro very short or absent ....
S. scaberrimus

Schismus barbatus (Loefl. ex L.) Thell.

Fig. 183. PI. 168.

Haasgras.

Annual; tufted; 50-250 mm
tall. Leaf blades 10-50 mm long
(rarely longer); involute, to 1.5
mm wide. Spikelets 4—7 mm
long; 1.5 mm wide. Panicle
10-50 mm long; spikelets 5-10-
flowered, usually light green,

sometimes purple, often long and narrowly lanceolate;
glumes usually closed; lemma backs pubescent, hairs usual-
ly club-shaped, lobes obtuse, with or without a short (less
than 1 mm long) mucro arising from the sinus.

Flowering June to December. Alluvial soils, disturbed
sandy areas. Common. Biome: Fynbos, Savanna, Grass-
land, Nama-Karoo, and Succulent Karoo. North Africa,
Middle East and SW Asia. Natural pasture (for sheep), or

weed (in gardens).

Description: Conert & Tuerpe 1974 (532), Stapf
1898-1900 (693), Chippindall 1955 (240). Illustration:
Conert & Tuerpe 1974 (fig. 6). Voucher: Oliver, Toelken&
Venter 367. PRECIS code 9904050—00100.

Schismus inermis (Stapf) C.E. Hubb.

Perennial; tufted; 120-400
mm tall. Leaf blades to 300 mm
long; about 1 mm wide. Spikelets
4. 5-7.0 mm long; 2.5^4.0 mm
wide. Panicle 25-70 mm long,
contracted and dense; spikelets
green or purple, 4-6-flowered,
ovate; glumes often open; lem-
mas pubesent, hairs 1.0-1. 5 mm

288

Fig. 183. Schismus barbatus

long, never club-shaped, lemma apex minutely lobed with
a short mucro (less than 1 mm long), arising from the sinus
between the lobes and seldom extending much beyond the
lobes.

Flowering June to February. On dense grassy slopes and
rocky areas. Common. Biome: Fynbos, Nama-Karoo, and
Succulent Karoo. Endemic. May be confused with Koeleria
capensis, which has a membranous ligule.

Description: Conert & Tuerpe 1974 (532), Stapf
1898-1900 (694), Chippindall 1955 (240). Illustration:
Conert & Tuerpe 1974 (fig. 12; spikelet only). Voucher:
Archibald 4541/41. PRECIS code 9904050-00200.

Schismus pleuropogon Stapf

Perennial; stoloniferous;

120-250 mm tall. Leaf blades
30-80 mm long; involute, to 1.5
mm wide. Spikelets 5-7 mm
long; to 2 mm wide. Panicle
15-80 mm long; spikelets 5-7-
flowered, ovate; glumes usually
open; lemma backs glabrous but
lower 1/3 of the margin is fringed
with hairs, lobes short, with a straight awn, 1.0-1. 5 mm
long, arising from between the lobes.

Flowering November. Moist areas. Rare. Biome: Fyn-
bos. Endemic. There are no specimens of this taxon in PRE.
The map locality was obtained from the type locality in the
original description.

Description: Conert & Tuerpe 1974 (532), Stapf 1916
Kew Bull. (234). Illustration: Conert & Tuerpe 1974 (fig.
17). PRECIS code 9904050-00300.

Schismus scaberrimus Nees

Perennial; tufted; 100^450
mm tall. Leaf blades 30-200 mm
long; to 1.5 mm wide (somewhat
scabrid). Spikelets 5-7 mm long;

1.5-2. 5 mm wide. Panicle 10-50
mm long; spikelets 4— 6-flowered,
ovate to broadly ovate; glumes
usually quite wide open; lemmas
sparsely pubescent with short,
scattered hairs across the back, margins tufted to densely
hairy, lemma lobes tend to be united with almost no sinus,
lacerate-tipped, occasionally with a short (less than 1 mm
long) mucro extending beyond the lobes.

Flowering September and October. Sandy areas such as
dry river beds. Infrequent. Biome: Fynbos and Succulent
Karoo. Endemic.

Description: Conert & Tuerpe 1974 (532), Stapf
1898-1900 (695). Illustration: Conert & Tuerpe 1974 (fig.
15). Voucher: De Winter and Verdoorn 9038. PRECIS code
9904050-00400.

Schizachyrium Nees

Pithecurus Kunth, Schizopogon Spreng.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms (50-)300-3200
mm high; herbaceous; branched above. Leaf blades linear.
Ligule a fringed membrane (short). Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant ; overtly heteromorphic (the pedicellate
broader and flatter or reduced, the G1 sometimes awned);
all in heterogamous combinations.

Inflorescence a single raceme, or paniculate (of single
racemes, sometimes solitary but usually in a spatheate false
panicle)', spatheate', a complex of ‘partial inflorescences’
and intervening foliar organs. Spikelet-bearing axes
peduncled ‘racemes' ; solitary (in their spathes, hut often
fascicled ); with substantial rachides\ disarticulating at the
joints.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, or sterile. Female-fertile spikelets
compressed dorsiventrally (or subterete below); falling with
the glumes (and the joint). Glumes present; two; more or
less equal; awnless; very dissimilar (lower bicarinate, upper
thinner and naviculate). Proximal incomplete florets /;
epaleate; sterile.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline, often stipitiform); incised (or rarely
merely prolonged into the awn, without teeth); awned.
Awns 1 ; median; from the sinus, or apical; geniculate; much
longer than the body of the lemma. Palea present, or absent;
when present very reduced (a minute, hyaline scale —
usually absent). Lodicules 2; fleshy; ciliate, or glabrous.
Stamens 2-3. Ovary glabrous. Hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. About 60 species.
Tropical. Helophytic, or mesophytic, or xerophytic; in open
habitats (savanna, rarely beaches or dunes); maritime-
arenicolous, or glycophytic. Namibia, Botswana, Transvaal,
Swaziland, Natal, and Cape Province. 6 indigenous species.

289

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Plant annual, base delicate 2

Plant perennial, tufted, base stout 3

2(1). Lower leaves with blade tips rounded; inflorescence
exserted from spathe; sessile spikelets 2. 5-3.0 mm

long S. brevifolium

All leaves with blade tips tapering; inflorescence
partially included in spathe; sessile spikelets 5-6

mm long S. exile

3(1). Racemes appearing nearly glabrous, hairs present
only along margins of rachis and pedicels; sessile
spikelets laterally compressed between rachis

intemode and pedicel; plants turning red

S. sanguineum

Racemes appearing hairy; sessile spikelets dorsally

compressed; plants not red 4

4(3). Raceme hairs yellowish; leaf blades hairy, folded,
curved; basal sheaths compressed, fanlike, yellow

turning brown S. ursulus

Raceme hairs white (rarely cream-coloured); leaf
blades glabrous, expanded; basal sheaths not

fanlike, not yellow 5

5(4). No barren branches present below the flowering
branches; female-fertile (upper) lemma only shortly

bifid; Namibia, Botswana, Transvaal

S. jeffreysii

Many barren branches present below the flowering
branches; female-fertile (upper) lemma bifid for
1/4- 1/3 of its length; Natal S. rupestre

Schizachyrium brevifolium (Swartz) Buese

Delicate annual; 50-600 mm
tall. Leaf blades to 70 mm long;

2-5 mm wide. Spikelets (sessile)

2. 5-3.0 mm long (pedicellate
reduced to a glume). Lower
leaves with rounded blade tips;
inflorescence exserted from
spathe.

Flowering February to April.

Usually grows in open, damp places such as vleis, often
shaded by taller grasses. Infrequent. Savanna. Throughout
the tropics.

Description: Chippindall 1955 (504). Voucher:

Scheepers 933. PRECIS code 9900680- 00100.

Schizachyrium exile (Hochst.) Pilg.

(=S. inclusum Stent) 2.

Annual; tufted; 500-1000 mm
tall. Leaf blades 20-150 mm
long; 2-3 mm wide. Spikelets
(sessile) 5-6 mm long (pedicel-
late much smaller). Leaf blades
with tapering tips; inflorescence
partly enveloped by spathe.

Flowering March to June. Open places, often in poor dry
soil. Infrequent. Biome: Savanna. Through tropical Africa
to Asia.

Description: Chippindall 1955 (504), Clayton et al.
1970-1982 (756). Voucher: Volk 363. PRECIS code
9900680-00200.

Schizachyrium jeffreysii (Hack.) Stapf

Perennial; loosely tufted;

600-1000 mm tall. Leaf blades to
200 mm long; 2-5 mm wide.

Spikelets (sessile) 7-8 mm long
(dorsally compressed; pedicellate
shorter). Leaves glabrous; inflor-
escence lacking barren branches;
racemes with conspicuous white
hairs; female-fertile lemma only
shortly bifid.

Flowering February to June. Open veld. Common.
Biome: Savanna. Southern tropical Africa. The inflores-
cence of plants with a single raceme may resemble
Elionurus muticus , which has leaf blades narrower than 2
mm and is densely tufted.

Description: Chippindall 1955 (503). Voucher: Giess
9926. PRECIS code 9900680-00300.

Schizachyrium rupestre (K. Schum.) Stapf

Perennial; tufted; 300-1500
mm tall. Leaf blades 150-300
mm long; 1-5 mm wide. Spike-
lets (sessile) 4.0-6. 5 mm long
(dorsally compressed; pedicellate
nearly as long). Inflorescence
with many barren branches;
racemes with conspicuous white
hairs; female-fertile lemma bifid
for 1/4-1/3 of its length.

Flowering March. Moist places in coastal bush. Rare.
North to Senegal, Nigeria and Tanzania. Known here only
by two collections of R.P. Ellis from St. Lucia, far south
of its previously reported range.

Description: Clayton et al. 1970—1982 (758). Voucher:
Ellis 4497. PRECIS code 9900680-00350.

290

Schizachyrium sanguineum (Retz.) Alst.

Fig. 184. PI. 169.

(=5. semiberbe Nees) 2.

Rooidekgras, red autumn
grass.

Perennial; shortly creeping
rhizomatous and tufted;

400-1200 mm tall. Leaf blades
60-300 mm long; to 7 mm wide.

Spikelets (sessile) 6-9 mm long (laterally compressed; ped-
icellate somewhat shorter). Plants conspicuously red in
autumn; ligule undivided, strongly curved, blade tips
rounded or abruptly pointed; racemes nearly glabrous.

Flowering January to May. Open veld. Very common.
Biome: Savanna and Grassland. Throughout tropics. Do-
mestic use (thatching). Vegetatively similar to and often
occurring with Heteropogon contortus, which has a slightly
curved ligule, and forms of Themeda triandra , which has
a divided ligule and tapering leaf tips.

Description: Chippindall 1955 (502), Clayton et al.
1970-1982 (756). Illustration: Chippindall 1955 (pi. 22),
Clayton etal. 1970-1982 (fig. 1 78). Voucher: Giess 10072.
PRECIS code 9900680-00400.

Schizachyrium ursulus Stapf

Perennial; tufted (in dense
round tufts); 300-700 mm tall.

Leaf blades to 400 mm long; 2-3
mm wide. Spikelets (sessile) 7-8
mm long (dorsally compressed;
pedicellate shorter). Basal
sheaths yellow, compressed,
fanlike, blades hairy, folded,
curved; racemes with conspicu-
ous yellowish hairs.

Flowering January to April. Open sour veld. Infrequent.
Biome: Savanna and Grassland. Southern tropical Africa.

Description: Chippindall 1955 (503). Voucher: De
Winter 273. PRECIS code 9900680-00500.

Schmidtia Steud.

Antoschmidtia Boiss.

Annual, or perennial (usually viscid); caespitose to
decumbent. Culms 150-1000 mm high; herbaceous;
branched above. Leaf blades linear to linear-lanceolate; flat,
or rolled. Ligule a fringe of hairs.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets solitary; 7-10 mm long; compressed laterally
(slightly so); disarticulating above the glumes; not disartic-
ulating between the florets. Hairy callus present. Glumes
two; very unequal to more or less equal; about equalling the
spikelets; awnless; similar (lanceolate, membranous,
usually green or grey). Incomplete florets distal to the
female-fertile florets, 1-2, merely underdeveloped, awned;
proximal incomplete florets absent.

Female-fertile florets 3-9. Lemmas decidedly firmer
than the glumes (subcoriaceous); without a germination
flap; 9 nerved; incised (6 lobed); awned. Awns 5 (one from
each sinus)-, median and lateral. The median awn similar in
form to the laterals; non-geniculate; about as long as the
body of the lemma. Palea present; relatively long (longer
than the body of the lemma). Lodicules 2; fleshy; ciliate
(sometimes glandular), or glabrous. Stamens 3. Ovary
glabrous. Fruit small (about 2.5 mm long); ellipsoid; hilum
short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines uneven. PCR sheath

extensions absent. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Chloridoideae; Pappophoreae. 2 species.
Tropical and southern Africa, Cape Verde Is., Pakistan.
Xerophytic; in open habitats; glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
and Cape Province. 2 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1965. Bol. Soc. Brot. 39: 303-31 1. 3. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Plant perennial, with a woody rootstock; culm bases
swollen, clad with white-hairy cataphylls; leaf

blades usually less than 7 mm wide

S. pappophoroides

Plant annual, with fibrous roots; culm bases not
swollen, not clad with hairy cataphylls; leaf blades
usually more than 7 mm wide . . S. kalihariensis

291

Schmidtia kalihariensis Stent

Annual; tufted; to 1000 mm
tall. Leaf blades 70-150 mm
long; 8-10 mm wide. Spikelets
6-17 mm long. Plant coarse,
hairy, viscid; culm bases not
swollen, lacking cataphylls; leaf
blades tapering abruptly at tip.

Flowering throughout the year
(but most commonly in mid to
late summer). Open veld, usually in poor sandy soils. Com-
mon, or locally dominant. Biome: Savanna, Nama-Karoo,
Succulent Karoo, and Desert. To Chad and Sudan. Hay and
pasture, has a strong unpleasant smell.

Description: Chippindall 1955 (234). Illustration: Chip-
pindall 1955 (fig. 207). Voucher: Theron 1985. PRECIS
code 9903610-00100.

Schmidtia pappophoroides Steud.

(=S. bulbosa Stapf) 2.

Kalahari sandkweek, vaalgras.

Perennial; stoloniferous and
tufted; 150-900 mm tall. Leaf
blades 50-160 mm long; 2-7 mm
wide. Spikelets 8-15 mm long.

Plant hairy to nearly glabrous; culm bases swollen, clad by
hairy cataphylls; leaf blades tapering gradually to a long
fine point.

Flowering throughout the year (but most commonly in
summer). Open veld in a variety of soils and habitats. Com-
mon, or locally dominant. Savanna, Nama-Karoo,
Succulent Karoo and Desert. To eastern and central tropical
Africa and in Cape Verde Islands. Variable in size,
hairiness and awn length.

Description: Chippindall 1955 (232), Clayton et al.
1970-1982 (165). Illustration: Chippindall 1955 (fig. 206),
Clayton et al. 1970-1982 (fig. 54). Voucher: Werdermann
& Oberdieck 2369. PRECIS code 9903610-00200.

Schoenefeldia Kunth

Annual, or perennial; caespitose. Culms 700-1200 mm
high; herbaceous. Leaf blades linear. Ligule a fringed
membrane (short).

Inflorescence of spike-like main branches ( usually 2-6
sessile, flexuous spikes)-, digitate or subdigitate; espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary; biseriate; all sessile-, strongly com-
pressed laterally, disarticulating above the glumes. Hairy
callus present. Glumes two; very unequal; long relative to
the adjacent lemmas (exceeding the lemma); awned (Gl,
sometimes), or awnless; similar (persistent, narrow or
setaceous, subhyaline). All florets female-fertile, or a
solitary distal incomplete floret present; proximal incom-
plete florets absent.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (often blackened at maturity); 3 nerved; incised;
awned. Awns 1; median; from the sinus; non-geniculate, or
geniculate; much longer than the body of the lemma (awns
very long, flexuous, tangling one another). Palea present.
Lodicules 2; fleshy; glabrous. Stamens 2-3. Ovary
glabrous. Fruit ellipsoid; hilum short; pericarp free; embryo
large.

Photosynthetic pathway and related features. C4;
XyMS+.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Tropical Africa, Asia. In
open habitats (savanna, hardpans and seasonally flooded
flats). Transvaal. 1 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Schoenefeldia transiens (Pilg.) Chiov.

Fig. 186. PI. 171.

{=Chloris transiensis

Pilg.) I-

Perennial; densely tufted;

700-1200 mm tall. Leaf blades to
350 mm long; 5 mm wide. Spike-
lets 3. 5-5.0 mm long. Spikelets
small in comparison to the awns,
awns of the female-fertile and
sterile lemmas 10-25 mm and 25-45 mm long respectively,
curving gracefully around the spike.

Flowering January to February. Heavy soils and
seasonally flooded flats. Rare. Biome: Savanna. Tropical
Africa. Reported to be cleistogamous.

Description: Clayton et al. 1970-1982 (309).

Illustration: Clayton et al. 1970-1982 (fig. 86). Voucher:
Gertenbach 4931. PRECIS code 9902950-00100.

Fig. 186. Schoenefeldia transiens

292

Secale L.

Annual (rarely perennial); caespitose (or solitary culms).
Culms 200-1500 mm high; herbaceous; unbranched above.
Leaves auriculate. Leaf blades linear; flat, or rolled
(convolute). Ligule an unfringed membrane.

Inflorescence a single spike (laterally compressed,
distichous)-, espatheate. Spikelet-bearing axes disarticu-
lating, or persistent (in cultivated forms); disarticulating at
the joints.

Spikelets solitary; distichous; 10-18 mm long; com-
pressed laterally; falling with the glumes (and the joint), or
not disarticulating (in cultivated forms). Glumes two; very
unequal, or more or less equal; decidedly shorter than the
adjacent lemmas; awned; similar (subulate). Upper glume

Fig. 187. Secale africanum

l nerved. All florets female-fertile, or distal incomplete
florets also present, merely underdeveloped (a single
rudiment); proximal incomplete florets absent.

Female-fertile florets 2-3. Lemmas less firm than the
glumes, or similar in texture to the glumes; 5 nerved; entire;
awned. Awns 1; median; apical; non-geniculate; much
longer than the body of the lemma. Palea present; relatively
long. Lodicules 2; membranous; ciliate. Stamens 3. Ovary
hairy. Fruit medium sized, or large; hilum long-linear;
embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 1. Pooideae; Triticodae; Triticeae. 5 species.
Mediterranean, eastern Europe to central Asia, and South
Africa. Mesophytic, or xerophytic; in open habitats (sandy
soils and dry hillsides); glycophytic. Cape Province. 1
indigenous species.

Intergeneric hybrids with Triticum (X Triticosecale
Wittmack ), Agropyron, Aegilops (X Aegi/oseca/e Ciferri &
Giacom.), Elytrigia. X Agrotrisecale Ciferri & Giacom. =
Agropyron x Secale x Triticum.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

Secale africanum Stapf

Fig. 187. Pi. 172.

Wild rye, wilderog.

Perennial; loosely tufted; to
1000 mm tall. Leaf blades
200-350 mm long; 4—9 mm wide.

Spikelets 10-15 mm long
(excluding awns). Spike 80-120
mm long, linear, very dense,
rachis fringed with short hairs,
breaking up at maturity; lemma keel minutely hairy, awn
up to 20 mm long, surface scabrid.

Flowering December. Undisturbed places on riverbanks.
Rare. Biome: Nama-Karoo. Endemic. Potential pasture
(liked by birds and stock). Reported to have occurred
abundantly during earlier years, but now apparently
restricted to rare patches on a farm Voelfontein in the
Sutherland district. Distinguished from cultivated rye, S.
cereale, which is annual, has the lemma keel fringed with
stiff hairs and the surface smooth, awns up to 50 mm long
and a rachis that does not break up at maturity.

Description: Stapf 1898-1900 (764), Chippindall 1955
(70). Illustration: Chippindall 1955 (fig. 43). Voucher:
Schweickerdt 1927. PRECIS code 9904390-00100.

Sehima Forssk.

Hologamium Nees.

Annual, or perennial; caespitose. Culms 200-1000 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear. Ligule a fringed membrane. Plants
bisexual, with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant ; overtly heteromorphic; all
in heterogamous combinations.

Inflorescence a single raceme (a single, curved, culm-
like ‘raceme’ with embedded spikelets)-, espatheate', not
comprising ‘partial inflorescences’ and foliar organs.
Spikelet-bearing axes ‘racemes’ (spiciform, laterally
compressed, curved); solitary; with substantial rachides
(compressed); disarticulating at the joints.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, or sterile, flat, often with G1 large and
strongly nerved, lemmas awnless. Female-fertile spikelets

293

compressed laterally (usually, more or less); falling with the
glumes. Glumes two; more or less equal; awned (G2 with
an apical bristle-like awn, G1 2-dentate or 2-mucronate);
very dissimilar (lower 2-keeled and 2-winged, upper
naviculate-subulate). Proximal incomplete florets I;
paleate, palea fully developed; male.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); incised; awned. Awns 1; median; from
the sinus; geniculate; much longer than the body of the
lemma. Palea present; relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Hilum short; embryo
large.

Cytology, classification, distribution. Chromosome base
number, a: = 10, 17, and 20. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 5 species. Warm Africa,
India, Australia. Helophytic to mesophytic; in open habitats
(savanna, sometimes on heavy clay); glycophytic. Namibia,
Botswana, Transvaal, Swaziland, and Natal. 2 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by G.E. Gibbs Russell.

Fig. 188. Sehima galpinii

1(0). Annual; lower glume of sessile spikelets deeply
grooved in lower half, tip membranous, deeply 2-

toothed S. ischaemoides

Perennial, densely tufted; lower glume of sessile
spikelets flat or slightly convex, tip not
membranous, not toothed

Sehima galpinii Stent

Dekgras.

Robust perennial; tufted; to
1800 mm tall. Leaf blades 3-6
mm wide. Spikelets (sessile)

12-15 mm long (pedicellate
somewhat shorter). Lower glume
of sessile spikelets flattish. tip not
toothed.

Flowering October to April. Black turf soil. Infrequent.
Biome: Savanna. Southern tropical Africa. Domestic use
(thatching). Can be distinguished from other robust grasses
with solitary racemes, such as Urelytrum agropyroides and
Trachypogon spicatus, by its ligule which is a fringe of
hairs.

Description: Chippindall 1955 (489). Illustration: Chip-
pindall 1955 (pi. 1 7). Voucher: Galpin M557. PRECIS code
9900130-00100.

Sehima ischaemoides Forssk.

Annual; 200-600 mm tall.

Leaf blades 50-300 mm long; 1-3
mm wide. Spikelets (sessile)

9-15 mm long. Lower glume of
sessile spikelets deeply grooved
below, tip deeply 2-toothed.

Flowering February to May.

Dry soils in savanna. Conserva-
tion status not known. Biome: Sa-
vanna. Tropical Africa to Pakistan.

Description: Clayton et al. 1970-1982 (750). Voucher:
De Winter 2932. PRECIS code 9900130-00200.

S. galpinii

Fig. 188. PI. 173.

Setaria P. Beauv.

Acrochaete Peter, Chaetochloa Scribn., Cymbosetaria
Schweick., Miliastrum Fabric., Tansaniochloa Rauschert.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 100-3200 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades occasionally sagittate or hastate — then
perhaps referable to Cymbosetaria ; flat, or folded; pseudo-
petiolate (occasionally), or not pseudopetiolate. Ligule a
fringed membrane (narrow), or a fringe of hairs. Plants
with hermaphrodite florets. The spikelets of sexually
distinct forms on the same plant (when clustered, often not
all fully developed), or all alike in sexuality.

Inflorescence of spike-like main branches ( not
uncommonly so — e.g., in Sect. Ptychophyllum — though
this is ignored in many published keys), or a false spike,
with clusters of spikelets on reduced axes, or paniculate ;
open, or contracted; axes not ending in spikelets ( produced
into 'bristles’ beyond the spikelets)-, espatheate. Spikelet-
bearing axes persistent.

Spikelets with ‘involucres’ of ‘bristles’ , or (at least some
of them) subtended by solitary ‘bristles’ (e.g., in Sect.
Ptychophyllum)-, not in distinct ‘long-and-short’ combina-
tions. Female-fertile spikelets 2-4 mm long; compressed
dorsiventrally; falling with the glumes, or not disarticu-
lating (in cultivated forms). Glumes two; relatively large;
very unequal; awnless; membranous. Proximal incomplete

294

florets 7; paleate, or epaleate, palea fully developed to
reduced; male, or sterile.

Female-fertile florets 7. Lemmas decidedly firmer than
the glumes (crustaceous); rugose; becoming indurated;
hairless; usually non-carinate (but cymbiform in species
perhaps referable to Cymbosetaria)', having the margins
tucked in onto the palea; with a clear germination flap; 1-5
nerved; entire; awnless (usually apiculate). Palea present;
relatively long. Lodicules 2; fleshy; glabrous. Stamens 3.
Ovary glabrous. Fruit small, ellipsoid to subglobose; hilum
short; embryo large.

Photosynthetic pathway. C4; NADP-ME (5 species);
XyMS- PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9 and 10. Panicoideae; Panicodae; Paniceae.
About 110 species. Tropical and warm temperate.
Generally mesophytic; in shade (e.g. S. palmifolia ), or in
open habitats (woodland, grassland, weedy places).
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. 19 indige-
nous species, 2 naturalized species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by M. Koekemoer.

1(0). Lower glume 1 -nerved 2

Lower glume 3-nerved 3

2(1). Bristles antrorsely barbed, with sparse long white

hairs; plants perennial S. rigida

Bristles retrorsely barbed, without hairs; plants annual

S. verticillata

3( 1). Culm nodes sparsely or densely pubescent 4

Culm nodes glabrous 7

4(3). Leaf blades saggitate at the base; panicle open ....

S. appendiculata

Leaf blades linear at the base; panicle spike-like or

open 5

5(4). Panicle open, with branches spreading or sub-erect;
leaf blades lanceolate, plicate at first and then flat;
spikelets with a solitary bristle . . . S. homonyma
Panicle densely spike-like; leaf blades linear, flat or

folded; spikelets with 4-8 bristles 6

6(5). Spikelets 2.5— 3.0(— 3.7) mm long; basal plant parts
with a light reddish or yellowish colour; rhizomes
not robust, mostly oblique; panicle often
interrupted at the base, tapering at the apex ....

S. incrassata

Spikelets 3. 5-5.0 mm long; basal plant parts dark
coloured; rhizomes very robust and branches far
apart; panicle usually dense, cylindrical to the apex

S. nigrirostris

7(3). Leaf blades pseudopetiolate at the base

S. sagittifolia

Leaf blades linear or tapering at the base 8

8(7). Plants annual 9

Plants perennial 12

9(8). Spikelets disarticulating above the glumes; lower
lemma smooth; panicle spike-like, 8-24 mm wide

S. italica

Spikelets disarticulating below the glumes; lower
lemma rugose; panicle spike-like or open, to 15 mm
wide 10

10(9). Panicle open and lax; spikelet tips mucronate and
deflexed; bristles delicate and solitary,

terminating the inflorescence branches

S. finita

Panicle cylindrical and spike-like; spikelet tips
acute and not deflexed; bristles rigid, in clusters

of 4-10 from the spikelet bases 11

11(10). Upper lemma coarsely rugose; bristles at least two
times the spikelet length, yellowish, brownish or
copper coloured; panicle usually ovate, to 5 times
longer than wide; growing under trees or bushes

in drier bushveld areas S. ustilata

Upper lemma finely rugose; bristles usually one and
a half times the spikelet length, mostly bright
yellow or purple-brown; panicle usually linear
and slender, about 10 times longer than wide;
common weed in moist disturbed areas and

cultivated lands S. pallide-fusca

12(8). Panicle open, usually lax; bristles solitary, delicate

13

Panicle cylindrical, spike-like; bristles in clusters of

295

6-10 (or in S. obscura solitary and very sparse)

16

13(12). Leaf blades flat, 2-6 mm wide . S. pseudaristata
Leaf blades plicate (sometimes only visible at the
base of young leaves), 5-1 10 mm wide .... 14
14(13). Plants robust, 900-3000 mm tall; culms 4-10 mm
in diameter; leaves 30-110 mm wide; leaf

sheaths usually densely pubescent

S. megaphylla

Plants not robust, 400-1500 mm tall; culms 2— 3(— 5)
mm in diameter; leaves 4-25(-30) mm wide; leaf

sheaths usually glabrous 15

15(14). Leaf blades narrowly lanceolate; 5-35 mm wide,
coarsely plicate, flat; plants loosely tufted; upper

lemma smooth or obscurely rugose

S. plicatilis

Leaf blades linear, 1 .5— 7.0(— 13.0) mm wide, finely
plicate, flat or involute; plants densely tufted;

upper lemma rugose S. lindenbergiana

16(12). Bristles solitary and sparse; upper lemma smooth
and deeply grooved; spikelets 4. 0-4. 6 mm long;
plants of the Natal Drakensberg ... S. obscura
Bristles in clusters of 6-10; upper lemma rugose
and rounded on the back; spikelets less than 4
mm long; very widespread distribution .... 17
17(16). Rhizome bare, very slender, knotty, branched;

culms wiry S. genicuiata

Rhizome covered with basal sheaths, thick, oblique
or creeping, not branching; culms not wiry . 18
18(17). Plants very robust, to 3000 mm tall, occasionally
with stilt roots; culms 6-10 mm in diameter; leaf
blades 8-17 mm wide; panicle 150-300 mm long

S. sphacelata var. splendida

Plants slender to fairly robust, 300-2000 mm tall,
lacking stilt roots; culms 1-6 mm in diameter;
leaf blades 2-10 mm wide; panicle 25-250 mm

long 19

19(18). Bristles mostly dark purple-brown or darkening
only towards the tips, occasionally yellowish;
leaves mostly basal, not more than 2 mm wide,
usually folded or inrolled, old leaves curly;

panicle 25 — 45 mm long

S. sphacelata var. torta

Bristles golden-yellow; leaves basal or cauline, 2-7
mm wide, usually flat; old leaves not curly;

panicle 30-250 mm long 20

20(19). Culms 4-10-noded, 3-6 mm in diameter, to 2000
mm tall; leaf blades 3-10 mm wide, often folded;

panicle 70-250 mm long

S. sphacelata var. sericea

Culms 2^4-noded, 1-3 mm in diameter, to 1000
mm tall; leaf blades 2-5 mm wide, flat; panicle

30-150 mm long

S. sphacelata var. sphacelata

Setaria appendiculata (Hack.) Stapf

Perennial; loosely tufted and
rhizomatous (rhizome oblique);

500-1000 mm tall. Leaf blades
70-300 mm long; 3-12 mm wide.

Spikelets 2. 0-2. 7 mm long. Culm
nodes hairy; leaf blades sagittate
at the base; panicle open, linear to
lanceolate; bristles solitary; lower
glume 3-nerved.

Flowering January to May. Rocky outcrops, among
bushes and in dry riverbeds, often in shady places. Locally
common (drier western parts). Biome: Savanna, Nama-
Karoo, and Desert. ?Endemic. Distinguished from other
hairy-noded Setaria species by the sagittate leaf bases. S.
sagittifolia also has sagittate leaf blades, but it has glabrous
culm nodes and pseudopetiolate leaf blades.

Description: Launert 1970 (160:171), Stapf 1930 (833),
Stapf 1898-1900 (422), Chippindall 1955 (343), Clayton et
al. 1970-1982 (532). Illustration: Chippindall 1955 (fig.
298). Voucher: Giess 8477. PRECIS code 9901280-00200.

Setaria finita Launert

Annual; loosely tufted (culms
erect or geniculate and rooting at
the nodes); 350-1000 mm tall.

Leaf blades 100-350 mm long;

4-12 mm wide. Spikelets 3. 2-3. 5
mm long. Culm nodes glabrous;
panicle open, up to 250 mm long;
branches delicate and lax; spike-
let tips mucronate and deflected
outwards; bristles solitary and delicate; lower glume 3-
nerved; lemmas finely rugose.

Flowering January to March. Mostly in the shade along
rivers, occasionally in disturbed places. Rare. Biome: Sa-
vanna. ?Endemic. An open, lax panicle, mucronate spike-
lets and delicate, solitary bristles distinguish this species
from S. italica , S. ustilata and S. pallide-fusca , which are
annual and have glabrous culm nodes.

Description: Launert 1970 (160:171). Voucher: Giess
7771, Smook 5076. PRECIS code 9901280-00600.

Setaria genicuiata (Lam.) Beauv.

Knotroot, bristle grass.

Perennial; (rhizome knotty,
slender and branching profusely);

300-800 mm tall. Leaf blades
1 50-300 mm long; 3^t mm wide.

Spikelets 2. 2-2. 7 mm long. Culm
nodes glabrous; panicle cylindri-
cal, spike-like; spikelets subtend-
ed by 2-3 bristles; bristles yellowish; lower glume 3-
nerved; lower lemma rugose.

Flowering December, January, and June. Mostly in
cultivated lands, occasionally adventive in disturbed areas.
Rare. Naturalized from tropical America. Biome: Fynbos
(urban areas). Tropical and temperate America. The panicle
is similar to that of S. pallide-fusca , which is annual.

Description: Hitchcock & Chase 1950 (697), Chippin-
dall 1955 (355). Illustration: Hitchcock & Chase 1950 (fig.
1564). Voucher: Taylor 7626. PRECIS code 9901280-
00800.

Setaria homonyma (Steud.) Chiov.

Fan-leaved bristle grass.

Annual; loosely tufted (culms
erect or geniculately ascending
and rooting at the nodes);

250-1000 mm tall. Leaf blades
45-300 mm long; 5-35 mm wide.

Spikelets 2. 3-2. 8 mm long. Culm
nodes pubescent; leaf blades
lanceolate, plicate at first; panicle open with stiff, spreading
or suberect branches; bristles solitary; lower glume 3-
nerved.

Flowering February to June. Shady places in woodlands,
forests, on riverbanks and floodplains on moist sandy soils,
often in disturbed areas and cultivation. Infrequent. Biome:
Savanna. Northwards to Cameroun and Ethiopia, also in
India. Natural pasture (average forage value), or weed (in
disturbed areas). Distinguished from other hairy-noded
Setaria species by plicate, lanceolate leaf blades.

Description: Launert 1970 (160:171), Stapf 1930 ; 857),
Chippindall & Crook 1976 (86), Chippindall 1955 (343),
Clayton et al. 1970-1982 (536). Voucher: Smook 1984.
PRECIS code 9901280-01000.

296

Setaria incrassata (Hochst.) Hack.

(=S. eylesii Stapf & C.E.

Hubb.) 2; ( =S . gerrardii Stapf)

2; (=S. holstii Herr.) 2; (=S.
pabularis Stapf) 2; (=S.
palustris Stapf) 2; (=S.
perberbis De Wit) 2; (=S.
phragmitoides Stapf) 2; (=S.
porphyrantha Stapf) 2; (=5.
rudifolia Stapf) 2; (=S. woodii
Hack, subsp. bechuanica De Wit) 2; (=S. woodii Hack,
var . fonssalutis De Wit) 2; (=S. woodii Hack. var. woodii)

2.

Perennial; rhizomatous and tufted (with an oblique
creeping rhizome); 300-2000 mm tall. Leaf blades 150—600
mm long; 3-14 mm wide. Spikelets 2.5-3.0(-3.7) mm long.
Basal parts showing a reddish straw colour; culm nodes
hairy; panicle spike-like, tapering towards the tip, often
interrupted in the lower part; lower glume 3-nerved.

Flowering October to May. Mostly on black clay in
moist areas such as swamps and vleis, but also on
streambanks, forest margins and rocky hillsides. Common.
Biome: Fynbos, Savanna, Grassland, and Nama-Karoo.
Tropical Africa. Very similar to S. nigrirostris, which has
its rhizome well developed and much branched, basal parts
dark coloured and spikelets 3. 5-5.0 mm long.

Description: Stapf 1930 (790), Stapf 1898-1900 (424),
Chippindall 1955 (346), Clayton et al. 1970-1982 (525).
Voucher: Smook & Gibbs Russell 2162. PRECIS code
9901280-01050.

Setaria italica (L.) Beauv.

Foxtail millet.

Annual; culms erect, solitary
to densely tufted; 350-1500 mm
tall. Leaf blades 150-450 mm
long; 6-20 mm wide. Spikelets
2. 0-3. 5 mm long. Culm nodes
glabrous; leaves mainly cauline;
panicle spike-like, 8-24 mm
wide; spikelets disarticulating above the glumes; lower
glume 3-nerved; lower lemma smooth.

Flowering January to April. In gardens or cultivation
and other disturbed areas. Rare (in the natural state). Nat-
uralized from Asia. Biome: Savanna. Tropical regions
worldwide. Cultivated mainly for bird seed, but not
extensively in South Africa. Uniquely distinguished in
Setaria by spikelets that disarticulate above the glumes.

Description: Stapf 1930 (820), Stapf 1898-1900 (428),
Chippindall 1955 (353), Clayton et al. 1970-1982 (520 &
524). Illustration: Hitchcock & Chase 1950 (fig. 1580).
Voucher: Van der Schijff PRE33148. PRECIS code
9901280-01100.

Setaria lindenbergiana (Nees) Stapf

Fig. 190. PI. 174.

(=S. phillipsii DeWet) 2.

Bergsetaria, mountain bristle
grass.

Perennial; rhizomatous (rhi-
zomes short and creeping), or
tufted (densely); 300—1200 mm
tall. Leaf blades 100-450 mm
long; 1 .5— 7.0(— 1 0.0) mm wide. Spikelets 2.0-3. 5 mm long.
Culm nodes glabrous; leaf blades linear, finely plicate, flat
or inrolled; panicle open or loosely contracted; bristles
solitary; lower glume 3-nerved; lower lemma rugose.

Flowering October to May. Usually in crevices on rocky
or stony hillsides but also in open woodland and forests.
Common. Biome: Fynbos, Savanna, and Grassland.
Northwards into tropical Africa. Pasture (palatable forage

and hay; fairly drought resistant but susceptable to frost).
Intergrades into S. plicatilis , which is loosely tufted and has
narrowly lanceolate, coarsely plicate leaf blades and
smooth to obscurely rugose upper lemmas.

Description: Stapf 1930 (848), Chippindall & Crook
1976 (89), Stapf 1898-1900 (422), Chippindall 1955 (343),
Clayton et al. 1970-1982 (537). Voucher: Smook 2664,
Smook 2831. PRECIS code 9901280-01200.

T ^ ,s

Fig. 190. Setaria lindenbergiana

297

Setaria megaphylla (Steud.) Dur. & Schinz

(=S. chevalieri Stapf ex Stapf
& C.E. Hubb.)2; (=S. insignis
De Wit) 1.

Riffelblaarsetaria, ribbon
bristle grass.

Very tall, robust perennial;
shortly rhizomatous and tufted
(culms erect and occasionally rooting at the nodes);
900-3000 mm tall. Leaf blades 150-800 mm long; 10-110
mm wide. Spikelets 2-3 mm long. Culms 4-10 mm in
diameter, nodes glabrous; leaf sheaths usually densely
pubescent or at least with hairy margins; leaf blades
lanceolate, conspicuously plicate; panicle open, 400-600
mm long; bristles solitary; lower glume 3-nerved.

Flowering September to May. Riverine or forest grass
on damp soils, mostly in shade, extending to forest margins
and disturbed places such as roadcuttings. Locally common.
Biome: Savanna, Grassland, and Desert. Tropical Africa
and America with a few records from India. Natural pasture
(very palatable), or ornamental (in gardens). Intergrades
into S. plicatilis, which is is a smaller plant with much
narrower leaves and a shorter, sparser panicle. Also closely
related to S. lindenbergiana, which has much narrower,
linear leaves that are finely plicate.

Description: Stapf 1930 (840), Chippindall & Crook
1976 (84), Chippindall 1955 (341), Clayton et al.
1970-1982 (539 & 541). Illustration: Chippindall 1955 (fig.
296 & 297). Voucher: Smook 5480. PRECIS code
9901280-01350.

Setaria nigrirostris (Nees) Dur. & Schinz

Perennial; rhizomatous (rhi-
zome very well developed and
much branched), or tufted (with
few basal leaves); 500-1200 mm
tall. Leaf blades 100-550 mm
long; 4-10 mm wide. Spikelets
3. 5-5.0 mm long. Basal plant
parts very robust and dark colour-
ed; culm nodes hairy; panicle

spike-like, usually cylindrical to the tip; spikelets with
distinct dark tips; lower glume 3-nerved.

Flowering October to April. Often on black turf in open
grassland or on riverbanks. Common. Biome: Savanna and
Grassland. USA. Very similar to S. incrassata , which has
its rhizome oblique and creeping, basal parts straw-
coloured and spikelets 2.5— 3.0(— 3.7) mm long.

Description: Stapf 1898-1900 (423), Chippindall 1955
(345). Illustration: Chippindall 1955 (fig. 300). Voucher:
Liebenberg 8373. PRECIS Code 9901280-01500.

Setaria obscura De Wit

Perennial: hard and densely
tufted; 500-1000 mm tall. Leaf
blades 100-350 mm long; 3^f
mm wide. Spikelets 4.0^4. 6 mm
long. Culm nodes glabrous; leaf
blades rigid and spiny-tipped;
panicle spike-like; bristles
solitary; spikelets disarticulating
below the glumes; lower glume
3-nerved; lower lemma smooth and deeply grooved.

Flowering November to April. Stream banks in high
mountain grassland above 2000 m. Rare. Biome: Grassland.
Endemic. The deep groove on the back of the lower lemma
and the notably long spikelets of this species are unique in
our Setaria species.

Description: Chippindall 1955 (344). Illustration: Chip-
pindall 1955 (fig. 299). Voucher: Killick 1614. PRECIS
code 9901280-01600.

Setaria pallide-fusca (Schumach.) Stapf & C.E. Hubb.

Garden bristle grass, tuin-
setaria.

Annual; loosely tufted; 300-
900 mm tall. Leaf blades 45-170
mm long; 5-9 mm wide. Spike-
lets 2.0-2. 8 mm long. Culm
nodes glabrous; panicle cylindri-
cal, spike-like, normally 10 times
longer than wide; spikelets subtended by 6-10 bristles;
bristles often bright yellow but sometimes dark purple-
brown; lower glume 3-nerved; lower lemma very finely
rugose.

Flowering December to April. In damp soils in
disturbed, weedy places and cultivation. Common. Biome:
Savanna, Grassland, Nama-Karoo and Succulent Karoo.
Tropics worldwide. Domestic use (twisted into ropes by
Basothos to bind grain sheaves), or pasture (natural;
average to good forage value), or weed (colonizer of bare
ground). Closely related to S. ustilata, which has a very
coarsely rugose lower lemma and grows in the shade of
bushes or trees in the drier bushveld regions.

Description: Launert 1970 (160:172), Chippindall 1955
(353), Clayton et al. 1970-1982 (531). Illustration: Chip-
pindall 1955 (fig. 305). Voucher: Smook & Gibbs Russell
2177. PRECIS code 9901280-01800.

Setaria plicatilis (Hochst.) Engl.

Breeblaarpolgras, folded leaf
tussock grass.

Loosely caespitose perennial;
shortly rhizomatous and tufted;

500-1500 mm tall. Leaf blades
100-350 mm long; 8-35 mm
wide. Spikelets 2. 5-3. 3 mm long.

Culm nodes glabrous; leaf blades
narrowly lanceolate and coarsely plicate; panicle open;
bristles solitary; lower glume 3-nerved; lower lemma
smooth or obscurely rugose.

Flowering October to March. Coastal and inland forests
in semi-shade, extending to forest margins and occasionally
into woodlands. Locally common. Biome: Savanna and
Forest. Tropical Africa to Sudan, Ethiopia and Yemen.
Intermediate between S. lindenbergiana , which is densely
caespitose, has much narrower, finely plicate, linear leaves
and a rugose lemma andS. megaphylla , which is very robust
with leaf blades up to 800 mm long and 1 10 mm wide.

Description: Stapf 1930 (847), Clayton et al. 1970-1982
(538). Voucher: Culverwell 643, Van Jaarsveld 177.
PRECIS code 9901280-02000.

Setaria pseudaristata (Peter) Pilg.

( =S . tenuiseta De Wit) 2.

Perennial; shortly rhizomatous
and tufted (culms erect but some-
times very slender); 500-1000
mm tall. Leaf blades 100-300
mm iong; 2-6 mm wide. Spike-
lets 3. 0-3. 5 mm long. Culm
nodes glabrous; leaf blades flat,
not plicate; panicle open, lax; bristles solitary, delicate;
lower glume 3-nerved; lemma rugose.

Flowering February to March. In the shade of riverine
forest. Rare. Biome: Savanna. Tropical Africa. Very similar
to S. plicatilis, which has plicate leaf blades.

Description: Chippindall 1955 (343), Clayton et al.
1970-1982 (535). Voucher: De Winter 782. PRECIS code
9901280-02250.

298

Setaria rigida Stapf

Robust, erect perennial; rhizo-
matous (rhizome stout and
oblique); 1000-1800 mm tall.

Leaf blades 100-300 mm long;

3-6 mm wide (often inrolled).

Spikelets 2. 3-2. 8 mm long. Pani-
cle spike-like, up to 200 mm long;
bristles often grooved and with
sparse long hairs in the lower
part; lower glume 1 -nerved.

Flowering February to March. On stream banks and in
swampy areas. Infrequent. Locally common. Biome:
Savanna and Grassland. Endemic. Distinguished from S.
verticil lata, which also has 1 -nerved lower glumes, but
which is annual and has retrorsely barbed bristles.

Description: Stapf 1898-1900 (426), Chippindall 1955
(352), Clayton et al. 1970-1982 (525). Voucher: Smook
5539. PRECIS code 9901280-02300.

Setaria sagittifolia (A. Rich.) Walp.

(=Cymbosetaria sagittifolia
(A. Rich.) Schweick.) 2.

Arrow grass.

Slender annual; loosely tufted;

120-800 mm tall. Leaf blades
50-300 mm long; (3— )5— 1 1 (— 1 8)
mm wide. Spikelets 2 mm long;

1- 2 mm wide. Culm nodes glabrous; leaf blades (at least
the lower ones) pseudopetiolate, bases sagittate with lobes

2- 30 mm long; panicle open, racemes secund; bristles
solitary; lower glume 3-nerved; lower lemma rugose.

Flowering January to March. Shady places in Savanna
woodland or open glades in forests. Infrequent. Biome: Sa-
vanna and Forest. Northwards to Sudan and Yemen. Natural
pasture. Our only other Setaria with sagittate leaf bases, S.
appendiculata , lacks pseudopetioles.

Description: Chippindall 1955 (355), Clayton et al.
1970-1982 (533). Illustration: Chippindall 1955 (fig. 306),
Clayton et al. 1970-1982 (fig. 128). Voucher: Smook 5397.
PRECIS code 9901280-02450.

Setaria sphacelata (Schumach.) Moss var. sphacelata

(=S. decipiens De Wit) 2;

{=S. flabellata Stapf subsp.
flabellata) 2; ( =S . neglecta De
Wit) 2; (=S. perennis Hack.) 2;

(=S. sphacelata (Schumach.)

Moss subsp. aquamontana De
Wit) 2; [=S. sphacelata
(Schumach.) Moss var.
stolonifera De Wit) 2; (=S.
stenantha Stapf) 2.

Fairly robust perennial; shortly or obliquely rhizomatous
and tufted; 400-1000 mm tall. Leaf blades 100-350 mm
long; 2-5 mm wide. Spikelets 1.5-3. 5 mm long. Culms 2-\
noded, 1-3 mm in diameter; nodes glabrous; sheaths and
leaves glabrous, sparsely or densely hairy; panicle spike-
like, 30-150 mm long; bristles golden-yellow to reddish-
brown; lower glume 3-nerved; lemma rugose.

Flowering October to June. Occupies a wide range of
habitats ranging from streamsides and moist places to rocky
hillsides, usually on well-drained soils. Common. Biome:
Fynbos, Savanna, and Grassland. Tropical east Africa.
Cultivated hay and pasture. Very difficult to separate from
S. sphacelata var. sericea, which is larger with wider leaf
blades. Intermediates are common.

Fig. 189.

Description: Launert 1970(160:172), Chippindall 1955
(351), Clayton et al. 1970-1982 (528). Illustration: Chip-
pindall 1955 (fig. 302 & 303). Voucher: Smook 5437, Codd
5373. PRECIS code 9901280-02500.

299

Setaria sphacelata (Schumach.) Moss var. sericea

(Stapf) Clayton

(=S. almaspicata De Wit) 2;

(=S. anceps Stapf ex Massey) 2:

(=S. cana De Wit) 2; (=S.
flabelliformis De Wit) 2; (=S.
sphacelata (Schumach.) Moss
subsp. nodosa De Wit) 2; (=S.
sphacelata (Schumach.) Moss
subsp. pyropea De Wit) 2.

Perennial; shortly rhizomatous and tufted; 1000-2000
mm tall. Leaf blades 100-500 mm long; 3-10 mm wide.
Spikelets 1.5-3. 5 mm long. Culms 4-10-noded, 3-6 mm in
diameter, nodes glabrous; basal sheaths and leaves gla-
brous, densely or sparsely pubescent; panicle spike-like,
70-250 mm long; bristles golden-yellow to reddish brown;
lower glume 3-nerved; lower lemma rugose.

Flowering October to June. Occupies a wide range of
habitats ranging from riversides and swampy areas to rocky
hillsides. Common. Biome; Savanna and Grassland.
Tropical Africa, cultivated elsewhere. Good hay and pas-
ture. Intergrades with S. sphacelata var. sphacelata , which
is a smaller plant with narrower leaves, and S. sphacelata
var. splendida , which is larger, robust and has wider leaves.

Description: Chippindall 1955 (349, 346, 351), Clayton
et al. 1970-1982 (529). Voucher; Smook 2583, Webster 8.
PRECIS code 9901280-02455.

Setaria sphacelata (Schumach.) Moss var. splendida

(Stapf) Clayton

( =S . splendida Stapf) 2.

Extremely robust, almost
reed-like perennial; shortly rhizo-
matous and tufted (usually with
only a few basal leaves);

1800-3000 mm tall. Leaf blades
300-800 mm long; 6-18 mm
wide. Spikelets 2. 3-2.7 mm long.

Plants and culm nodes glabrous; panicle spike-like,
1 50— 300(— 500) mm long; bristles golden-yellow, 7-15 per
spikelet cluster; lower glume 3-nerved; upper lemma
rugose.

Flowering January to June. Swampy areas or
floodplains, often in the water. Rare (in the wild but often
cultivated). Biome: Savanna. Scattered localities

northwards through east Africa to Sudan. Good hay and
pasture. The most robust variety in the S. sphacelata
complex.

Description: Stapf 1930 (799), Chippindall 1955 (351),
Clayton et al. 1970-1982 (530). Voucher: Killick &
Leistner 3412. PRECIS code 9901280-02570.

Setaria sphacelata (Schumach.) Moss var. torta (Stapf)

Clayton PI

(=S. flahellata Stapf subsp.
natalensis De Wit) 2; (=S.
homblei De Willd.) 2; ( =S . torta
Stapf) 2.

Twisted leaf bristle grass.

Perennial; shortly rhizomatous
and tufted (most leaves basal;
culms occasionally flat and rooting at the nodes);
300-500(-1000) mm tall. Leaf blades 100-300 mm long;
1-3 mm wide (mostly folded or inrolled). Spikelets2. 5-3.0
mm long. Basal sheaths usually strongly keeled and
flabellate; old leaves curly; culm nodes glabrous; panicle
spike-like, 25^4-5 mm long; bristles 7-15 per spikelet
cluster, usually dark purple-brown but occasionally
yellowish; lower glume 3-nerved; lemma rugose.

Flowering September to March. On rocky outcrops,
hillsides, open woods and grassland on well-drained soils.

Common (usually scattered amongst other grasses). Biome:
Fynbos, Savanna, Grassland, Nama-Karoo, and Succulent
Karoo. Tropical Africa. Well eaten natural pasture. Very
slender and small when compared to the other varieties,
representing the smallest extreme of the S. sphacelata
complex.

Description: Stapf 1930 (801), Chippindall & Crook
1976 (92), Clayton et al. 1970-1982 (529). Voucher:
Scheepers 1451, Manders 4. PRECIS code 9901280-02590.

300

Setaria ustilata De Wit

Annual; loosely tufted (culms
erect or geniculate); 120-650 mm
tall. Leaf blades 50-170 mm
long; 8-12 mm wide. Spikelets
2. 3-2. 7 mm long. Culm nodes
glabrous; panicle spike-like,
usually ovate but up to 5 times
longer than wide; bristles 6-10
per spikelet; lower glume 3-
nerved; lemma very coarsely rugose.

Flowering January to May. In the drier bushveld regions
in the shade of trees and bushes. Infrequent. Biome: Savan-
na. ?Endemic. Closely related to S. pallide-fusca , which has
a very finely rugose lemma and is a common weed in moist
areas and cultivation.

Description: Chippindall 1955 (355), Clayton et al.
1970-1982 (531). Voucher: Smook 2619, Hardy, Retief &
Herman 533 1 . PRECIS code 990 1 280-03 1 00.

Setaria verticillata (L.) Beauv.

Klitssetaria, bur bristle grass.

Annual; loosely tufted (often
sprawling); 300-1000 mm tall.

Leaf blades 50-300 mm long;

6-22 mm wide. Spikelets 1.5-2. 5
mm long. Panicle spike-like,
often shortly branched and
interrupted in the lower part,

20- 1 50 mm long; bristles retorsely barbed, often entangled;
lower glume I -nerved.

Flowering December to May. Ruderal in disturbed areas,
cultivation, cattle kraals and along paths, often in damp,
shady places. Common. Biome: Fynbos, Savanna, Grass-
land, and Nama-Karoo. Old world tropics, introduced to the
USA. Domestic use (inflorescences used in east Africa to
keep rats from harvested grains. They adhere to small
animals and cause much suffering. Plants are also used for
weaving hats and toys), or pasture (good palatable hay and
forage), or weed (troublesome in gardens and cultivation).
Uniquely distinguished from our other Setaria species by
the retrorsely barbed bristles.

Description: Launert 1970 (160: 171), Stapf 1930 (823),
Chippindall & Crook (84), Stapf 1898-1900 (429), Hitch-
cock & Chase 1950 (699), Chippindall 1955 (355), Clayton
et al. 1970-1982 (522). Illustration: Clayton et al.
1970-1982 (fig. 127), Hitchcock & Chase 1950 (fig. 1566).
Voucher: Dahlstrand 2485, Du Toit 174. PRECIS code
9901280-03200.

Fig. 192. PI. 176.

Sorghastrum Nash

Dipogon Steud., Poranthera Raf.

Annual, or perennial; caespitose. Culms 700-1500 mm
high; herbaceous; branched above, or unbranched above.
Ligule an unfringed membrane to a fringed membrane.
Plants bisexual , with bisexual spikelets. The spikelets of
sexually distinct forms on the same plant ( hermaphrodite
and sterile)-, overtly heteromorphic (in that the sterile
spikelets are reduced to pedicels), or homomorphic (rarely
the pedicellate spikelets are well developed and simiar to
the sessile ones).

Inflorescence paniculate (narrowly elongated, more or
less unilateral panicles of much-reduced, capillary
‘racemes')-, open (usually narrow); espatheate ; not com-
prising ‘partial inflorescences’ and foliar organs. Spikelet-
bearing axes very much reduced (the ultimate units with
very few spikelets, often only one accompanied by the
sterile pedicel)-, disarticulating at the joints (but the
disarticulating units much reduced) or falling entire (when
reduced to one joint).

Spikelets nearly always in pairs (but ostensibly solitary,
by virtue of the ‘pedicellate’ member being reduced to its
pedicel — by contrast with Sorghum)-, consistently in ‘long-
and-short’ combinations (but the sterile member of each
combination is nearly always reduced to its pedicel).
Pedicels free of the rachis. The sessile spikelets hermaphro-
dite. The ’pedicellate spikelets’ sterile (usually reduced to
pedicels). Female-fertile spikelets 5-8 mm long; com-
pressed dorsiventrally (plump); falling with the glumes (and
the joint). Glumes two; more or less equal; awnless; very
dissimilar (the lower flattened and often hairy on the back,
the upper glabrous and slightly keeled above). Lower glume
9 nerved. Proximal incomplete florets l\ epaleate; sterile.

Female-fertile florets 1 . Lemmas less firm than the
glumes; incised; awned. Awns 1; median; from the sinus;
geniculate; much longer than the body of the lemma. Palea
present, or absent; when present conspicuous but relatively
short, or very reduced. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. About 20 species. Mainly
tropical and subtropical Africa and America. Helophytic to
mesophytic; in shade, or in open habitats (savanna and
woodland margins, often in wet places); glycophytic.
Namibia, Botswana, Transvaal, and Natal. 2 indigenous
species.

301

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Leaf sheath not appendaged at apex; leaf blade base
expanded; lowest node of culm to 5 cm long; awns
3— 8(— 1 0) mm long, straight or bent and twisted .

S. friesii

Leaf sheath with erect appendages at apex; leaf blade
base tightly rolled, narrower than middle portion of
blade; lowest node of culm to 10 mm long; awns
8-16 mm long, never straight, always bent and
twisted S. stipoides

Sorghastrum friesii (Pilg.) Pilg.

Perennial; tufted; 700-1200
mm tall. Leaf blades to 200 mm
long; 2-6 mm wide. Spikelets
5-7(-8) mm long (all alike, each
with an empty pedicel). Leaf
blade base expanded, sheath
mouth not appendaged; awns
3— 8(— 10).

Flowering January to April.

Wet places. Infrequent. Biome: Savanna. Endemic.

Description: Chippindall 1955 (467). Illustration: Chip-
pindall 1955 (fig. 382). Voucher: Pole Evans 17-4-1934.
PRECIS code 9900461-00100.

Sorghastrum stipoides (Kunth) Nash

(=S. rigidifolium (Stapf)

Chippind.) 2.

Perennial; tufted; to 1500 mm
tall. Leaf blades to 450 mm long;

2-5 mm wide. Spikelets 4-7 mm
long (all alike, each with an
empty pedicel). Leaf blade base
tightly rolled, sheath mouth with
erect appendages; awns 8-16 mm long.

Flowering December to April. Wet places. Infrequent.
Tropical Africa and South America.

Description: Chippindall 1955 (468), Clayton et al.
1970-1982 (732). Illustration: Clayton et al. 1970-1982
(fig. 169). Voucher: Ward 6085. PRECIS code 9900461-
00200.

Fig. 193. PI. 177.

Sorghum Moench

Blumenbachia Koel., Sarga Ewart & White.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 600-3000 mm
high; herbaceous. Leaf blades usually flat. Ligule an
unfringed membrane to a fringed membrane, or a fringe of
hairs (rarely). Plants bisexual, with bisexual spikelets; with
hermaphrodite florets. The spikelets of sexually distinct
forms on the same plant ; overtly heteromorphic (the
pedicellate much narrower and awnless).

Inflorescence paniculate ( the primary branches usually
whorled)\ open, or contracted; espatheate ; not comprising
‘partial inflorescences’ and foliar organs. Spikelet-bearing
axes very much reduced ‘racemes’ (with 1 — 6(-8) articles
only); with very slender rachides; disarticulating, or
persistent (in cultivated forms); falling entire (when
reduced to one joint), or disarticulating at the joints.
‘Articles’ without a basal callus-knob.

Spikelets not secund\ consistently in ‘long-and-short’
combinations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The ‘longer’
spikelets male-only, or sterile (but not reduced to the

pedicel, by contrast with Sorghastrum). Female-fertile
spikelets compressed dorsiventrally, falling with the
glumes, or not disarticulating (in cultivated forms). Glumes
two; more or less equal; awnless; very dissimilar (lower flat
or rounded on the back save at summit, upper naviculate).
Proximal incomplete florets 1\ epaleate; sterile.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline, ciliate); incised; awnless to mucronate
(rarely), or awned. Awns when present 1; from the sinus;
geniculate; much shorter than the body of the lemma, to
much longer than the body of the lemma. Palea present, or
absent; when present relatively long, or conspicuous but
relatively short, or very reduced. Lodicules 2; fleshy; ciliate
(usually). Stamens 3. Ovary glabrous. Fruit small, or
medium sized, or large; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5. Panicoideae; Andropogonodae; Andropo-
goneae; Andropogoninae. 24 species. Tropical and
subtropical. Mesophytic; in shade, or in open habitats
(savanna and forest margins, alluvial plains and disturbed
ground); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. Indigenous species (2), naturalized species (2),
cultivated (species 1).

Intergeneric hybrids with Saccharum.

References. 1. De Wet 1978. Amer. J. Bot. 65: 477. 2.
Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Nodes with a ring of spreading white hairs; sessile

spikelets black at maturity S. versicolor

Nodes lacking spreading white hairs; sessile spikelets

variously coloured at maturity 2

2(1). Plants perennial, with long rhizomes . S. halepense
Plants annual or short-lived perennial, without

rhizomes 3

3(2). Leaf blades less than 15 mm across

S. bicolor subsp. drummondii

Leaf blades more than 20 mm across

S. bicolor subsp. arundinaceum

302

Sorghum bicolor (L.) Moench subsp. arundinaceum

(Desv.) De Wet & Harlan

(=S. verticilliflorum (Steud.)

Stapf) 1.

Common wild sorghum, wilde-
graansorghum.

Short-lived perennial (without
rhizomes), or annual; to 2500 mm
tall. Leaf blades 20-30 mm wide.

Spikelets (sessile) 5-7 mm long.

Disturbed places. Locally common. Biome: Savanna,
Grasslartd, and Nama-Karoo. Throughout tropical Africa to
Australia, introduced to tropical America.

Description: Chippindall 1955 (460). Voucher: De
Winter & Leistner 5 163. PRECIS code 9900460-00300.

PI. 178.

Sorghum bicolor (L.) Moench subsp. drummondii

(Steud.) De Wet

(=S. sudanense (Piper)

Stapf) 1.

Sudan grass, witkafferkoring,
shattercane.

Annual; to 3000 mm tall. Leaf
blades 8-15 mm wide. Spikelets
(sessile) 6-7 mm long.

Infrequent. Naturalized, native to tropical north Africa.
Pasture (planted as a fodder), or weed.

Description: Chippindall 1955 (459). Voucher: Nat.
Herb Pretoria B. PRECIS code 9900460-00350.

Sorghum halepense ( L. ) Pers.

Fig. 194 .

(=S. almum Parodi) 1.

Johnson grass.

Perennial; usually strongly
long-rhizomatous; to 2500 mm
tall. Leaf blades 10-30 mm wide.

Spikelets (sessile) 4.0-5.5(-7.0)
mm long (pedicellate spikelet
longer).

Flowering usually December to May (occasionally at
other times). In disturbed places. Common. Naturalized
from the Mediterranean region, now widely naturalized.
Biome: Fynbos, Savanna, Grassland, and Nama-Karoo.
Worldwide in warm areas. Weed (especially difficult to
eradicate because of its long, deeply buried rhizomes).

Description: Chippindall 1955 (460). Illustration:
Chippindall 1955 (fig. 377), Hitchcock & Chase 1950 (fig.
1177). Voucher: Webber 2-2-23. PRECIS code 9900460-
02600.

Sorghum versicolor Anderss.

Swartsaadgras, black wild
sorghum.

Perennial, or annual; 600-
1200 mm tall. Leaf blades 4-8
mm wide. Spikelets (sessile) 4—7
mm long. Nodes with spreading
white hairs, spikelets black at
maturity.

Flowering December to May. Black turf soil. Common.
Biome: Savanna and Grassland. Tropical Africa.

Description: Chippindall 1955 (459), Clayton et al.
1970-1982 (729). Illustration: Chippindall 1955 (pi. 14).
Voucher: De Winter 2915. PRECIS code 9900460-03700.

Spartina Schreber

Chauvinia Steud., Limnetis Rich., Ponceletia Thours,
Psammophila Schult., Solenachne Steud., Trachynotia
Michaux, Tristania Poir.

Perennial; long-rhizomatous to long-stoloniferous, or
caespitose. Culms 200-3000 mm high; herbaceous. Leaf
blades flat, or rolled. Ligule a fringe of hairs.

Fig. 195. Spartina maritima

303

Inflorescence of spike-like main branches (with 2 to
many long or short spikes, borne racemosely on the main
axis)', axes not ending in spikelets (their slender, naked tips
often prolonged)', espatheate. Spikelet-bearing axes
persistent.

Spikelets biseriate; 6-18 mm long; compressed laterally;
falling with the glumes. Glumes two; very unequal (the
upper longer); long relative to the adjacent lemmas (i.e., the
upper, which often exceeds the lemma); awned (shortly),
or awnless; similar (coriaceous or membranous). All florets
female-fertile; proximal incomplete florets absent.

Female-fertile florets I . Lemmas 1-3 nerved; entire, or
incised; awnless. Palea present; relatively long. Stamens 3
(the anthers relatively long). Ovary glabrous. Fruit medium
sized; fusiform; hilum short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4; PCK
(anglica)', XyMS+. PCR sheath outlines uneven. PCR
sheath extensions present. Maximum number of extension
cells 7-8. PCR cell chloroplasts with well developed grana;
centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number,* = 7 and 10. Chloridoideae; Chlorideae sensu lato.
16 species. Temperate America, coastal Europe, Africa,
Tristan da Cunha. Commonly adventive. Hydrophytic to
helophytic; in open habitats; halophytic. Namibia and Cape
Province. 1 indigenous species.

References. 1. Launert. 1970. FSWA.

Species treatment by M. Koekemoer.

Spartina maritima (Curtis) Fernald

Fig. 195. PI. 179.

(=S. capensis Nees ex
Trin.) 1.

Cape cord grass, strandkweek.

Perennial; hydrophyte and
rhizomatous, or stoloniferous;

200-800 mm tall. Leaf blades
120-190 mm long. Spikelets
12-15 mm long. Leaf blades inrolled; spikes robust, one-
sided, usually 2-3 (rarely 1), not spreading.

Flowering November to April. Along coasts on intertidal
mud flats, around estuaries or submerged in lagoons. Local-
ly common. Biome: Fynbos and Succulent Karoo. Atlantic
coastlines. Hybridization, back-crossing and polyhaploidy
complicate taxonomy in this genus, which has 16 species
worldwide but only one representative here.

Description: Chippindall 1955 (208). Illustration: Chip-
pindall 1955 (fig. 185). Voucher: Boucher 2999. PRECIS
code 9902930-00200.

Sphenopus Trin.

Annual', caespitose. Culms 70-300 mm high; herba-
ceous. Leaf blades linear; flat (to almost filiform). Ligule
an unfringed membrane.

Inflorescence paniculate', open; with conspicuously di-
varicate branchlets (spikelets numerous and very small);
espatheate. Spikelet-bearing axes persistent.

Spikelets 1. 5-2.5 mm long; compressed laterally; disar-
ticulating above the glumes. Glumes two; minute to rela-
tively large (G2 0.5-0. 9 mm long in S. divaricatus)', very
unequal; markedly shorter than the spikelets; awnless;
similar (hyaline, rounded to emarginate). Lower glume 0
nerved. All florets female-fertile; proximal incomplete
florets absent.

Female-fertile florets 2-7 . Lemmas decidedly firmer
than the glumes; 3 nerved; entire; awnless; non-carinate
(but keeled on all three veins). Palea present; relatively
long. Lodicules 2; membranous; glabrous. Stamens 3.

Ovary glabrous. Fruit small; hilum short.

Cytology, classification, distribution. Chromosome base
number, x = 6 and 7. Pooideae; Poodae; Poeae. 2 species.
Mediterranean to western Asia. In open habitats; maritime-
arenicolous and halophytic. Cape Province. 1 naturalized
species.

References. 1. Linder. Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

Fig. 196. Sphenopus divaricatus

304

Sphenopus divaricatus (Gouan) Reichb.

Fig. 196. PI. 180.

( -Poa divaricata Gouan) 1.

Annual; tufted (slender, outer
stems geniculate and spreading
from the base); 70-200(-300)
mm tall. Leaf blades 30-70 mm
long; setaceous or rolled, to 1 mm
wide. Spikelets 2-3 mm long.

Ligule membranous, to 4 mm
long; panicle open, delicate, with multiple branching;
spikelets 2-5-flowered, pedicellate, pedicels 1. 5-7.0 mm
long; glumes unequal, lower scale-like, upper to 1 mm long;
lemmas 1.25-1.50 mm long, obtuse or rounded.

Flowering August to October. Coastal areas on mudflats
along rivers or salty marshes or in hollows between dunes.
Rare. Locally common. Naturalized from Europe. Biome:
Fynbos and Succulent Karoo. Southwestern Europe,
Mediterranean region eastwards to central Asia, introduced
to South Africa and Australia.

Description: Bor 1985 (1740), Linder (59). Voucher:
Smook 3650. PRECIS code 9903790-00100.

Sporobolus R.Br.

Agrosticula Raddi, Bauchea Fourn., Cryptostachys
Steud., Diachyrium Griseb., Spermachiton Llanos,
Triachyrum A. Br.

Annual (rarely), or perennial; long-rhizomatous, or
long-stoloniferous, or caespitose, or decumbent. Culms
50-1600(-3000) mm high; herbaceous. Leaf blades linear;
flat, or folded, or rolled. Ligule a fringed membrane
(narrow), or a fringe of hairs.

Inflorescence a false spike, with clusters of spikelets on
reduced axes, or a single raceme (rarely), or paniculate ;
open, or contracted; espatheate. Spikelet-bearing axes
persistent.

Spikelets 0.8-3. 5 mm long (rarely, to 6 mm)\ compressed
laterally to not noticeably compressed (often fusiform); dis-
articulating above the glumes. Hairy callus absent. Glumes
two; very unequal (G1 often very short), or more or less
equal; decidedly shorter than the adjacent lemmas, or long
relative to the adjacent lemmas; awnless; persistent or
subpersistent, thinly membranous or hyaline. Lower glume
1 nerved. All florets female-fertile, or rarely distal incom-
plete florets also present; proximal incomplete florets
absent.

Female-fertile florets 1 . Lemmas 1 nerved, or 2 nerved;
entire; awnless. Palea present; relatively long. Lodicules
when present 2; fleshy; glabrous. Stamens 2-3. Ovary
glabrous. Fruit small (0.3-2 mm); hilum short; pericarp free
(commonly swelling when wet, forcibly ejecting the seed);
embryo large.

Photosynthetic pathway and related features. C4; PCK
(6 species), or NAD-ME (4 species); XyMS+. PCR sheath
outlines uneven, or even. PCR sheath extensions present,
or absent. Maximum number of extension cells when
present 2-5. PCR cell chloroplasts ovoid, or elongated; with
well developed grana; centrifugal/peripheral, or centripetal.

Cytology, classification, distribution. Chromosome base
number,* = 9 and 10. Chloridoideae; Chlorideae sensu lato.
160 species. Tropical and warm temperate. Mesophytic, or
xerophytic; in diverse habitats; maritime-arenicolous, or
halophytic, or glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 39 indigenous species.

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1974 FTEA. 3. De Winter & Vorster. 1974. Bothalia
1 1: 295. 4. Goossens. 1938. Trans. Roy. Soc. S.A. 26: 173.

Species treatment by M. Koekemoer.

1(0). Panicle with primary branches in whorls, or with at
least the lowermost branches in a single whorl . 2
Panicle with primary branches not whorled .... 15
2(1). Plants less than 180 mm tall at maturity; leaf blades
10-30 mm long, 1-5 mm wide, forming a basal
rosette, the margins with stiff spreading hairs;
culms 1-noded; spikelets pendulous at maturity,

grains discoid S. discosporus

Plants not as above 3

305

3(2). Plants annual 4

Plants perennial 6

4(3). Spikelets longer than 2 mm, fewer than 4 per primary
branch; grains 1.2-1. 9 mm in diameter;
inflorescence lacking viscid patches on the central

axis and primary branches S. panicoides

Spikelets shorter than 2 mm, more than 4 per primary
branch; grains less than 1 mm in diameter;
inflorescence with viscid patches on the central axis

and primary branches 5

5(4). Lower glume linear, more than 0.5 mm long and at
least half the length of the spikelet; viscid patches

rounded to ovate S. stolzii

Lower glume obovate, shorter than 0.5 mm, reduced
to a tiny scale, or absent; viscid patches more than
3 times longer than wide . . . S. coromandelianus
6(3). Spikelets shorter than 1.5 mm; upper glume
prominently keeled, keel scabrid, central nerve
lighter in colour than the rest of the glume; primary

branches with spikelets in the upper half

S. nitens

Spikelets longer than 1.5 mm; upper glume rounded,
central nerve not distinctly coloured; primary
branches with spikelets over the whole length or

only in the upper part 7

7(6). Leaf blades rounded at the base; upper glume 2/3— 3/4
the spikelet length; lower glume 1/4 the spikelet

length . S. kentrophyllus

Leaf blades tapering at the base; upper glume as long
as or slightly longer than the spikelet; lower glume

1/4 to as long as the spikelet 8

8(7) Lower glume longer than 2/3 the spikelet length; leaf
blades inrolled; upper glume longer than the

spikelet S. centrifugus

Lower glume shorter than 2/3 the spikelet length; leaf
blades flat or sometimes inrolled; upper glume as

long or longer than the spikelet 9

9(8). Leaf blade margins with flexuous hairs longer than

0.5 mm 10

Leaf blade margins glabrous or with scabrid hooks
less than 0.5 mm long 12

10(9). Spikelets very densely clustered on the upper 1/3
of the primary branches; upper glume as long as
spikelet; lower glume 1/2 the spikelet length . .

S. pectinatus

Spikelets not clustered, covering the whole length
or only the upper half of the branches,
overlapping not more than half the spikelet
length; upper glume longer than spikelet; lower

glume 1/2— 3/4 the spikelet length 11

11(10). Panicle pyramidal, 100-200 mm long, with fewer
than 10 whorls; lower glume L'7^1.0 mm long,

1/2-3/4 the spikelet length S. congoensis

Panicle linear to lanceolate, 200-430 mm long, with
more than 10 whorls; lower glume 0.5-1. 6 mm

long, shorter than 1/2 the spikelet length

S. sanguineus

12(9). Leaf blades rigid, shorter than 30 mm; rhizomes
with very short intemodes, creeping horizontally
and branching profusely; plants mat-forming .

S. ludwigii

Leaf blades not rigid, longer than 30 mm, rhizome
not horizontally creeping or profusely branched;

plants rhizomatous or tufted 13

13(12). Spikelets 2. 5-5.0 mm long; lower glume 2/3— 3/4

the spikelet length; panicle contracted

S. mauritianus

Spikelets 1.5-2. 7 mm long; lower glume 1/4 — 1/3

the spikelet length; panicle open 14

14(13). Basal leaf sheaths yellow, hard, glossy and brittle;

culms erect; plants seldom stoloniferous; leaf
blades less than 3 mm wide, often inrolled,

30-140 mm long S. rangei

Basal leaf sheaths dull and papery; culms
geniculate; plants almost always stoloniferous;
leaf blades flat, more than 5 mm wide, 20-300

mm long S. ioclados

15(1). Panicle branches not dichotomous; spikelets borne
on short pedicels along the length of the

branchlets 16

Panicle branches dichotomous; spikelets solitary at

the tips the of branchlets 31

16(15). Panicle narrow and spike-like 17

Panicle linear to open, not spike-like 21

17(16). Upper glume as long as, or slightly longer than the
spikelet; lower glume 3/4 the spikelet length . .

S. virginicus

Upper glume to 2/3 the spikelet length; lower glume

less than 1/2 the spikelet length 18

18(17). Spikelets very densely clustered so that central axis
is not visible; primary branches shorter than 3

mm; panicle to 5 mm wide S. spicatus

Spikelets not clustered to hide the central axis;
primary branches to 30 mm long; panicle wider

than 5 mm 18

19(18). Leaf blades longer than 200 mm; upper glume 1/2

the spikelet length; spikelets very dense

S. africanus

Leaf blades shorter than 200 mm; upper glume
about as long as the spikelet; spikelets somewhat

loose 20

20(19). Plants 200-360 mm tall; spikelets 2. 2-2. 5 mm long;

glumes and lemmas membranous . S. albicans
Plants 500-800 mm tall; spikelets 2. 5-3.0 mm long;
glumes and lemmas cartilaginous

S. bechuanicus

21(16). Panicle pyramidal or ovate, open, not more than

three times longer than wide 22

Panicle linear, open or contracted, more than five
times longer than wide (in S.fourcadii sometimes

ovate) 23

22(21 ). Glumes keeled along the whole length or only at the
tips, keel scabrid; rhizome long and deeply
buried; leaves rigid . . . . S. sp. (=Smook 3429)
Glumes not keeled; rhizome creeping near ground
level, internodes short; leaves fine and curly . .

S. nervosus

23(21). Lemma notably long and fine, to 1.5 times the
length of the spikelet; plants annual . S. molleri
Lemma as long as the spikelet; plants perennial .

24

24(23). Upper and lower glumes more or less equal and as
long as the spikelet; glumes keeled, keel scabrid;

plants reed-like S. consimilis

Upper glume half to as long as spikelet and lower
glume less than 1/2 the spikelet length; glumes

not keeled; plants not reed-like 25

25(24). Glumes subequal, both glumes less than 1/3 the

spikelet length S. pyramidalis

Glumes unequal, upper glume 1/2 to slightly longer
than the spikelet; lower glume shorter than the

spikelet 26

26(25). Old leaf sheaths splitting into fibres with age;

leaves setaceous, relatively short and forming a

cushion at the base S. pellucidus

Old leaf sheaths not splitting into fibres; leaves not
setaceous or forming a cushion at the base . 27
27(26). Culms more than 3 mm in diameter ( 1 00 mm above
base); panicle 350-750 mm long, much
branched, branches lax; upper glume almost as

long as the spikelet S. maeranthelus

Culms less than 3 mm in diameter (100 mm above
base); panicle shorter than 500 mm, branches
usually firm; upper glume less than 3/4 the

spikelet length 28

28(27). Spikelets very densely clustered on relatively short
primary branches; branches rigid and contracted;
panicle almost spike-like; grains ellipsoid,

1.1-1. 2 mm long S. africanus

Spikelets not clustered; branches lax and spreading;
panicle linear to pyramidal; grains to 1 mm long

306

29(28). Plants usually small, 250-400(-700) mm tall;

panicles with a few branches far apart, branches
almost horizontally spreading at maturity; upper
glume 2/3 and lower glume 1/3 the spikelet

length S. fourcadii

Plants usually taller than 1000 mm, often robust;
panicle fairly dense with numerous racemes;
racemes never 'spreading more than 60 degrees
from the main axis at maturity; glume length very
variable, upper glume 1/2-3/4 and lower glume

1/4— 3/4 the spikelet length 30

30(29). Plants with characteristic oblique rhizomes; lower
leaf sheaths herbaceous; upper glume about 2/3

the spikelet length S. fimbriatus

Plants lacking obvious rhizomes; lower leaf sheaths
papery; upper glume about 1/2 the spikelet length

S. natarensis

31(15). Panicle with long stiff hyaline hairs in all or some

of the axils (2-15 hairs per axil) 32

Panicle lacking hairs in the axils 34

32(31). Panicle with very few hairs occuring in some axils;

glumes unequal, upper glume 2/3 and lower
glume 1/2 the spikelet length . . . S. welwitschii
Panicle with many hairs in almost all the axils;
glumes more or less equal and 1/2 the spikelet

length 33

33(32). Rachilla extention present; plants not fibrous at the

base S. subtilis

Rachilla extention absent; plant base fibrous ....

S. conrathii

34(31). Spikelets 2.0-2. 9 mm long S. salsus

Spikelets shorter than 2 mm 35

35(34). Basal leaf sheaths splitting into fibres at maturity

36

Basal leaf sheaths not splitting into fibres .... 37

36(35). Fibrous remains of leaf sheaths with a dense mass

of woolly hairs between the fibres

S. stapfianus

Fibrous remains of leaf sheaths lacking woolly hairs

between the fibres S. festivus

37(35). Plants annual; leaf blades to 4 mm wide

S. engleri

Plants perennial; leaf blades setaceous or to 3 mm

wide 38

38(37). Leaf blades setaceous, rigid and forming a very
dense tuft; old dead leaves and sheaths are
persistent and form a dense cushion below the

new growth S. nebulosus

Leaf blades flat; plants not tufted, with a much-

branched creeping rhizome 39

39(38). Leaf blades shorter than 35 mm, rounded at the tips;

culms mostly 1-noded; leaves mostly basal with
cauline leaves much shorter than the basal ones

S. tenellus

Leaf blades 40-120 mm long, tapering to a fine
point; culms 2-5-noded; leaves basal and cauline
with cauline leaves usually much longer than the
basal ones S. acinifolius

Sporobolus acinifolius Stapf

Kalkgras, limestone dropseed.

Mat-forming perennial; rhizo-
matous (rhizome long and much
branched); 150-430 mm tall. Leaf
blades 40-120 mm long; to 3 mm
wide. Spikelets 1-2 mm long.

Cauline leaves usually much
longer than the basal leaves, leaf
tips tapering to a fine point; panicle dichotomously
branched, axils without hairs; spikelets solitary on branchlet
tips.

Flowering February to May. Brackish calcarous soil on
pans or at the edge of water. Locally common. Biome: Fyn-
bos, Savanna, and Nama-Karoo. ?Endemic. Very similar to
S. tenellus, which has leaf blades shorter than 35 mm and
leaf tips rounded, and S. salsus, which has larger spikelets.

Description: Goosens 1938 (191), Stapf 1898-1900
(581), Chippindall 1955 (214). Illustration: Chippindall
1955 (fig. 189). Voucher: Smith 195. PRECIS code
9902830-00100.

Sporobolus africanus (Poir.) Robyns & Tournay

Fig. 197.

(=S. capensis (Willd.)

Kunth) 2.

Dropseed, taaipol.

Perennial; rhizomatous and
tufted: 280-1500 mm tall. Leaf
blades 200-400 mm long; 1—4
mm wide. Spikelets 2.0-2. 8 mm
long. Panicle dense, not whorled, almost spike-like,
branches relatively short and rigid, central axis usually
visible; lower glume 1/4-1/2 the spikelet length; upper
glume 1/2 the spikelet length; grain ellipsoid, 1.1-1. 2 mm
long.

Flowering October to April. Mainly in disturbed places
and along streams. Common. Biome: Fynbos, Savanna, and
Grassland. Northwards through tropical east Africa to
Ethiopia. Traditional medicine (local application to wounds
and snake bite). The typical form can be distinguished from
S . fourcadii , S. fimbriatus, S. pyramidalis and S. natalensis,
by its contracted, almost spike-like panicle with short firm
branches and longer grains. S. africanus, S. fimbriatus, S.
natalensis and S. pyramidalis form an interlaced group of
species, in which the typical forms are overshadowed by a
large number of intermediates, for which Clayton(1974)
suggests a hybrid origin. Further research is needed to
distinguish these specimens satisfactorily.

Description: Goossens 1938 (213), Launert 1970
(160:181), Chippindall 1955 (225), Clayton et al.
1970-1982 (375). Illustration: Chippindall 1955 (fig. 198).
Voucher: Smook 5456, Pole Evans 139. PRECIS code
9902830-00200.

Sporobolus albicans Nees

Mat-forming perennial; rhizo-
matous; 200-360 mm tall. Leaf
blades 5-12 mm long; 1 mm
wide. Spikelets 2. 0-2. 5 mm long.

Inflorescence 25-30 mm wide,
spike-like but spikelets not
densely clustered to hide the
central axis; glumes and lemmas
membranous, pale yellow.

Flowering February to April. Limestone pans or dried
up depressions. Locally common. Biome: Grassland and
Nama-Karoo. Endemic. Very similar to S. virginicus, which
has longer glumes, and S. bechuanicus, which has longer
spikelets and cartilaginous lemmas.

Description: Goossens 1938 (194), Launert 1970
(160:181), Stapf 1898-1900 (580), Chippindall 1955 (214).
Voucher: Smook & Gibbs Russell 2439. PRECIS code
9902830-00300.

Sporobolus bechuanicus Goossens

Perennial; tufted; 500-800
mm tall. Leaf blades 120-200
mm long; 3.0-3. 5 mm wide.

Spikelets 2. 5-3.0 mm long. Pani-
cle spike-like, more than 5 mm
wide; spikelets not hiding the
central axis; glumes and lemmas
cartilaginous.

Flowering January to April.

307

Brackish soil on seasonally flooded pans. Rare. Biome: Sa-
vanna. ?Endemic. Very similar to S. albicans, which has
smaller spikelets and membranous lemmas, and to S.
virginicus, which has longer glumes.

Description: Goossens 1938 (210), Chippindall 1955
(226). Voucher: Pole Evans 3277. PRECIS code

9902830-00500.

Sporobolus centrifugus (Trin.) Nees

(=S. schlechteri
Schweick.) 2.

Olive dropseed.

Perennial; rhizomatous (rhi-
zome can be long and creeping);

180-900(-1060) mm tall. Leaf
blades 60-300 mm long, filiform;

1.0-1. 5 mm wide. Spikelets 2.5 — 4.2 mm long. Basal sheaths
hard, brittle, glossy, yellow or brown; panicle contracted
and unobtrusively whorled; spikelets in lowest whorl often
sterile and disarticulating at maturity; lower glume slightly
shorter than spikelet; upper glume longer than spikelet.

Flowering October to April. High mountainveld or
highveld on humiferous well-drained soils. Locally com-
mon. Biome: Savanna, Grassland, and Afromontane.
Tropical east Africa. Similar to S. mauritianus, S.
sanguineus and S. congoensis, which have shorter glumes,
and female-fertile spikelets in the lowest whorl.

Description: Goossens 1938 (182), Stapf 1898-1900
(584), Chippindall 1955 (220), Clayton et al. 1970-1982
(365). Voucher: Wedermann & Oberdieck 2188, Davidse
6790. PRECIS code 9902830-00600.

Sporobolus congoensis Franch.

(=S. eylesii Stent &

Rattray) 2.

Perennial; rhizomatous; 380-
920 mm tall. Leaf blades 60-220
mm long; 3-10 mm wide. Spike-
lets 3.0-4. 5 mm long. Leaf blades
ciliate on the margins; panicle
100-200 mm long, whorled,
with fewer than 10 whorls; spikelets loosely clustered on
the upper two thirds of the branches; lower glume 1 .7^4.0
mm long, 1/2-3/4 the spikelet length; upper glume longer
than spikelet.

Flowering November to January. Shallow rocky soil on
sandstone and quartzite. Infrequent. Biome: Grassland.
Tropical east Africa. Similar to S. sanguineus, which has a
larger panicle with more than 10 whorls and shorter lower
glumes, S. mauritianus, with glabrous or scabrid leaf blade
margins, and to S. centrifugus, which has a longer lower
glume and sterile spikelets in the lowest whorl.

Description: Chippindall & Crook 1976 (1 14), Chippin-
dall 1955 (222), Clayton et al. 1970-1982 (365). Voucher:
Acocks & Hafstrom 54. PRECIS code 9902830-00650.

Sporobolus conrathii Chiov.

Perennial; tufted; 200-480
mm tall. Leaf blades 100-180
mm long; 1.0-1. 5 mm wide.

Spikelets 1.5-1. 9 mm long. Plant
base fibrous; panicle dichoto-
mously branched with stiff long
hairs in the axils; glumes about e-
qual, 1/3-1/2 the spikelet length.

Flowering December to
March. Shallow soil on rocky slopes or outcrops. Locally
common. Biome: Savanna and Grassland. Endemic. Similar
to S. welwitschii, which has a panicle with very few hairs
in some of the axils, unequal glumes and a longer upper

glume, and to S. subtilis, which has a rachilla extention and
lacks a fibrous base.

Description: Goossens 1938 (188), Chippindall 1955
(212). Voucher: Du Toil 80. PRECIS code 9902830-00700.

Fig. 198. Sporobolus centrifugus

308

Sporobolus consimilis Fresen.

Sporobolus engleri Pilg.

(=S. robust us sensu
Chippind., non Kunth) 2.

Vleigras.

Robust and reed-like perenni-
al; rhizomatous; 880-1600 mm
tall. Leaf blades 300-600 mm
long; 6-10 mm wide. Spikelets
1.7-2. 5 mm long. Panicle not whorled, linear, more than
five times longer than wide; lower and upper glumes more
or less as long as the spikelet; glumes keeled, keel scabrid.

Flowering November to May. On sand or turf soils in
river beds, on sand banks and near brackish springs. Com-
mon. Biome: Savanna, Grassland, and Nama-Karoo.
Tropical Africa to Somalia and Chad. Distinguished from
other species which have linear panicles with non-whorled
branches by its reed-like appearance and glumes that are
keeled and about the length of the spikelet.

Description: Launert 1970 ( 160: 1 82), Chippindall 1955
(225), Clayton et al. 1970-1982 (371). Voucher: De Winter
& Leistner 5804. PRECIS code 9902830-00800.

Sporobolus coromandelianus (Retz.) Kunth

(=S. argutus (Nees) Kunth)

2; ( =S . pyramidatus sensu
Chippind., non (Lam.)

Hitchc.) 2.

Small dropseed.

Annual; tufted; 100-340 m
tall. Leaf blades 20-100 mm

long; 2-4 mm wide. Spikelets 1.0-1. 5 mm long. Panicle
with lowest branches in a single whorl; central axis and
branches with viscid patches more than three times longer
than wide; lower glume oblong, 0. 1-0.5 mm long, less than
1/3 the spikelet length; upper glume as long as spikelet.

Flowering February to April. In or near brackish pans,
usually on fine clayey soil. Locally common. Biome: Sa-
vanna, Grassland, and Nama-Karoo. Tropical Africa to
India. Similar to S. nitens and S. ludwigii , which are
perennial. Some specimens of S. coromandelianus have
previously been wrongly identified as Sporobolus
cordofanus (Steud.) Coss. andS. uniglumis Stent & Rattray.

Description: Launert 1970 (160:182), Chippindall 1955
(220), Clayton et al. 1970-1982 (363). Voucher: Giess &
Loutit 14146. PRECIS code 9902830-00900.

Sporobolus discosporus Nees

Oortjiesgras, disc dropseed.

Perennial, or annual; shortly
rhizomatous, or tufted; 55-180
mm tall. Leaf blades 10-30 mm
long; 2-5 mm wide. Spikelets
1. 0-1.7 mm long. Leaf blades
short and broad, in a basal rosette,
margins pectinately ciliate; pani-
cle whorled; spikelets pendulous at maturity; grains discoid.

Flowering November to May. Sandy depressions on
Cave sandstone or other exposed bedrock; often on clayey
soil in bare patches or wet areas. Locally common. Biome:
Grassland, and Nama-Karoo. East Africa to Ethiopia. Easily
distinguished by its small size, short, wide, pectinately
ciliate leaf blades, pendulous spikelets and discoid grains.

Description: Goossens 1938 (218), Stapf 1898-1900
(582), Chippindall 1955 (219), Clayton et al. 1970-1982
(358). Illustration: Chippindall 1955 (fig. 194). Voucher:
Smook & Gibbs Russell 2341. PRECIS code 9902830-
01000.

Fig. 199. PI- 182.

Annual; tufted; 120-600 mm
tall. Leaf blades 40-150 mm
long; 2-5 mm wide. Spikelets
1.0-1. 5 mm long. Leaf blades
flat; culms 1-2 mm in diameter;
panicle dichotomously branched,
without hairs in the axils; spike-
lets solitary at the branchlet tips.

Flowering March to May.

Deep sand on dunes and in dry riverbeds, also on rocky soil
and often in shady places. Locally common. Biome: Savan-
na, Nama-Karoo, and Desert. Endemic. Very closely related
to S. nebulosus, which is perennial and has setaceous leaf
blades, also similar to S. festivus and S. stapfianus, which
have the fibrous remains of old leaf sheaths at the base.

Description: Launert 1970 (160:182). Voucher: Van
Vuuren & Giess 1167. PRECIS code 9902830-01100.

Sporobolus festivus A. Rich.

(=S. festivus A. Rich. var.
fibrosus Stent) 2.

Rooigras.

Perennial; tufted: 100-550

mm tall. Leaf blades 20-70 mm
long; 1-2 mm wide. Spikelets
0.8-1. 5 mm long. Old leaf
sheaths splitting into fibres, lacking woolly hairs between
them; panicle dichotomously branched without hairs in the
axils.

Flowering December to May. On exposed bedrock in
sandfilled depressions, also in vleis, on pans edges and in
mopane woodland. Locally common to common. Biome:
Savanna, Grassland, and Nama-Karoo. Tropical east Africa
to Mauritania and Somalia. Pasture (food for warthog).
Similar to S. stapfianus, which has dense woolly hairs
between the basal fibres, S. engleri , which is annual, and
5. nebulosus, which lacks fibres at the base.

Fig. 199. Sporobolus discosporus

309

Description: Goossens 1938 (195), Chippindall & Crook
1976 (113), Launert 1970 (160:183), Stapf 1898-1900
(582), Chippindall 1955 (213), Clayton et al. 1970-1982
(384). Voucher: Story 6236. PRECIS code 9902830-01300.

Sporobolus fimbriatus (Trin.) Nees

(=S. fimbriatus (Trin.) Nees
var. latifolius Stent) 2.

Blousaadgras, dropseed.

Perennial; densely tufted and
rhizomatous (rhizome character-
istically oblique); 240-1600 mm
tall. Leaf blades to 300 mm long;

2^1 mm wide. Spikelets 1.4-2. 2 mm long. Panicle fairly
dense, open, branches numerous, not whorled and not
spreading more than 60 degrees; lower glume 1/4-3/4 the
spikelet length; upper glume about 2/3 the spikelet length.

Flowering December to May. Sandy well-drained loam
near water, often in disturbed areas or in shady spots. Com-
mon. Biome: Fynbos, Savanna, Grassland, and Nama-
Karoo. Tropical east Africa to Sudan and Somali. Food and
drink (seeds pulverised for porridge in times of famine), or
chemicals (hydrocyanic acid in wilted plants). See the
comment as S. africanus. Two varieties have previously
been recognized on leaf width, but this distinction is not
clear and the varieties therefore are not upheld.

Description: Goossens 1938 (205), Launert 1970
(160:183), Stapf 1898-1900 (585), Chippindall 1955 (224),
Clayton et al. 1970-1982 (377). Illustration: Chippindall
1955 (fig. 197), Clayton et al. 1970-1982 (fig. 101).
Voucher: Smook 2779. PRECIS code 9902830-01400.

Sporobolus fourcadii Stent

Perennial; rhizomatous and
tufted; 250-400(-700) mm tall.

Leaf blades 80-350 mm long; 4-7
mm wide. Spikelets 2. 0-2. 6 mm
long. Panicle much longer than
wide, branches not whorled,
spreading almost horizontally at
maturity; lower glume 1/3, upper
glume 2/3 the spikelet length.

Flowering November to March. On the edge of
floodplains or on forest margins. Locally common. Biome:
Savanna. Possibly endemic. Distinguished from S.
fimbriatus, S. pyramidalis,S. natalensis and 5. africanus by
the key characters.

Description: Goossens 1938 (191), Stent 1927 Bothalia
2 (269), Chippindall 1955 (223). Voucher: Giffen 658.
PRECIS code 9902830-01600.

Sporobolus ioclados (Trin.) Nees

similar to S. ludwigii, which is mat-forming, has a
horizontally creeping rhizome and shorter leaf blades, also
similar to S. rangei, which has shorter and narrower leaf
blades and is seldom stoloniferous.

Description: Chippindall & Crook 1976(115), Goossens
1938 (198), Launert 1970 (160:183), Stapf 1898-1900
(583), Chippindall 1955 (216), Clayton et al. 1970-1982
(367). Illustration: Chippindall 1955 (fig. 192). Voucher:
De Winter & Codd 340. PRECIS code 9902830-01700.

Sporobolus kentrophyllus (K. Schum.) Clayton

Tussocky perennial; rhizomat-
ous and stoloniferous; 130-950
mm tall. Leaf blades 30-180 mm
long; 4-8 mm wide. Spikelets
1.5-2. 5 mm long. Leaf blades
rounded at the base; panicle
partly whorled (at least the lowest
branches); lower glume 1/4 the
spikelet length; upper glume
2/3— 3/4 the spikelet length.

Flowering January to March. Powdery loam or
calcareous soil on lake beds, also in moist depressions on
old lands. Rare. Biome: Savanna. Tropical east Africa to
Somalia. The two specimens recorded for our area do not
match the type specimen of S. verdcourtii (a synonym)
satisfactorily although they match other cited specimens
from east Africa.

Description: Clayton et al. 1970-1982 (369). Voucher:
Killick & Leistner 3422. PRECIS code 9902830-01800.

Sporobolus ludwigii Hochst.

Brakvleigras.

Mat-forming perennial; rhizo-
matous (rhizome long, creeping
and profusely branched);

100-450 mm tall. Leaf blades
10-30 mm long; 2-3 mm wide.

Spikelets 1. 5-2.0 mm long. Leaf
blade margins not ciliate; panicle
whorled; lower glume 1/3 the spikelet length; upper glume
as long as the spikelet.

Flowering January to May. Fine, damp calcareous soils
in vleis or near pans. Locally common. Biome: Savanna and
Grassland. Possibly endemic. Superficially similar to S.
nitens, which has smaller spikelets, S. coromandelianus ,
which is annual, and S. ioclados, which has longer leaf
blades and is usually stoloniferous.

Description: Goossens 1938 (201), Stapf 1898-1900
(583), Chippindall 1955 (216). Illustration: Chippindall
1955 (fig. 191). Voucher: Esterhuysen 2018. PRECIS code
9902830-02000.

(=S. ioclados (Trin.) Nees
var. usitatus (Stent) Chippind.)

2; (=S. marginatus Hochst. ex
A. Rich.) 2; (=S. smutsii Stent)

2; (=S. usitatus Stent) 2.

Pan dropseed.

Perennial (often mat-forming); rhizomatous and stolon-
iferous; 250-1000 mm tall. Leaf blades 20-300 mm long;
2-12 mm wide. Spikelets 1.5-2. 5 mm long. Basal leaf
sheaths papery; culms geniculate; leaf blades flat; panicle
whorled; lower glume 1/4— 1/3 the spikelet length; upper
glume as long as, or longer than the spikelet.

Flowering January to April. A variety of soil types,
including black turf and sand, often in disturbed places.
Common. Biome: Savanna, Grassland, and Nama-Karoo.
Tropical Africa to India. Well eaten natural pasture. Panicle

Sporobolus macranthelus Chiov.

Robust perennial; rhizomat-
ous; 1050-1700 mm tall. Leaf
blades to 450 mm long; 4-7 mm
wide. Spikelets 1.6-2.4 mm long.

Culms 3-7 mm in diameter; pani-
cle lax, much branched, linear
and not whorled, 350-750 mm
long.

Flowering January to Feb-
ruary. Often in the shade of riverine woodland or on the
edge of floodplains on fertile loam. Rare. Biome: Savanna.
Tropical east Africa to Sudan and Somalia. Distinguished
from S. africanus, S. fourcadii, S. fimbriatus and S
natalensis by its robust habit and large panicle.

Description: Clayton et al. 1970-1982 (380). Voucher.
Smith 803. PRECIS code 9902830-02050.

310

Sporobolus mauritianus (Steud.) Dur. & Schinz

(=S. artus Stent) 2.

Perennial; densely tufted and
rhizomatous; 170-430 mm tall.

Leaf blades 50-250 mm long; 1-6
mm wide (often filiform). Spike-
lets 2.5-5. 0 mm long. Basal
sheaths papery, not glossy; pani-
cle with lowest branches whorled,
primary branches short and contracted; lower glume
2/3— 3/4 the spikelet length; upper glume longer than the
spikelet.

Flowering October to January. Poorly drained soil in
marshy areas or on coastal sandflats, fairly frequent in
sourveld. Infrequent. Biome: Savanna and Grassland.
Tropical Africa, Madagascar and Mauritius. Similar to S.
congoensis and S. sanguineus, which have flexuous hairs
longer than 0.5 mm on the leaf margins, and S. centrifugus,
which has sterile spikelets in the lowest whorl and longer
lower glumes.

Description: Goossens 1938 (181), Chippindall 1955
(221), Clayton et al. 1970-1982 (366). Voucher: Schrire
613. PRECIS code 9902830-02150.

Sporobolus molleri Hack.

Annual; loosely tufted;

110-360 mm tall. Leaf blades
20-250 mm long; 1-5 mm wide.

Spikelets 1. 7-2.0 mm long. Pani-
cle linear, more than five times
longer than wide, not whorled;
lemma narrow and needle-like,
notably longer (to 1.5 times) than
the rest of the spikelet.

Flowering February to April. Well-drained soil on
abandoned or cultivated lands. T^are. Biome: Savanna.
Tropical Africa south of the Congo River. Weed (easily
controlled by cultivation). A single collection is known
from near Tzaneen. The long narrow lemma of this species
is unique for the genus.

Description: Chippindall & Crook 1976 (111), Clayton
et al. 1970-1982 (372). Voucher: Retief 33. PRECIS code
9902830-02160.

Sporobolus natalensis (Steud.) Dur. & Schinz

Perennial; tufted; 550-1450
mm tall. Leaf blades 250-500
mm long; 2-4 mm wide. Spike-
lets 1.6-2. 3 mm long. Panicle
fairly dense, branches numerous,
not whorled, contracted or some-
times spreading; lower glume
1/3— 1/2 the spikelet length; upper
glume 1/2-2/3 the spikelet length.

Flowering December to April. Sandy well-drained soil
near water or in woodlands, often in disturbed places. Infre-
quent. Biome: Savanna and Grassland. Tropical east Africa
to Ethiopia. See the comment at S. africatius.

Description: Launert 1970 (160:184), Clayton et al.
1970-1982 (374). Voucher: Liebenberg 8661. PRECIS
code 9902830-02170.

Sporobolus nebulosus Hack.

Perennial; densely tufted and
rhizomatous; 70-300 mm tall.

Leaf blades filiform and rigid,

15-50 mm long; 0. 3-1.0 mm
wide. Spikelets 0.8-1 .4 mm long.

Old dead leaves and sheaths form
a hard, dense cushion below the
new growth; panicle dichoto-
mously branched, without hairs in
the axils; spikelets solitary at the branchlet tips.

Flowering January to May. In depressions or moist
places in deep sand. Locally common. Biome: Savanna,
Nama-Karoo, and Desert. Endemic. Closely related to S.
engleri, which is annual, and S. festivus and S. stapfianus
in which the old leaf sheaths split into fibres.

Description: Goossens 1938 (220), Launert 1970
(160:184), Chippindall 1955 (213). Illustration: Chippindall
1955 (fig. 188). Voucher: Volk 58. PRECIS code
9902830-02200.

Sporobolus nervosus Hochst.

(=S. lampranthus Pilg.) 2;

(=5. sladenianus Bol. f.) 2.

Perennial; rhizomatous (rhi-
zomes compact with short
intemodes, creeping horizontally
at ground level); 180-530 mm
tall. Leaf blades 40-100 mm
long; 1-3 mm wide. Spikelets
1.7-2. 4 mm long. Leaves fine and curly; panicle not
whorled, pyramidal to ovate, not more than three times
longer than wide; spikelets loosely grouped at the branchlet
tips; glumes not keeled, lower glume 1/2-2/3 the spikelet
length, upper glume 3/4— 4/5 the spikelet length.

Flowering February to May. On flats or in moist
depressions in sandy red soil, limestone or shale. Locally
common. Biome: Savanna and Nama-Karoo. Tropical east
Africa to Arabia.

Description: Launert 1970 (160:184), Chippindall 1955
(215), Clayton et al. 1970-1982 (380). Illustration: Chip-
pindall 1955 (fig. 190), Clayton et al. 1970-1982 (fig. 102).
Voucher: De Winter & Hardy 8014. PRECIS code
9902830-02250.

Sporobolus nitens Stent

Fig. 200.

Perennial; rhizomatous and
stoloniferous; 190-520 mm tall.

Leaf blades 35-90 mm long; 4-8
mm wide. Spikelets 1.2-1. 5 mm
long. Leaf blade margins wavy
and scabrid or ciliate; panicle
with lowest branches whorled;
spikelets clustered on upper half
of primary branches; lower glume
1/2 the spikelet length; upper glume as long as spikelet,
acuminate, prominently keeled, keel scabrid.

Flowering November to April. In bare patches and in
overgrazed veld, also in gardens and other disturbed places.
Common. Biome: Savanna and Grassland. Possibly
endemic. Superficially similar to S. coromandelianus,
which is annual, and toS. ludwigii, which has larger spike-
lets.

Description: Goossens 1938 (197), Chippindall 1955
(218). Illustration: Chippindall 1955 (fig. 193). Voucher:
De Winter & Codd 510. PRECIS code 9902830-02300.

Sporobolus panicoides A. Rich.

Famine grass.

Annual; loosely tufted (erect
and slender); 190-960 mm tall.

Leaf blades 50-300 mm long; 2-6
mm wide. Spikelets 2. 0-3. 3 mm
long. Inflorescence branches
whorled; spikelets large, sparse,
sterile in the lowest whorl; grains
almost spherical, bright brown or orange coloured, 1.2- 1.9
mm in diameter.

Flowering December to May. Sandy, rocky areas on
steep slopes or flats, most often on roadsides or in other
disturbed areas, sometimes in the shade. Locally common.
Biome: Savanna. Tropical east Africa to Ethiopia. Food and
drink (grains used as food in times of famine).

311

Fig. 200. Sporobolus nitens

Characterized by the few large spikelets and brightly
coloured grains.

Description: Goossens 1938 (217), Chippindall & Crook
1976 (112), Launert 1970 (160:185), Chippindall 1955
(223), Clayton et al. 1970-1982 (359). Illustration: Chip-
pindall 1955 (fig. 196), Clayton etal. 1970-1982 (fig. 100).
Voucher: Fourie 2541. PRECIS code 9902830-02400.

Sporobolus pectinatus Hack.

Fringed dropseed, kamme-
tjiesgras.

Perennial; rhizomatous (older
plants with long horizontally
creeping rhizomes); 240-740 mm
tall. Leaf blades 50-300 mm
long; 5-8 mm wide. Spikelets
3. 0-3. 7 mm long. Leaf blades
pectinately ciliate; panicle whorled; spikelets very densely
clustered on the upper third of the branches, leaving the
lower part bare; lower glume 1/2 the spikelet length; upper
glume as long as spikelet.

Flowering November to February. Shallow rocky soil on
outcrops or quartzite ridges. Infrequent. Biome: Grassland.
Endemic. Characterized by the spikelet arrangement in the
panicle.

Description: Goossens 1 938 ( 1 86), Chippindall & Crook
1976 (114), Chippindall 1955 (221). Illustration: Chippin-
dall 1955 (fig. 195). Voucher: Louw 3924. PRECIS code
9902830-02500.

Sporobolus pellucidus Hochst.

Perennial; densely tufted and
rhizomatous; 150-640 mm tall.

Leafblades40-150mm long; 1-2
mm wide. Spikelets 1. 7-2.0 mm
long. Leaf sheaths splitting into
fibres with age; leaves filiform
and forming a cushion at the base;
panicle 60-200 mm long, not
whorled, linear, more than five
times longer than wide.

Flowering January to March. Calcareous soils. Rare.
Biome: Savanna. Tropical east Africa to Ethiopia.

Description: Clayton et al. 1970-1982 (374). Voucher:
Giess & Loutit 14102. PRECIS code 9902830-02600.

Sporobolus pyramidalis Beauv.

Catstail grass, vleigras,
taaipol.

Perennial; densely tufted and
rhizomatous; 700-1600 mm tall.

Leaf blades 100-500 mm long;

3-10 mm wide. Spikelets 1. 7-2.0
mm long. Panicle linear, more
than five times longer than wide;
both glumes less than 1/3 the spikelet length.

Flowering November to May. Vleis, watercourses,
periodically flooded areas or near dams on sandy soil or
heavy clay. Common. Biome: Savanna and Grassland.
Tropical Africa, Madagascar, Mauritius and Yemen. Tough
and very unpalatable, erosion control (trampled areas), or
indicator (of overgrazing), or weed (in pastures). See
comment at S. africanus. Vegetatively very similar to the
other ‘taaipol’, Eragrostis plana , which has several florets
in each spikelet.

Description: Goossens 1938 (210), Launert 1970
(160:185), Chippindall 1955 (224), Clayton et al.
1970-1982 (373). Voucher: Smook 5043. PRECIS code
9902830-02700.

312

Sporobolus rangei Pilg.

Perennial (usually robust);
rhizomatous (rhizome usually
horizontal), or stoloniferous
(seldom); 350^460 mm tall. Leaf
blades 30-140 mm long;
setaceous or to 3 mm wide.

Spikelets 1.7-2. 7 mm long. Basal
leaf sheaths hard, glossy and
brittle; culms erect; panicle with
lowest branches whorled; lower glume less than 1/2 the
spikelet length; upper glume as long as the spikelet.

Flowering November to March. Calcareous sandy soil
in shallow pans or near watercourses. Infrequent. Biome:
Savanna and Nama-Karoo. Tropical east Africa. Similar to
S. ioclados, which has geniculate culms and wider and
longer leaf blades.

Description: Launert 1970 (160:185), Chippindall 1955
(217), Clayton et al. 1970-1982 (368). Voucher:

Merxmuller 1051. PRECIS code 9902830-02800.

Sporobolus salsus Mez

Perennial; rhizomatous; 270-
700 mm tall. Leaf blades 45-150
mm long; 1^4 mm wide. Spike-
lets 2. 0-2. 9 mm long. Panicle di-
chotomously branched, without
long hairs in the axils; spikelets
large and solitary at the branchlet
tips.

Flowering January to Septem-
ber. Seasonally flooded brackish pans and near hot springs
or rivers. Locally common. Biome: Savanna and Nama-
Karoo. Possibly endemic. Very similar to S. tenellus and S.
acinifolius, which have smaller spikelets.

Description: Feddes Rep. 1921 17 (296). Voucher: De
Winter & Codd 339. PRECIS code 9902830-02900.

Flowering December to June. Grassy vleis, brackish
sandy soil to very saline soils on pans or in river beds. Lo-
cally common. Biome: Savanna and Succulent Karoo.
Tropical east Africa and drier regions of Africa, from the
Mediterranean coast to India. Very similar to S. albicans
and S. bechuanicus, which have wider panicles, and S.
virginicus, which has longer glumes and a wider panicle.

Description: Goossens 1938 (209), Launert 1970
(160:185), Chippindall 1955 (226), Clayton et al.
1970-1982 (369). Illustration: Chippindall 1955 (fig. 199
at fig. 187). Voucher: Codd & Dyer 3804. PRECIS code
9902830-03200.

Sporobolus stapfianus Gand.

Fynblousaadgras, fibrous
dropseed.

Perennial; tufted; 150-550
mm tall. Leaf blades 30-150 mm
long; 1-2 mm wide. Spikelets
1. 4-2.1 mm long. Old leaf
sheaths fibrous with a mass of
woolly hairs between the fibres;
panicle dichotomously branched, without long hairs in the
axils.

Flowering October to March. Sandy well-drained to very
compacted soils on rocky outcrops or near streams. Com-
mon. Biome: Savanna, Grassland, and Nama-Karoo.
Tropical east Africa to Nigeria and Ethiopia, also in
Madagascar. Very closely related to S.festivus, which lacks
woolly hairs between the fibres, S. engleri, which is annual,
andS. nebulosus, which does not have a fibrous base.

Description: Chippindall & Crook 1976 (113), Chippin-
dall 1955 (213), Clayton et al. 1970-1982 (384).
Illustration: Chippindall 1955 (fig. 187). Voucher: Burtt-
Davy 2683. PRECIS code 9902830-03300.

Sporobolus stolzii Mez

Sporobolus sanguineus Rendle

(-S. rhodesiensis Stent &

Rattray) 2.

Perennial; loosely tufted and
rhizomatous; 600-1020 mm tall.

Leaf blades 100-400 mm long;

.1.5-6. 0 mm wide. Spikelets
2.0-3. 5 mm long. Leaf blade mar-
gins ciliate; panicle 200-430 mm
long, linear to lanceolate, whorled, with more than 10
whorls; lower glume less than 1/2 the spikelet length; upper
glume longer than the spikelet.

Flowering November to April. Stony hillslopes and
highlying grasslands. Infrequent. Biome: Savanna. Tropical
Africa. Similar to S. congoensis, which has a shorter,
pyramidal panicle with fewer than 10 whorls, S.
mauritianus, which has glabrous or scabrid leaf blade
margins, and S. centrifugus, which has sterile spikelets in
the lowest whorl and glumes longer.

Description: Chippindall & Crook 1976 (116), Chippin-
dall 1955 (222), Clayton et al. 1970-1982 (364). Voucher:
Van der Schijff 4060. PRECIS code 9902830-03000.

Sporobolus spicatus (Vahl) Kunth

Mat-forming, wiry perennial;
rhizomatous and stoloniferous;

250-1000 mm tall. Leaf blades
rigid and spiny-tipped, 20-300
mm long; 1^4 mm wide. Spike-
lets 1.4-2. 8 mm long. Panicle
spike-like, not whorled, less than
5 mm wide; spikelets very
densely clustered around the
central axis and hiding it completely.

Annual; tufted (erect);

290-950 mm tall. Leaf blades
10-60 mm long; 2-6 mm wide.

Spikelets 0.9-1. 6 mm long. Leaf
margins pectinately ciliate; in-
florescence branches in whorls,
rachis and branches with
abundant round to ovate viscid
patches; lower glume linear-
lanceolate, longer than 0.5 mm and at least 1/2 the length
of the spikelet; upper glume slightly shorter than the spike-
let.

Flowering January to May. Amongst trees on sandy soil.
Rare. Biome: Savanna. Tropical Africa to Senegal and
Ethiopia. One specimen at PRE provided seed from which
the remainder of our specimens were cultivated.

Description: Chippindall & Crook 1976 (109), Clayton
et al. 1970-1982 (358). Voucher: De Winter 9261. PRECIS
code 9902830-03350.

Sporobolus subtilis Kunth

Fig. 201.

Misty dropseed.

Perennial; rhizomatous (rhi-
zome slender and creeping), or
stoloniferous (sometimes), or
tufted; 320-600 mm tall. Leaf
blades 40-150 mm long. Spike-
lets 1. 5-3.0 mm long. Culms wi-
ry; few basal leaves; panicle
dichotomously branched with long stiff hairs in the axils;
rachilla extending into a rudimentary floret between the
upper glume and the palea.

Flowering November to January. Shallow sandy soil in
moist areas. Locally common (often in pure stands). Biome:
Savanna and Grassland. Tropical Africa to Sierra Leone,

313

also in Madagascar. The rachilla extention in this species
is unique for the genus. Similar to S. conrathii, which has
a fibrous base, and S. welwitschii, which has a panicle with
few hairs in the axils.

Description: Goossens 1938 (221), Stapf 1898-1900
(588), Chippindall 1955 (212), Clayton et al. 1970-1982
(386). Illustration: Chippindall 1955 (fig. 186), Clayton et
al. 1970-1982 (fig. 103). Voucher: Huntley 700. PRECIS
code 9902830-03400.

Sporobolus tenellus (Spreng.) Kunth

Pankweek.

Mat-forming perennial; rhizo-
matous (rhizome long and
profusely branched); 60-280 mm
tall. Leaf blades 5-35 mm long;

1-3 mm wide. Spikelets 1.5-1. 8
mm long. Leaf blades short and
rounded at the tips, mostly basal;
culms usually one-noded; panicle dichotomously branched
with spikelets solitary at the branch tips.

Llowering November to April. Shallow soils at pan
edges or in moist depressions. Locally common. Biome: Sa-
vanna and Nama-Karoo. Possibly endemic. Very similar to
S. salsus , which has larger spikelets, and S. acinifolius ,
which has leaf blades 40-120 mm long and tapering to a
fine point.

Description: Goossens 1938 (194), Launert 1970
(160:185), Stapf 1898-1900 (580), Chippindall 1955 (214).
Voucher: Acocks 12511. PRECIS code 9902830-03500.

Pig. 201. Sporobolus subtilis

Sporobolus virginicus (L.) Kunth

Seaside rush grass.

Mat-forming perennial; sto-
loniferous, or rhizomatous (rhi-
zomes extensively creeping);

110-770 mm tall. Leaf blades
50-150 mm long; 1-7 mm wide.

Spikelets 1.7-2. 5 mm long. Leaf
blades convolute and pungent;
panicle spike-like, branches not whorled; lower glume 3/4
the spikelet length; upper glume as long or slightly longer
than the spikelet.

Llowering October to April. On dunes, beaches and
along tidal streams on sand. Mostly along the coast but also
inland at saline water edges. Common (along coasts).
Biome: Pynbos, Savanna, Succulent Karoo, and Desert.
Tropical and subtropical regions worldwide. Erosion con-
trol (on sand dunes). Specimens of this species can vary
from soft, very fine and delicate plants to large and robust
plants. Similar to S. albicans, S. bechuanicus and S.
spicatus, which all have shorter glumes.

Description; Goossens 1938 (207), Launert 1970
(160:186), Chippindall 1955 (227), Clayton et al.
1970-1982 (370). Illustration: Chippindall 1955 (fig. 200).
Voucher: Strey 7325. PRECIS code 9902830-03600.

Sporobolus welwitschii Rendle

(=.S\ macrothrix Pilg.) 2; {=S.
baumianus Pilg.) 3.

Perennial; rhizomatous and
tufted; 600-700 mm tall. Leaf
blades 40-80 mm long; mostly
filiform, but to 2 mm wide.

Spikelets 0.8-2. 1 mm long.

Plants wiry with few leaves;
leaves mostly cauline; panicle dichotomously branched
with a few long stiff hairs in some of the axils; spikelets
solitary on the tips of delicate branches; glumes unequal,
lower glume 1/2, upper glume 2/3 the spikelet length.

Flowering December to February. Brackish sandy loam
on the edge of pans or woodlands. Rare. Locally common.
Biome: Savanna. Similar to S. conrathii and S. subtilis,
which have panicles with many hairs in almost all the axils
and glumes more or less equal and 1/2 the spikelet length.

Description: Launert 1970 (160:186), Chippindall 1955
(212). Voucher: Rogers 25106. PRECIS code 9902830-
03700.

Sporobolus sp. (=Smook 3429)

Mat-forming perennial; rhizo-
matous (rhizome long and deeply
buried); 130-320 mm tall. Leaf
blades 20-100 mm long; 3.0- 4.5
mm wide (mostly inrolled).

Spikelets 1.8-2. 9 mm long.

Leaves rigid; panicle ovate, not
whorled, less than three times
longer than wide; glumes usually
keeled along the whole length or at least at the tip, keel
scabrid; lower glume 2/3 to slightly shorter than the spike-
let; upper glume 3/4 to slightly longer than spikelet.

Flowering November to April. Brackish soils in or near
salt pans. Locally common (brackish soils). Biome: Savan-
na. Endemic. Although this group of specimens is habitat
specific and can easily be distinguished from other
Sporobolus species, its status is a little uncertain. Spikelet
characters are very variable within a single panicle and of
little use in identification, possibly indicating a hybrid

origin.

Voucher: Smook 3429.

PRECIS code 9902830-99999.

314

Stenotaphrum Trin.

Diastemenanthe Steud., Ophiurinella Desv.

Perennial; long-rhizomatous, or long-stoloniferous, or
caespitose. Culms 100-600 mm high; herbaceous; branched
above. Leaf blades lanceolate to elliptic; flat, or folded
(when young). Ligule a fringed membrane. Plants bisexual,
with bisexual spikelets. The spikelets all alike in sexuality.

Inflorescence of spike-like main branches, or a false
spike, with clusters of spikelets on reduced axes, or panicu-
late (spikelets 1 to several, in very short spike -like racemes
embedded in hollows of the common axis , or in longer
racemes closely oppressed to it); spatheate (the small
racemes subtended/enclosed by spathes which are laterally
adnate to the rachis), or espatheate. Spikelet-bearing axes
very much reduced (or coalesced with the main axis); disar-
ticulating; falling entire (the free racemes falling with the
joint of the main axis), or disarticulating at the joints (when
the ‘spikelet bearing unit’ consists of a coalesced main axis
and branches).

Spikelets abaxial; compressed dorsiventrally; falling
with the glumes. Glumes two; very unequal; awnless; very
dissimilar (lower minute, scale-like, upper large,
substantial), or similar (both small, scale-like). Proximal in-
complete florets 1; paleate, or epaleate, palea when present
fully developed; male, or sterile (rarely). Proximal lemmas
7-9 nerved.

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes (papery to subcoriaceous); smooth to striate; not

Fig. 202. Stenotaphrum secundatum

becoming indurated; hairless; having the margins lying flat
and exposed on the palea; with a clear germination flap; 3-5
nerved; entire; awnless. Palea present; relatively long. Lod-
icules 2; fleshy. Stamens 3. Ovary glabrous. Fruit small,
ellipsoid; hilum short; embryo large.

Photosynthetic pathway. C4; XyMS- PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number,* = 9. Panicoideae; Panicodae; Paniceae. 7 species.
Tropical and subtropical. Mesophytic; in open habitats
(usually maritime); maritime-arenicolous to halophytic, or
glycophytic. Namibia, Transvaal, Swaziland, Natal, and
Cape Province. 2 indigenous species.

References. 1 . Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by H.M. Anderson.

1(0). Axis of inflorescence almost cylindrical, without
notches; spikelets 1-3, more of less embedded in

the axis S. secundatum

Axis of inflorescence flat on one surface, with widely
spaced notches along the edges; spikelets 2-7, each
fitting into a shallow cavity S. dimidiatum

Stenotaphrum dimidiatum (L.) Brongn.

Perennial; extensively stolon-
iferous (forming dense swards);

60-400 mm tall. Leaf blades
50-80 mm long (keeled); 8-12
mm wide. Spikelets 4-5 mm
long; 1-2 mm wide. Leaf-sheaths
strongly flattened, folded, often
grouped in fan-shaped arrange-
ments; the spikelike raceme
compact and compressed, central axis thick, swollen, flat
on one surface, hollowed out on the other, each cavity con-
taining 2-7 spikelets in short racemes borne alternately on
either side of a wavy midrib, the edge of each cavity with
a broad acute tooth.

Flowering October to May. A coastal pioneer along
beaches and marshes, by saline and fresh water. Infrequent.
Pan-tropical and in warm temperate areas. Pasture and
ornamental (lawns).

Description: Chippindall & Crook 1976 (187). Voucher:
Eglington 34412. PRECIS code 9901080-00100.

Stenotaphrum secundatum (Walt.) Kuntze

Fig. 202. PI. 184.

Buffalo grass.

Perennial, extensively stolon-
iferous (forming dense swards);

60-400 mm tall. Leaf blades
50-150 mm long (keeled); 4—10
mm wide. Spikelets 4—5 mm
long; 1-2 mm wide. Leaf sheaths
strongly flattened, folded and of-
ten grouped in fan-shaped arrangements; the spike-like ra-
ceme compact and cylindrical, central axis thick, swollen,
flat on one surface, deeply hollowed out on the other, each
cavity usually containing one spikelet (sometimes 2-3)
borne alternately on either side of a wavy midrib.

Flowering October to May. A coastal pioneer along,
beaches and marshes, by saline and fresh water. Locally
common. Pan-tropical and in warm temperate areas. Pasture
and ornamental (lawns). The Cape deme (a sterile triploid
clone) is widely cultivated.

Description: Chippindall 1955 (367). Illustration: Chip-
pindall 1955 (fig. 316). Voucher: Smook 3130. PRECIS
code 9901080-00200.

315

Stereochlaena Hackel

Chloridion Stapf.

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 600-1500 mm high; herbaceous; branched above, or
unbranched above. Leaf blades linear to linear-lanceolate;
flat. Ligule a fringed membrane. Plants bisexual, with
bisexual spikelets.

Inflorescence of spike-like main branches (slender
spike-like racemes ); digitate or subdigitate\ espatheate.
Spikelet-bearing axes persistent.

Spikelets in pairs; biseriate; consistently in ‘long-and-
short’ combinations (but homogamous). Spikelets 2-4.5
mm long; abaxial; compressed dorsiventrally; falling with
the glumes. Glumes one or two per spikelet; minute, or rela-
tively large (G1 is minute or absent, while G2 may be
minute to almost as long as the spikelet); when both present
very unequal; awned (G2 only, sometimes), or awnless;
when both present very dissimilar. Lower glume when
present 0 nerved. Proximal incomplete florets 1 ; paleate, or
epaleate (?), palea when present reduced; sterile. Proximal
lemmas awned ( the terminal awn from 3-30 mm long).

Female-fertile florets 1. Lemmas decidedly firmer than
the glumes (papery); not becoming indurated (brown);
hairless; having the margins lying flat and exposed on the
palea; with a clear germination flap; 3 nerved (faintly);
entire; awnless (sometimes apiculate). Palea present; rela-
tively long. Stamens 3. Ovary glabrous. Fruit small (about
1.7 mm long), elongate ellipsoid; hilum short (punctiform);
embryo large (about 1/3 the fruit length).

Photosynthetic pathway. C4; XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 5 species. Tropical east Africa. Meso-
phytic; in open habitats (savanna grasslands); glycophytic.
Botswana (?) and Transvaal. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.
Species treatment by G.E. Gibbs Russell.

Stereochlaena cameronii (Stapf) Pilg.

Fig. 203. PI. 185.

Perennial; sometimes stolon-
iferous and tufted; 600-1000
(-1200) mm tall. Leaf blades
80-250 mm long; 2-8 mm wide.

Spikelets 2. 0-3. 5 mm long. Basal
sheaths hairy; lower lemma with
a straight awn 5-20 mm long;
female-fertile floret dark brown
at maturity.

Flowering January to May. Dry sandy grassland. Infre-
quent. Biome: Savanna. To east tropical Africa. Related to
Digitaria , which does not have awned lemmas. The awned
digitate racemes give a superficial resemblance to Chloris,
which has laterally compressed spikelets that fall above the
glumes.

Description: Chippindall 1955 (425), Clayton et al.
1970-1982 (656). Illustration: Chippindall 1955 (fig. 353).
Voucher: Galpin 1 1345. PRECIS code 9900910-00100.

Stiburus Stapf

Sometimes included in Eragrostis Wolf.

Annual ; caespitose. Culms 100-630 mm high; herba-
ceous; unbranched above. Ligule a fringed membrane (very
narrow ), or a fringe of hairs.

Inflorescence paniculate', contracted; elongated-sym-
metrical, spike-like (purplish)', non-digitate; espatheate.
Spikelet-bearing axes persistent.

Spikelets 4 mm long', compressed laterally; disarticu-
lating above the glumes; disarticulating between the florets.
Rachilla prolonged beyond the uppermost female-fertile
floret. Hairy callus present (but minute ). Glumes two; very
unequal, or more or less equal; decidedly shorter than the
adjacent lemmas, or long relative to the adjacent lemmas;
awnless; similar. Upper glume 1 nerved. Incomplete florets
distal to the female-fertile florets, merely underdeveloped;
proximal incomplete florets absent.

Female-fertile florets 1-5. Lemmas similar in texture to
the glumes (thin); without a germination flap; 3 nerved;
entire', mucronate (excurrent into the mucro). Palea present;
relatively long, or conspicuous but relatively short. Lodi-
cules 2; fleshy (tiny); glabrous. Stamens 3 (anthers minute).
Ovary glabrous. Fruit small (about 2 mm); hilum short;
pericarp fused (probably).

Photosynthetic pathway and related features. C4;
XyMS-i- (the ms cells very large, larger than the per cells,
with very thick walls). PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 2 species. Southern Africa.
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 2 indigenous species.

References. 1 . Chippindall. 1 955. Gr. & Past. 2. Phillips.
1982. Kew Bull. 37:133.

Species treatment by M. Koekemoer.

1(0). Glumes and lemmas very densely hairy, dark purple
to black, tips acuminate; panicle 30-90 mm long;
leaves almost always overtopping the
inflorescences; plants flowering February to May

S. alopecuroides

Glumes and lemmas hairy, light purple to yellow, tips
acute; panicle less than 25 mm long; leaves seldom
overtopping the inflorescences; plants flowering
August to December S. conrathii

316

Stiburus alopecuroides (Hack.) Stapf

Perennial; rhizomatous and
tufted; 170-630 mm tall. Leaf
blades 90-360 mm long; 2 mm
wide. Spikelets 2. 7-4.0 mm long.

Inflorescences rarely overtopping
the leaves; glumes and lemmas
very densely hairy, dark purple to
black, tips acuminate.

Flowering February to May.

Open veld, mostly sourveld, at fairly high altitudes, in
fertile soil and wet areas. Locally common. Biome: Savanna
and Grassland. Zimbabwe. Phenologically distinct from S.
conrathii, which flowers from August to December.

Description: Stapf 1898-1900 (697), Chippindall 1955
(186). Illustration: Chippindall 1955 (fig. 161). Voucher:
Mohle 351. PRECIS code 9904000-00100.

Stiburus conrathii Hack.

Perennial; rhizomatous and
tufted; 100-410 mm tall. Leaf
blades 30-100 mm long. Spike-
lets 1. 7-3.0 mm long. Inflores-
cences usually overtopping the
leaves; glumes and lemmas hairy,
light purple to yellow, tips acute.

Flowering August to Decem-
ber. Damp or wet areas in
mountain sourveld. Locally common (to infrequent).
Biome: Grassland (in mountains). Phenologically distinct
from S. alopecuroides, which flowers from February to
May.

Description: Chippindall 1955 (186). Voucher: Rogers
24049. PRECIS code 9904000-00200.

Fig. 204. PI. 186.

Fig. 204. Stiburus alopecuroides

Stipa L.

Achnatherum P. Beauv., Aristella Bertol., Jarava Ruiz
& Pavon, Lasiagrostis, Macrochloa Kunth, Orthoraphium
Nees, Ptilagrostis Griseb., Sparteum P. Beauv, Timouria
Roshev.

Perennial (rarely annual — e.g. S. capensis, S. parvulaf
caespitose. Culms 100-2500 mm high; woody and
persistent (rarely), or herbaceous; branched above, or un-
branched above. Leaf blades linear; flat, or folded, or rolled.
Ligule an unfringed membrane, or a fringed membrane.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets not in distinct ‘long-and-short’ combinations;
3-12 mm long (narrow); compressed laterally; disarticu-
lating above the glumes. Rachilla terminated by a female-
fertile floret. Hairy callus present (long and sharp-pointed,
except in Ptilagrostis). Glumes two; more or less equal;
about equalling the spikelets to much exceeding the
spikelets; awnless, or awned (sometimes aristate); similar.
All florets female-fertile only; proximal incomplete florets
absent.

Female-fertile florets I . Lemmas decidedly firmer than
the glumes (narrow, convolute); hairy, or hairless (rarely);
without a germination flap; 3-7 nerved; entire, or incised
(shortly 2-toothed in Ptilagrostis); awned. Awns /; median;
from the sinus ( Ptilagrostis ), or apical; geniculate (or
sometimes bi-geniculate); hairless, or hairy, or long-
plumose; much shorter than the body of the lemma, to much
longer than the body of the lemma. Palea usually present
(enclosed by the lemma); relatively long (usually), or con-
spicuous but relatively short to very reduced (rarely); indu-
rated (more or less, at least the exposed part)', 2-nerved,
or nerveless (rarely). Lodicules 2 (rarely), or 3; fleshy, or
membranous (stipoid); glabrous. Stamens 3. Ovary
glabrous. Fruit small, or medium sized, or large; fusiform;
hilum long-linear; pericarp fused; embryo small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number,* = 9, 10, 1 1, 12, and 22. Arundinoideae; Stipeae.
300 species. Tropical and temperate. Mesophytic to xero-
phytic. Shade species and in open habitats; glycophytic.
Transvaal, Orange Free State, Natal and Cape Province.
Indigenous species (3), naturalized species (4).

Intergeneric hybrids with Oryzopsis — X Stiporyzopsis
B.L. Johnson & Rogler.

References. 1. De Winter. 1965. Bothalia 8: 212. 2.
Caro. 1966. Kurtziana 3: 79. 3. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Glumes less than 5 mm long; leaf blades tightly
rolled; plants very densely tufted; perennial ....

S. tenuissima

Glumes more than 5 mm long; leaf blades expanded
or rolled; plants loosely to densely tufted; annual

or perennial 2

2(1). Glumes about 15 mm long 3

Glumes 5-10 mm long 4

3(2). Glumes colourless, translucent, shining; lemma
lacking a raised collar around base of awn; plant

annual, to 500 mm tall, usually shorter

S. capensis

Glumes dark purple; lemma with a raised collar
around base of awn; plant perennial, about 1000

mm tall S. neesiana

4(2). Lemma with a conspicuous brush of shining white
hairs 4-5 mm long at upper end; glumes shorter

than floret S. papposa

Lemma shortly hairy all over, lacking a brush of long
hairs at upper end; glumes longer than floret . . 5
5(4). Glumes 9-10 mm long; awns sinuous, to 50 mm long
S. variabilis

317

Glumes 5-7 mm long; awns straight or bent and

twisted, to 20 mm long 6

6(5). Leaf blades narrowly rolled, to 1 mm across; open

veld S. clandestina

Leaf blades expanded, flat, to 12 mm across; forest

7

7(6). Panicle open, with long spreading flexuous branches

bearing spikelets only towards the end

S. dregeana var. elongata

Panicle dense, with short ascending branches bearing

spikelets from near the base

S. dregeana var. dregeana

Stipa capensis Thunb.

( =S . tortilis Desf.) 1.

Annual; 100-500(-1000) mm
tall. Leaf blades 50-200 mm
long; to 3 mm wide. Spikelets
12-16 mm long (excluding the
bent and twisted awn 50-80 mm
long). Glumes colourless,
translucent, shining.

Flowering August to November. Open veld and
disturbed places in arid winter rainfall regions. Infrequent.
Biome: Nama-Karoo and Succulent Karoo. Also in north
Africa and the Middle East.

Description: De Winter 1965 (217), Chippindall 1955
(290). Illustration; Chippindall 1955 (fig. 259). Voucher:
Acocks 14736. PRECIS code 9902630-00100.

Stipa clandestina Hack.

Perennial; densely tufted (in
big hard tufts); 500-1500 mm
tall. Leaf blades to 750 mm long;
about 1 mm wide (rolled,
setaceous). Spikelets 5-7 mm
long (excluding bent and twisted
awn to 20 mm long). Lemmas
shortly hairy all over.

Flowering November to May.

Disturbed places in veld. Infrequent. Naturalized from
Mexico. Biome: Nama-Karoo. Pasture (readily eaten, green
in winter), or weed.

Description: Hackel 1910 Feddes Rep. 8 (516).
Voucher: Acocks 19284. PRECIS code 9902630-00150.

Stipa dregeana Steud. var. dregeana

Fig. 205.

Similar to var. elongata but
with panicle narrow, branches to
100 mm long, bearing spikelets
nearly from the base.

Biome: Forest. Endemic.

Description: De Winter 1965
(215), Chippindall 1955 (289).
Voucher: Brynard 30. PRECIS
code 9902630-00200.

Stipa dregeana Steud. var. elongata (Nees) Stapf

PI. 187.

Perennial; tufted and rhizo-
matous (rhizomes short, knotted);
900-1200 mm tall. Leaf blades to
60 mm long; to 12 mm wide.
Spikelets 5-7 mm long
(excluding straight or bent and
twisted awn to 18 mm long).
Glumes equal, longer than the
lemmas; panicle open, the

branches slender, to 200 mm long, spreading, drooping,
with spikelets in the upper half only.

Flowering August to May (most commonly in summer).
Moist places in forests. Locally common. Biome: Forest.
Also in east African highlands. Easily mistaken for another

318

erect, broadleaved, long-awned forest grass, Festuca
africana, which has unequal glumes shorter than the
lemmas.

Description: De Winter 1965 (216), Chippindall 1955
(289). Illustration: Chippindall 1955 (fig. 258). Voucher:
Killick & Vahrmeijer 4050. PRECIS code 9902630-00300.

Stipa neesiana Trin. & Rupr.

Perennial; tufted (erect);

300-1000 mm tall. Leaf blades to
300 mm long; to 3 mm wide.

Spikelets 15-17 mm long
(excluding scabrous bent and
twisted awn to 100 mm long).

Glumes dark purple; lemma with
a raised collar at junction of awn.

Flowering November to Dec-
ember. Disturbed places. Infrequent, or locally common.
Naturalized from South America. Weed (in cultivated
lands).

Voucher: Fanshawe 1976-11-05. PRECIS code

9902630-00400.

Stipa papposa Nees

Perennial; tufted; to 600 mm
tall. Leaf blades to 200 mm long;

1-2 mm wide (rolled). Spikelets

8- 10 mm long (excluding fine
bent and twisted awn to 30 mm
long). Glumes shorter than floret;
lemma with a conspicuous brush
of shining white hairs at upper
end.

Flowering December to January. Roadsides. Rare. Nat-
uralized from South America. Biome: Fynbos. Weed.
Known only from a single population on the University of
Cape Town campus, collected in 1963 and 1980.

Voucher: Esterhuysen 30599a. PRECIS code

9902630-00450.

Stipa tenuissima Trin.

Perennial; very densely tufted;

250-1000 mm tall. Leaf blades to
700 mm long; 0.5 mm wide
(tightly rolled, setaceous). Spike-
lets 4.0-5. 0(-5. 5) mm long
(excluding fine bent and twisted
awn to 30 mm long). Glumes not
swollen around floret at base; flo-
ret symmetrical, tapering at both
ends, awn centrally placed.

Flowering January. Open veld. Rare. Naturalized and
invader from South America. Biome: Nama-Karoo.
Delcared weed. Very similar vegetatively to Nassella
trichotoma , which has the floret rounded at the upper end
and the awn asymmetrically placed.

Voucher: Van Graan 411. PRECIS code 9902630-
00500.

Stipa variabilis Hughes

Perennial; tufted; to 700 mm
tall. Leaf blades to 150 mm long;

1-2 mm wide (rolled). Spikelets

9- 10 mm long (excluding sinuous
awn to 50 mm long). Lemma
shortly hairy all over.

Flowering October. Road-
sides. Rare. Naturalized from
Australia. Biome: Fynbos. Po-
tential weed. So far known only from a single specimen
collected at Atlantis.

Voucher: Smook 3617. PRECIS code 9902630-00700.

Stipagrostis Nees

Schistachne Fig. & De Not.

Annual (rarely), or perennial; caespitose. Culms
100-2000 mm high\ herbaceous; branched above, or un-
branched above. Leaf blades linear (narrowly); flat (rarely),
or folded, or rolled (or subterete). Ligule a fringe of hairs.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets solitary; 7-20 mm long (?); compressed
laterally to not noticeably compressed; disarticulating
above the glumes. Rachilla terminated by a female-fertile
floret. Glumes two; very unequal, or more or less equal;
long relative to the adjacent lemmas (usually exceeding it);
awnless; similar (scarious). Lower glume 3 nerved
(usually). All florets female-fertile only; proximal incom-
plete florets absent.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (leathery, the glumes membranous); hairless
(usually glabrous or scabrid); with a clear germination flap;
3 nerved; awned. Awns usually triple or trifid, commonly

Fig. 206. Stipagrostis uniplumis var. uniplumis

319

with a basal column, or not of the triple/trifid, basal column
type ( S . anomala)', apical; non-geniculate (at least, not
geniculate in the usual sense); long-plumose (usually, at
least on the median branch), or hairless ( S . anomala)', much
longer than the body of the lemma. Palea present (but
small); conspicuous but relatively short (usually less than
half lemma length); 2-nerved. Lodicules when present 2;
membranous; glabrous. Stamens 3. Ovary glabrous. Fruit
fusiform; hilum long-linear; pericarp fused; embryo large.

Photosynthetic pathway. C4; XyMS+ (and PCR sheath
single, by contrast with Aristida).

Cytology, classification, distribution. Chromosome base
number, x = 11. Arundinoideae; Aristideae. 50 species.
Africa, southwest Asia, northwest India. Xerophytic; in
open habitats (desert and semidesert, sometimes dunes — -
e.g. S. ciliata being a sandbinder). Namibia, Botswana,
Transvaal, Orange Free State, Natal, Lesotho, and Cape
Province. 27 indigenous species.

References. l.De Winter. 1965. Bothalia 8: 199. 2. Kers.
1971. Svensk. Bot. Tidskr. 65: 199.

Species treatment by L. Smook.

1(0). Awn solitary (protruberances indicating two lateral
awns sometimes present), awn not plumose except
for a pencil of long white hairs around the base of
the column; articulation present between the apex

of the lemma and the base of the column

S. anomala

Awns three, either all three or only the central awn
distinctly and densely plumose; articulation absent

or present 2

2(1). Lemma articulation absent

S. zeyheri subsp. sericans

Lemma articulation present 3

3(2). Callus minutely bifid S. obtusa

Callus never bifid 4

4(3). All three awns distinctly plumose with long hairs
(lateral awns indistinctly plumose with short and/or

scattered, long hairs) 5

Only central awn distinctly plumose, lateral awns
glabrous or with short hairs and/or scattered long

hairs, not distinctly plumose 15

5(4). Lemma articulation near the middle of the lemma • 6
Lemma articulation near the apex of the lemma . . 7
6(5). Spikelets to 14 mm long (including awns); upper

glume 7-9 mm long S. proxima

Spikelets 15-30 mm long (including awns); upper

glume 10-14 mm long S. namaquensis

7(5). Plants delicate, culms to 1.2 mm wide; upper glume
7-9 mm long; lower leaf surface rough, densely

covered with prickles S. ramulosa

Plants robust or reed-like, culms 1. 2-5.0 mm wide;
upper glume 9-25 mm long; lower leaf surface
smooth, prickles absent, or present only on the side

of the nerves in the intercostal cavities 8

8(7). Plant reed-like; leaves rigid, straight and pungent,
overtopping the narrow dense inflorescence;

column very short and stout S. sabulicola

Plants robust, but not reed-like; inflorescence usually
extending beyond the leaves, if overtopped by the
leaves, then leaves flaccid and often curling with
age; column long, or if short, then slender .... 9
9(8). Column hairy with long hairs for at least some
distance below branching point of the awns . . 10
Column glabrous or scaberulous or with only a few
scattered hairs around the branching point of the
awns 12

1 0(9). Glumes densely hairy with long hairs, or sometimes

only hairy at the glume apex

S. zeyheri subsp. sericans

Glumes glabrous, puberulous or scabrid 11

1 1(10). Leaves erect and rigid; inflorescence usually open;

glumes usually purple; plant often tinged purple;

mainly from the winter rainfall area

S. zeyheri subsp. zeyheri

Leaves flaccid, often curling; inflorescence usually
narrow; glumes pallid, slightly darker at the base;
plant sometimes faintly flushed with purple;
coastal areas of eastern Cape and northern Natal

S. zeyheri subsp. barbata

12(9). Inflorescence narrow, compact, the branches
appressed to the main axis; callus bluntly
rounded; awns plumose with long silver-white

hairs S. damarensis

Inflorescence open; callus pungent; awns plumose
with long dirty-white, yellow or silver hairs • 13
13(12). Glumes longer than 15 mm; awns plumose with

long dirty-white or yellow hairs

S. zeyheri subsp. macropus

Glumes shorter than 15 mm; awns plumose with

long silvery hairs 14

14(13). Axils of inflorescence branches glabrous; culm
nodes glabrous . . . . S. lutescens var. lutescens
Axils of inflorescence branches distinctly bearded;
culm nodes hairy . S. lutescens var. marlothii
15(4). Lemma articulation at or just above the middle of

the body of the lemma 16

Lemma articulation at the apex of the lemma . 22
16(15). Lower glume narrowly oblong to oblong, apex firm,

obtuse to truncate 17

Lower glume linear to lanceolate, apex

membranous, acute to long-acuminate 18

17(16). Culm nodes bearded with a ring of long, spreading
white hairs; lower leaf sheaths not covered with
a mat of woolly hairs . S. ciliata var. capensis
Culm nodes glabrous; lower leaf sheaths sparsely
to densely covered with matted woolly hairs . .

S. schaeferi

18(16). Plants suffrutescent; culms with branches fascicled;

central awn usually to 35 mm long . S. amabilis
Plants tufted, not woody; culms branched or
unbranched, branches not fascicled; central awn

usually 40-100 mm long 19

19(18). Inflorescence narrow, spike-like, unbranched . 20
Inflorescence narrow, interrupted, branched . . 21

20(19). Lower glume with long, rigid, erect hairs

. . . S. hochstetteriana var. hochstetterriana

Lower glume without long erect hairs

S. hochstetteriana var. secalina

21(19). Glumes softly pilose, especially along the margins
near the apex; central awn plumose right to the
tip; lateral awns up to 1/3 the length of the central
awn; culms with striations indistinct or widely

separate, densely scabrid, covered with

conspicuous prickles S. dinteri

Glumes not softly pilose; central awn excurrent into
a delicate naked tip; lateral awns at least 1/2 the
length of the central awn; culms with striations
distinct and close together, smooth or minutely
scaberulous, with small inconspicuous prickles
(Note: 5. giessii X hochstetteriana has glumes

with long stiff hairs) S. giessii

22( 15). Leaves mainly cauline, with well to poorly
developed leaf blades; plants usually suffruticose

Leaves apparently mainly basal, with well
developed leaf blades; plants not woody ... 28

23(22). Glumes with long white hairs 24

Glumes glabrous or with only very short hairs . 25
24(23). Inflorescence not or only slightly exserted from
uppermost leaf sheath, spikelet fascicles densely
clustered, lowermost sometimes separated from

the rest; leaf blades poorly developed

S. getninifolia

Inflorescence well exserted from uppermost leaf
sheath; spikelet fascicles much interrupted along
the main axis; leaf blades well developed ....
S. fastigiata

320

25(23). Plants with raised, round glands . . . S. brevifolia

Plants without raised, round glands 26

26(25). Plants slender in the upper parts; leaves flexuous,
usually held at an angle of 45 degrees from the

culm, to 1 mm wide S. garubensis

Plants robust in upper parts; leaves rigid, usually at
an angle of 90 degrees from the culm, 1-2 mm

wide 27

27(26). Axils of inflorescense branches glabrous; culm
nodes glabrous . . . . S. lutescens vari lutescens
Axils of inflorescence branches distinctly bearded;
culm nodes hairy . S. lutescens var. marlothii
28(22). Glumes with obvious, long hairs, though sometimes

along the margins only 29

Glumes puberulous, scabrid or glabrous 33

29(28). Inflorescence spiciform, subsecund; culms not
visibly or obviously striate, usually densely

scabrid S. gonatostachys

Inflorescence open or contracted but not spiciform
and subsecund; culms conspicuously striate,

smooth 30

30(29). Callus with short hairs along the entire length
(except for the naked tip), meeting the long hairs
at the junction between the lemma and the callus;
plants annual (Note: S. uniplumis X hirtigluma
is perennial) . . . S. uniplumis var. intermedia
Callus with a distinct glabrous break between the
short hairs along the length of the callus and the
long hairs at the junction of the lemma and the

callus; plants annual or perennial 31

31(30). Inflorescence narrow, when fully exserted much

longer than wide; plants annual

S. hirtigluma subsp. hirtigluma

Inflorescence open, spreading, when fully exserted
not much longer than wide; plants annual or

perennial 32

32(31). Plants annual with very few leaves at the base . .

S. hirtigluma subsp. pearsonii

Plants perennial with a dense tuft of basal leaves

S. hirtigluma subsp. patula

33(28). The branching point of the awns and a short
distance down the column with hairs longer than

1.5 mm 34

The branching point of the awns and a short
distance down the column glabrous, scabrid or

with hairs shorter than 1 .5 mm 35

34(33). Inflorescence with numerous spikelets; glumes
usually to 10 mm long; central awns usually

straight S. uniplumis var. uniplumis

Inflorescence with a few spikelets; glumes 10 mm
or longer; central awns bent at right angles . . .

S. uniplumis var. neesii

35(33). Inflorescence subsecund, branched only in the
lower part, spikelets in the upper part solitary,
borne on robust, rigid pedicels directly on the

main axis S. gonatostachys

Inflorescence not subsecund, much branched for
most of its length, spikelets paired or solitary,
borne on slender, usually flexuous pedicels from

the branches 36

36(35). Inflorescence contracted and very dense, main axis

hidden 37

Inflorescence open or contracted, interrupted, main

axis clearly visible 38

37(36). Column densely short-hairy at the swollen
branching point of the awns; callus 1 .5 mm long;
culms well developed and extending somewhat
beyond the basal tuft of leaves . . S. hermannii
Column glabrous, sometimes with a few scattered
hairs around branching point of awns; callus
0.8-1. 0 mm long; culms very poorly developed

and short S. subacaulis

38(36). Column hairy; plants annual S. namibensis

Column smooth, glabrous or densely scabrid; plants
usually perennial 39

39(38). Inflorescence open, branches with long naked basal

parts; glumes dark; column smooth

S. dregeana

Inflorescence contracted, interrupted, branches
bearing spikelets to near the base; glumes pallid;

column usually densely scabrid 40

40(39). Lower leaf sheaths densely covered all over with
matted woolly hairs; plants rare .... S. lanipes
Lower leaf sheaths glabrous, if hairy these not
matted woolly hairs and present only at the very
base; plants common S. obtusa

Stipagrostis amabilis (Schweick.) De Winter

(=Aristida amabilis
Schweick.) 1.

Kalahari dune bushman grass,
duinekweek.

Shrub or dwarf shrub; rhizo-
matous (rhizomes long, creep-
ing), or tufted (culms erect or
horizontal); 1500-2000 mm tall. Leaf blades curved, sharp,
to 250 mm long; to 2.5 mm wide. Spikelets 1 1-14 mm long
(excluding awns). Culms with branches fascicled at nodes,
internodes distinct; inflorescence narrow, interrupted, with
spikelets crowded, pedicels erect; glumes unequal; lower
glume lanceolate, long-acuminate; lemmas smooth, articu-
lation at about the middle of the lemma; column short to
almost absent; central awn to 35 mm long, plumose for the
upper 2/3, lateral awns not plumose; callus 1.5 mm long,
tip naked, pungent.

Flowering sporadic from August to May. On the crest
of Kalahari sand dunes. Locally common. Biome: Savanna
and Nama-Karoo. Endemic. Erosion control (sand binder on
dune crests). Similar toS. namaquensis , which has all three
awns plumose and shorter leaves with more pungent tips.

Description: De Winter 1965 (324). Voucher: Leistner
1365. PRECIS code 990261 1-00100.

Stipagrostis anomala De Winter

{=Stipa namaquensis Pilg.,
non Stipagrostis namaquensis
(Nees) De Winter) 1.

Torro-boesmangras.

Weakly perennial or annual;
densely tufted (erect or slightly
geniculate near base); 100-600
mm tall. Leaf blades scabrid 10-200 mm long; setaceous,
to 1.5 mm wide. Spikelets 9-12 mm long (excluding awns).
Leaves mainly basal, often curved; inflorescence narrow,
interrupted, with spikelets erect, crowded along the main
axis; glumes unequal, scaberulous; lemma articulation
between the apex of the lemma and the base of the column;
column twisted and with long stiff hairs at the base; awn
solitary, not plumose, diverging at right angles at maturity,
protruberances indicating rudimentary lateral awns are
sometimes present; callus 1.5 mm long, pungent.

Flowering January to June (and August and September).
Shallow sandy soils over rocks on slopes and gravel plains.
Locally common. Biome: Nama-Karoo. Endemic. Although
this species has only a single glabrous awn, the three-nerved
glumes, hairs at the base of the column and anatomical
characters place it in Stipagrostis rather than Aristida. It
differs from the genus Stipa , which has a membranous
ligule.

Description: De Winter 1965 (375). Illustration: Muller
1984 (fig. 121), Chippindall 1955 (fig. 260). Voucher:
Leistner 2362. PRECIS code 990261 1-00200.

Fig. 207. PI. 188.

321

Stipagrostis brevifolia (Nees) De Winter

( =Aristida brevifolia (Nees)

Steud.) 1.

Langbeentwagras, kortblaar-
boesmangras.

Robust shrub or dwarf shrub
(culms much branched); rhizo-
matous (rhizomes branched and woody); to 1000 mm tall.
Leaf blades usually very short, 5-80(-120) mm long; rolled
or expanded, 1-3 mm wide. Spikelets 12-15 mm long (ex-
cluding awns). Vegetative parts with raised round glands;
nodes densely covered with woolly hairs; leaves mainly
cauline; inflorescence narrow, sometimes interrupted,
branches appressed to main axis; glumes long acuminate,
glabrous; lemma articulation between the apex of lemma
and the base of the column; column distinct, scabrid; only
the central awn plumose with the lower 1/4 scabrid and the
apex not plumose; callus 2.0-2. 5 mm long, with a naked,
pungent tip.

Flowering September to May. Sand over rocks on plains
and especially in drainage areas. Locally common. Biome:
Nama-Karoo and Succulent Karoo. Endemic. Drought
resistant pasture (palatable only when green). Hybridizes
with S. namaquensis, (De Winter 3266).

Description: De Winter 1965 (338), Stapf 1898-1900
(570). Illustration: Muller 1984 (fig. 122), Chippindall 1955
(fig. 270). Voucher: De Winter & Hardy 7852. PRECIS
code 9902611-00300.

Stipagrostis ciliata (Desf.) De Winter var. capensis

(Trin. & Rupr.) De Winter

(=Aristida ciliata sensu
Desf., non Steud.& Hochst. ex
Steud.) 1; (-Aristida ciliata
Desf. var. capensis Trin. &

Rupr.) 1; ( =Aristida ciliata Desf.
var . pectinata Henr.) 1;

(= Aristida ciliata Desf. var.
tricholaena Hack.) 1; (-Aristida
ciliata Desf. var. villosa
Hack.) 1.

Langbeenboesmangras, tall bushman grass.

Densely or laxly tufted (erect or occasionally genicu-
late); 850-1000 mm tall. Leaf blades to 300 mm long; to
1.5 mm wide. Spikelets 6.5-12 mm long (excluding awns).
Leaves mainly basal; sheaths hairy but not woolly, or gla-
brous; culm nodes with long stiff spreading hairs; inflores-
cence narrow or open, branches flexuous; spikelets variable
in size, straw coloured, often purple at the base; glumes
equal to subequal, lower glume oblong to narrowly oblong,
apex obtuse to truncate, firm; articulation about in the
middle of the lemma; column length variable; only central
awn plumose, hairs usually silvery (occasionally golden);
callus 2. 0-2. 5 mm long, with pungent, naked point.

Flowering August to October, and February to June.
Coarse sandy soils especially in river beds or on gravel
plains. Locally common. Biome: Savanna, Nama-Karoo,
Succulent Karoo, and Desert. Also in Tunisia and Egypt.
Variable, sometimes hybridizes with S. zeyeri subsp.
macropus (Acocks 14817). Closely allied to S. schaeferi,
which has glabrous nodes and is far less common.

Description: De Winter 1965 (316), Stapf 1898-1900
(563), Chippindall 1955 (299). Illustration: Muller 1984
(fig. 213), Chippindall 1955 (fig. 265). Voucher: Smook
2896, Oliver, Muller & Steenkamp 6612. PRECIS code
990261 1-00400.

Stipagrostis damarensis (Mez) De Winter

( =Aristida damarensis
Mez) 1 .

Robust perennial; laxly tufted
(much branched near the base), or
rhizomatous (rhizome well devel-
oped); to 1200 mm tall. Leaf
blades to 300 mm long; 2-3 mm
wide. Spikelets 12-14 mm long
(excluding awns). Culms 1. 2-5.0 mm wide; lower leaf
surface with prickles on the nerves but sunk in the intercost-
al cavities; inflorescence extending beyond the leaves,
elongate, narrow, compact, often interrupted, branches
appressed to the main axis; spikelets erect; glumes glabrous
or pilose near apex and on the margins; upper glume 10-15
mm long; lemma articulation between the apex of the lem-
ma and the base of the column; column long, glabrous, well
developed; all three awns completely plumose, with long
silver-white hairs; callus 1 mm long, hairy almost to the tip,
tip bluntly rounded.

Flowering March to June. River beds and drainage lines.
Locally common. Biome: Savanna and Desert. Endemic.

322

Resembles S. namaquensis, which has the articulation in the
middle of the lemma and the column not as well developed.

Description: De Winter 1965 (329). Voucher: De Winter
& Hardy 8131, Giess 7912. PRECIS code 990261 1-00500.

Stipagrostis dinteri (Hack.) De Winter

( =Aristida dinteri Hack.) 1.

Slender perennial; densely
tufted; to 400 mm tall. Leaf
blades to 150 mm long; to 1 mm
wide. Spikelets 15-16 mm long
(excluding awns). Culms, if
branched, not in fascicles; vegeta-
tive parts densely scabrid, of-
ten with round, usually crateriform glands; culms indistinct-
ly striate, or striations widely separate, densely covered
with conspicuous prickles; inflorescence narrow, branched,
interrupted; glumes pilose with short hairs (at least on the
margins); lower glume linear to lanceolate, tapering to a
long acuminate apex; lemma articulation just above the
middle of the lemma; column of variable length; central
awn 40-100 mm long, plumose to the lower 1/3 and to the
apex, lateral awns to 1/3 the length of the central awn, not
plumose; callus 2 mm long, with a distinct, naked, pungent
tip.

Flowering November and February to May. Loose sand
in riverbeds and on hills. Locally common. Biome: Nama-
Karoo and Desert. North to Angola.

Description: De Winter 1965 (320), Chippindall 1955
(300). Voucher: Giess, Volk & Bleissner 6249, Giess 7984.
PRECIS code 990261 1-00600.

Stipagrostis dregeana Nees

{-Aristida dregeana (Nees)

Trin. & Rupr.) 1 .

Rock bushman grass.

Laxly to densely tufted (erect
to geniculate, branched at base);
to 300 mm tall. Leaf blades to 135
mm long (smooth with scabrid
margins); setaceous. Spikelets to 12 mm long (excluding
awns). Leaves mainly basal; inflorescence open, nearly as
long as wide, with main axis visible, branched, branches
somewhat flexuous, with long naked basal parts, pedicels
slender; glumes glabrous, dark; lemma articulation between
the apex of the lemma and the base of the column; column
smooth and glabrous to the branching point of the awns; all
three awns, or only the central awn plumose; callus 1-3 mm
long, tip naked, pungent.

Flowering August and April. Coarse, sandy soils or
shallow soils, between rocks and in depressions along
roadsides. Infrequent. Biome: Succulent Karoo. Endemic.

Description: De Winter 1965 (344), Stapf 1898-1900
(569). Voucher: Giess & Van Vuuren 682. PRECIS code
9902611-00700.

Stipagrostis fastigiata (Hack.) De Winter

( =Aristida fastigiata
Hack.) 1.

Shrub or dwarf shrub (suf-
fruticose, culms fascicled); rhizo-
matous (rhizomes thick and much
branched); to 600 mm tall. Leaf
blades to 80 mm long (often much
shorter); 2-3 mm wide. Spike-
lets 15 mm long (excluding awns). Leaves mainly cauline.

blades well developed; inflorescence usually elongate, ex-
serted from the uppermost leaf sheaths; spikelets in fascic-
les which are lax and interrupted along the inflorescence;
glumes densely hairy with long white hairs; lemmas
smooth, articulation between the apex of the lemma and the
base of the column; column well developed, twisted; only
the central awn plumose; callus 2 mm long with a pungent
naked tip.

Flowering February to June. Sandy and alkaline soils.
Locally common. Biome: Nama-Karoo, Succulent Karoo,
and Desert. Endemic. Pasture. Closely related to S.
geminifolia , which is usually smaller than 300 mm, with the
inflorescence as long as wide and the lower part enclosed
in the swollen upper leaf sheath, and with spikelet fascicles
densely clustered.

Description: De Winter 1965 (338). Illustration: Muller
1984 (fig. 124). Voucher: Acocks 21790, Giess 13436.
PRECIS code 990261 1-00800.

Stipagrostis garubensis (Pilg.) De Winter

( =Aristida garubensis
Pilg.) L

Shrub or dwarf shrub; base
robust, woody and branched; to
600 mm tall. Leaf blades to 120
mm long; to 1 mm wide. Spike-
lets 12-14 mm long (excluding
awns). Plants slender in the upper
parts; leaves flexuous, held at 45 degree angles from the
culms, pungent; glumes often dark at the base; lemma apex
densely tuberculate, articulation between the apex of the
lemma and the base of the column; column well developed,
slender; only the central awn plumose; callus 1.0-1.5 mm
long, with a pungent, naked tip.

Flowering June, July and September. Between rocks,
especially granite, on hillslopes and in riverbeds. Locally
common. Biome: Succulent Karoo. Endemic.

Description: De Winter 1965 (343). Voucher: Kinges
2289. PRECIS code 990261 1-00900.

Stipagrostis geminifolia Nees

(= Aristida geminifolia (Nees)

Trin. & Rupr.) 1 .

Shrub or dwarf shrub; erect or
geniculate; to 250 mm tall. Leaf
blades usually very short, ex-
panded and rigid 1 0(— 20) mm
long; to 2 mm wide. Spikelets
10-14 mm long (excluding
awns). Leaves mainly cauline, blades not well developed;
inflorescence ovate to oblong, not or only slightly exserted
from the uppermost leaf sheath; spikelet fascicles densely
clustered, the uppermost sometimes separated from the rest;
glumes densely covered with long white hairs; lemma
smooth, articulation between the apex of the lemma and the
base of the column; column short, glabrous; only the central
awn plumose, hairs usually golden; callus 2 mm long, with
a pungent, naked tip.

Flowering August to October, and January to June.
Coarse sandy soils of watercourses and in open places on
gravel plains. Infrequent to locally common. Biome: Suc-
culent Karoo. Endemic. Pasture (eaten by stock). Closely
related to S. fastigiata , which has an elongated
inflorescence exserted from the upper leaf sheath and
spikelet fascicles not densely clustered.

Description: De Winter 1965 (341), Stapf 1898-1900
(570). Illustration: Chippindall 1955 (fig. 268). Voucher:
Ellis 2181. PRECIS code 990261 1-01000.

323

Stipagrostis giessii Kers

Perennial; tufted; to 800 mm
tall. Leaf blades 70-250 mm
long; to 2 mm wide. Spikelets
13-22 mm long (excluding
awns). Vegetative parts scabrid,
culm striations distinct and close
together, smooth or scaberulous
with minute prickles; leaves
mainly basal; inflorescence nar-
row, with branches of variable lengths; pedicels short,
thick, erect; glumes papery, lower glume linear to lanceo-
late, glabrous, acuminate, tuberculate; lemma articulation
is at about the middle of the lemma; column long, slender
and twisted; central awn 40-100 mm long, plumose with
long hairs on the upper half and excurrent into a delicate
naked tip; lateral awns not plumose, stout and at least 1/2
the length of the central awn; callus 2 mm long, with a pun-
gent, naked tip.

Flowering November and March to June. Sandy river
beds, stony hills and gravel plains. Locally common.
Biome: Savanna, Nama-Karoo, and Desert. North to
Angola. Variable; differs from S.hochstetteriana, which has
an unbranched inflorescence. Hybrids with S.
hochstetteriana have been reported (De Winter & Hardy
8058).

Description: Kers 1971 (199). Voucher: De Winter &
Hardy 8185, Giess & Leippert 7413. PRECIS code
9902611-01100.

Stipagrostis gonatostachys (Pilg.) De Winter

( -Aristida gonatostachys

Pilg.) 1.

Rough-leaved bushman grass.

Perennial; densely tufted; to
200 mm tall. Leaf blades 40-50
mm long; folded. Spikelets 8-10
mm long (excluding awns).

Leaves mainly basal; vegetative parts densely scabrid;
culms not conspicuously striate; inflorescence elongate,
narrow, spiciform and subsecund, base usually enclosed in
upper leaf sheath; glumes scabrid or densely covered with
long hairs; lemma smooth, articulation between the apex of
the lemma and the branching point of the column; column
well developed, scaberulous, if hairy at branching point,
hairs less than 1.5 mm long; only the central awn plumose;
callus 1.5 mm long with a pungent, naked tip.

Flowering September to December and March to June.
Coarse to fine sand between rocks on mountain slopes, and
in depression on plains where water collects. Infrequent.
Biome: Desert. Endemic. Easily confused with the more
widespread S. ohtusa , which has a shorter callus, an inflor-
escence which is not subsecund and is well exserted from
the upper leaf sheath.

Description: De Winter 1965 (353). Voucher: Giess
13421, De Winter & Hardy 8098. PRECIS code
990261 1-01200.

Stipagrostis hermannii (Mez) De Winter

(-Aristida hermannii Mez) 1.

Laxly tufted (geniculate to
prostrate); to 150 mm tall. Leaf
blades densely scabrid 10-20 mm
long; to 2 mm wide. Spikelets
9-14 mm long (excluding awns).

Culms well developed and ex-
tending somewhat from the basal
tuft of leaves; inflorescence narrow, dense, much branched
for most of its length, with the main axis hidden, the base
partly enclosed in the leaf sheath; glumes glabrous, with a

long tapering acuminate apex; lemma articulation between
the apex of the lemma and the base of the column; column
length variable, densely short-hairy, with hairs shorter than
1.5 mm long on the swollen branching point of the awns;
only the central awn plumose; callus 1.5 mm long with a
pungent, naked tip.

Flowering January to August. Sandy areas on hills and
plains. Locally common. Biome: Desert. Endemic. Closely
allied to S. subacaulis, which has a glabrous column and a
callus 0.8-1. 0 mm long.

Description: De Winter 1965 (369). Voucher: Giess &
Robinson 13212, Dinter 6396. PRECIS code 9902611-
01300.

Stipagrostis hirtigluma (Trin. & Rupr.) De Winter subsp.

hirtigluma

( =Aristida hirtigluma Steud.
ex Trin. & Rupr.) 1.

Annual; tufted (erect); to 500
mm tall. Leaf blades 60-200 mm
long; setaceous. Spikelets 10-15
mm long (excluding awns).

Culms conspicuously striate,
smooth; leaves mainly basal;
inflorescence narrow, much longer than wide; glumes hairy,
with long hairs on both sides of central nerve along inner
and outer surfaces; lemma articulation between the apex of
the lemma and the base of the column; column variable in
length and hairiness; only the central awn plumose; callus
0.4 mm long, with a distinct glabrous break between the
short hairs on the callus body and the long hairs at the
junction between the lemma and the callus, and with a long
naked, pungent tip.

Flowering April to May. Not habitat selective, grows on
sandy or gravelly soils as well as rocky substrates. Locally
common. Biome: Savanna, Nama-Karoo and Desert. To
Angola, North Africa, and the desert areas of the Middle
East and west Africa. Barely distinguishable from subsp.
pearsonii, which has an open, spreading inflorescence not
much longer than wide, and from subsp. patula, which is
perennial.

Description: De Winter 1965 (361). Voucher: Giess
3034. PRECIS code 990261 1-01400.

Stipagrostis hirtigluma (Trin. & Rupr.) De Winter subsp.

patula (Hack.) De Winter

(-Aristida gracilior Pilg. var.
gracilior) 1.

Perennial; densely tufted (e-
rect); to 600 mm tall. Leaf blades
to 200 mm long; setaceous.

Spikelets 12-13 mm long (ex-
cluding awns). Culms con-
spicuously striate, smooth; leaves mostly in dense basal
tufts; inflorescence open, spreading, not much longer than
wide; glumes brown, usually flushed with purple, densely
hairy; lemmas densely tuberculate, articulation between the
apex of the lemma and the base of the column; column well
developed, stout; only the central awn plumose, the long
spreading hairs cover the entire awn and extend for varying
lengths down the column; callus 0.7 mm long, with a dis-
tinct glabrous break between the short hairs on the callus
body and the long hairs at the junction of the lemma and
the callus, with a pungent, naked tip.

Flowering February to July. Calcareous soils or sandy
areas near limestone around pans, on floodplains or rocky
ridges. Locally common. Biome: Savanna. Also in tropical
Africa. Hybrids between this var. and S. uniplumis var.
neesii have been reported. They are recognized by being
perennial, with hairy glumes and a S. uniplumis type callus
(Volk 2338).

Description: De Winter 1965 (361 & 365). Voucher: De
Winter & Giess 7135. PRECIS code 9902611-01500.

324

Stipagrostis hirtigluma (Trin. & Rupr.) De Winter subsp.

pearsonii (Henr.) De Winter

( =Aristida gracilior Pilg. var.
pearsonii Henr.) 1.

Annual; densely tufted (erect
to geniculate); to 800 mm tall.

Leaf blades to 200 mm long; set-
aceous. Spikelets 10-13 mm long
(excluding awns). Culms con-
spicuously striate, smooth; leaves
sparse at the base; inflorescence open, spreading, not much
longer than wide; glumes densely hairy; lemma articulation
between the apex of the lemma and the base of the column;
column thick; callus 0.7 mm long, with a distinct glabrous
break between the short hairs on the body of the callus and
the long hairs at the junction of the lemma and the callus,
with a pungent, naked tip.

Flowering January to May. Sandy soils on gravel flats,
and on stony hills. Locally common. Biome: Savanna,
Nama-Karoo and Desert. To Angola. Pasture (not highly
palatable but utilized). Barely distinguishable from subsp.
hirtigluma, which has a narrower inflorescence that is
longer than wide when fully exserted, and from subsp.
patula, which is a more definite perennial.

Description: De Winter 1965 (361). Illustration: Muller
1984 (fig. 125). Voucher: Oertendahl 158, De Winter &
Leistner 5621. PRECIS code 9902611-01600.

Stipagrostis hochstetteriana (Beck ex Hack.) De

Winter var. hochstetteriana

{-Aristida hochstetteriana
Beck ex Hack.) 1 .

Spike bushman grass.

Perennial; densely tufted (e-
rect); to 900 mm tall. Leaf blades
250^400 mm long; 2. 0-2. 5 mm
wide. Spikelets 15-20 mm
long (excluding awns). Leaves mainly basal; inflorescence
spike-like, narrow, unbranched; pedicels thick; lower
glume linear to lanceolate, with long, rigid, erect hairs
usually either side of the median keel, apex long acuminate,
membranous; lemma articulation around the middle of the
lemma; column well developed, slender; central awn
40-100 mm long, plumose in the upper 2/3; lateral awns
not plumose; callus 2 mm long with naked, pungent tip.

Flowering November to June. Sandy to clay soils on
rocky slopes and on gravels, often calcareous, disturbed
areas at roadsides or in river courses. Locally common.
Biome: Nama-Karoo and Desert. Endemic (possibly
occuring in Angola). Distinguished from var. secalina,
which has no hairs on the lower glumes, and from S. giessii,
which has the inflorescences branched. Hybrids with S.
giessii have been reported (De Winter & Hardy 8097).

Description: De Winter 1965 (313), Stapf 1898-1900
(571), Chippindall 1955 (297). Illustration: Chippindall
1955 (fig. 263). Voucher: Muller 116, De Winter 3254.
PRECIS code 990261 1-01700.

Stipagrostis hochstetteriana (Beck ex Hack.) De

Winter var. secalina (Henr.) De Winter

( =Aristida secalina Henr.) 1.

Rye bushman grass.

Perennial; densely tufted (e-
rect); to 900 mm tall. Leaf blades
to 400 mm long; 2 mm wide.

Spikelets 15-20 mm long (ex-
cluding awns). Leaves mainly
basal; inflorescence spike-like, narrow, unbranched, pedi-
cels thick; lower glume linear to lanceolate, apex long
acuminate, membranous, without long, erect, rigid hairs;
lemma articulation around the middle of the lemma; column

well developed, slender; central awn 40-100 mm long,
plumose, hairs only in the upper 2/3; lateral awns not
plumose; callus 2 mm long, with a naked pungent tip.

Flowering February to June. Sandy soil and rocky
slopes, especially limestone. Locally common. Biome: Sa-
vanna, Nama-Karoo, Desert. To Angola. May cause grass
balls in sheep. Similar to var. hochstetteriana, which has
erect hairs on the back of the lower glume, and toS. giessii,
which has branched inflorescences.

Description: De Winter 1965 (313). Voucher: Theron
3834, De Winter 3048. PRECIS code 990261 1-01800.

Stipagrostis lanipes (Mez) De Winter

{-Aristida lanipes Mez) 1 .

Woolly bushman grass.

Perennial; densely tufted; to
600 mm tall. Leaf blades to 25
mm long. Spikelets to 9.5 mm
long (excluding awns). Leaves
mainly basal, lower leaf sheaths
densely woolly-hairy; inflorescence contracted, interrupted,
much branched, branches bearing spikelets to near the base;
glumes glabrous, pallid; lemma articulation between the
apex of the lemma and the base of the column; column
densely scabrid; only central awn distinctly plumose; callus
1 .4—1.6 mm long.

Flowering August. Sandy soils on slopes. Rare. Biome:
Succulent Karoo. Endemic if distinct from S. obtusa, which
occurs in North Africa, Sinai Peninsula and Iraq to
Pakistan. There are very few specimens that can definitely
be identified as S. lanipes and its status as a species distinct
from S. obtusa is somewhat doubtful.

Description: De Winter 1965 (355). Voucher: De Winter
& Giess 6133. PRECIS code 990261 1-01900.

Stipagrostis lutescens (Nees) De Winter var. lutescens

(=Aristida lutescens (Nees)

Trin. & Rupr.) 1.

Shrub or dwarf shrub; rhizo-
matous (rhizome long, branched);

700-1000 mm tall. Leaf blades to
100 mm long; to 2 mm wide
(flat). Spikelets 12-14 mm long
(excluding awns). Plants robust in
upper parts; culms stout, much branched and fascicled in
the lower parts, nodes glabrous; leaves mainly cauline,
rigid, usually held at 90 degree angles from the culm,
relatively short, flat or folded, distinctly pungent; inflores-
cence open, axils of branches glabrous; glumes 8-14 mm
long, glabrous; lemma articulation between the apex of
the lemma and the base of the column; column well
developed, glabrous or occasionally hairy; all three awns
or only the central awn plumose; callus 2. 0-2. 5 mm long,
with a narrow, pungent, naked tip.

Flowering March, and July to October. Sandy soils. In-
frequent. Biome: Succulent Karoo. Endemic. Similar to var.
marlothii , which has hairy nodes and hairs in the axils of
the inflorescence branches.

Description: De Winter 1965 (334), Stapf 1898-1900
(567). Voucher: Schlieben 1 1480. PRECIS code 990261 1-
02000.

Stipagrostis lutescens (Nees) De Winter var. marlothii

(Hack.) De Winter

(=Aristida marlothii
Hack.) 1.

Leeugras.

Shrub or dwarf shrub; rhizo-
matous (rhizomes long, strong
and branched; culms branched
and fascicled in lower part); to

325

1500 mm tall. Leaf blades rigid, to 150 mm long (usually
shorter); 2-3 mm wide (flat). Spikelets 12-14 mm long (ex-
cluding awns). Plants robust in the upper parts; culm nodes
bearded with long spreading hairs; leaves mainly cauline,
rigid, usually held at 90 degree angles from the culm,
relatively short, flat or folded, distinctly pungent; inflores-
cence open, hairy in axils of branches; glumes 8-14 mm
long, glabrous; lemma articulation between the apex of the

Fig. 208. Stipagrostis namaquensis

lemma and the base of the column; column well developed,
glabrous or with occasional hairs; all three awns or only the
central awn plumose, hairs long, silvery; callus 2. 0-2. 5 mm
long, narrow, with naked, pungent tip.

Flowering throughout the year. Locally common.
Biome: Desert. Endemic. Differs from var. lutescens , which
has glabrous nodes and no hairs in the axils of the inflores-
cence branches.

Description: De Winter (334). Voucher: De Winter &
Hardy 7891. PRECIS code 9902611-02100.

Stipagrostis namaquensis (Nees) De Winter

Fig. 208.

(-Aristida namaquensis
(Nees) Trin. & Rupr.) 1.

Steekrietboesmangras, river
bushman grass.

Robust to slender shrub or
dwarf shrub; rhizomatous (rhi-
zomes long and strong), or tuft-
ed (densely, sprawling to erect); to 2000 mm tall. Leaf
blades 60-100 mm long; setaceous. Spikelets 10-14 mm
long (excluding awns). Plants glabrous and smooth, except
for upper and basal leaf sheaths, which are appressed wool-
ly-hairy; culm branches usually fascicled from the nodes,
rarely solitary; leaf blades break off early, leaving the
sheaths, and exposing the upper part of the internode, which
is usually dark, giving it the characteristic banded
appearance; inflorescence elongate, narrow, interupted,
with spikelets clustered; spikelets 15-30 mm long, in-
cluding the awns; glumes straw-coloured; upper glume
10-14 mm long; lemma articulation at about the middle of
the lemma, a short, twisted beak present; all three awns
plumose; callus 1.5 mm long, with naked, pungent tip.

Flowering February to May and July to December. Dry
river courses, on loose gravelly soils, rarely on sand dunes.
Locally common. Biome: Savanna, Nama-Karoo, Succulent
Karoo, and Desert. Endemic. Drought resistant pasture
(grazed by karakul), or erosion control (sand binder and silt
catcher). Hybridizes with S. brevifolia (De Winter 3266).
Similar to S. amahalis, which has only the central awn
plumose and longer, less pungent leaves.

Description: De Winter 1965 (325), Stapf 1898-1900
(566), Chippindall 1955 (298). Illustration: Muller 1984
(fig. 127), Chippindall 1955 (fig. 264). Voucher: Ward 254,
Smook 4488. PRECIS code 990261 1-02200.

Stipagrostis namibensis De Winter

Annual; tufted (sprawling and A

lax); to 300 mm tall (usually \. r'v\^ j

smaller). Leaf blades scabrid, to \r J
30 mm long; to 1 mm wide (flat f ,/ \

or slightly folded). Spikelets 8-9 \ Jj

mm long (excluding awns). { Qr1

Leaves mainly basal; inflores- \ 7

cence narrow but open (main axis
visible), much branched; glum-
es firm, glabrous, apex membranous; lemma granular,
especially around the articulation, articulation between the
apex of the lemma and the base of the column; column
length variable, hairy, with hairs shorter than 1.5 mm; only
the central awn plumose; callus 1.0-1. 6 mm long, with
naked, pungent tip.

Flowering March to June. In depressions where water
collects, on sandy and gravel plains. Locally common.
Biome: Desert. Endemic, or possibly also in Angola.

Description: De Winter 1965 (370). Voucher: Giess,

Volk & Bleissner 5739. PRECIS code 990261 1-02300.

326

Stipagrostis obtusa (Del.) Nees

Fig. 209.

(= Aristida obtusa Del.) 1.

Kortbeenboesmangras.

Compact and densely tufted;
to 600 mm tall. Leaf blades
10-250 mm long; to 1 mm wide.

Spikelets 11-12 mm long (ex-
cluding awns). Basal sheaths may
be glabrous or hairy, but not with densely matted, woolly
hairs; leaves markedly basal, often curved, glabrous or with
scattered long bulbous-based hairs; inflorescence usually
contracted, interrupted, much branched, branches bearing
spikelets to near the base; glumes pallid, firm, with
membranous apices and margins, glabrous, scabrid; lemma
articulation between the apex of the lemma and the base of
the column; column length variable, scabrid to and at the
branching point of the awns; only the central awn plumose;
callus 1.5 mm long, with a naked tip varying from pungent
to minutely bifid.

Flowering July to May. Mainly sandy soils in dry areas.
Common. Biome: Savanna, Nama-Karoo, Succulent Karoo

Fig. 209 . Stipagrostis obtusa

and Desert. Disjunct distribution, also in North Africa,
Sinai Peninsula and Iraq to Pakistan. Pasture (good fodder),
or erosion control (sand binder).

Description: De Winter 1965 (355), Stapf 1898-1900
(567). Illustration: Chippindall 1955 (fig. 269). Voucher:
Smook 4515, Oliver, Muller & Steenkamp 6615, Skarpe
5240. PRECIS code 990261 1-02400.

Stipagrostis proxima (Steud.) De Winter

( =Aristida proxima Steud.) 1.

Shrub or dwarf shrub; long
rhizomatous and tufted (erect); to
600 mm tall. Leaf blades to 100
mm long (usually less); setace-
ous, rolled. Spikelets 8-9 mm
long (excluding awns). Culms
and leaves with dense ap-
pressed hairs in the grooves, hairs becoming more dense,
longer and more obvious below the nodes; inflorescence
narrow, branches densely hairy, hairs longer and thicker on
swollen part just below spikelet; spikelets to 14 mm long
(including the awns); glumes firm, acute; upper glume 7-9
mm long; lemma articulation about in the middle of the
lemma; column short and stout; all three awns plumose
down to branching point and just below on the column;
callus 1. 5-2.0 mm long, narrow, very short pungent tip.

Flowering November. Sandy soils in disturbed areas.
Rare. Infrequent. Biome: Nama-Karoo. Endemic.

Description: De Winter 1965 (322), Stapf 1898-1900
(566), Chippindall 1955 (297). Voucher: Vorster 85,
Flanagan 1657. PRECIS code 990261 1-02500.

Stipagrostis ramulosa De Winter

Slender perennial; densely
tufted; 400-600 mm tall. Leaf
blades to 1 10 mm long. Spikelets
to 9 mm long (excluding awns).

Plants delicate; culms slender, to
1.2 mm wide, fascicled from the
nodes; leaves pungent, erect,
usually overtopping the inflores-
cence, lower surface rough,
densely covered with appressed prickles; inflorescence
elongate, narrow, sparsely branched with a few spikelets;
upper glume 7-9 mm long; lemma articulation near the
apex of the lemma; column very short; all three awns
plumose to the branching point of the awns, apices not
plumose; callus 2 mm long, narrow, with a naked pungent
tip.

Flowering November, January, and April. Sandy soils
between small dunes, in river beds, near water. Infrequent.
Biome: Nama-Karoo. Endemic. Pasture (grazed by game).

Description: De Winter 1965 (333). Voucher: Giess,

Volk & Bleissner 6294, sheet 1 & 2. PRECIS code
9902611-02600.

Stipagrostis sabulicola (Pilg.) De Winter

( =Aristida sabulicola
Pilg.) 1.

Namib dune bushman grass.

Reed-like shrub or dwarf
shrub; rhizomatous (rhizomes
robust and much branched), or
tufted (lax to densely); to 2000
mm tall. Leaf blades 250-600 mm long; to 3 mm wide
(folded). Spikelets 8-14 mm long (excluding awns). Culms
1 .2—5.0 mm wide, fasciculately branched from the nodes;
leaves erect, straight, rigid, pungent, overtopping the inflor-
escence; inflorescence elongate, narrow, spike-like, dense,
usually partly enclosed in the uppermost leaf sheath;

327

glumes straw-coloured, turning brown with age; lemma ar-
ticulation near apex of lemma; column short and stout; all
three awns plumose down to and around the branching
point, equal to subequal; callus 1. 5-2.0 mm long, tip pun-
gent but barely naked.

Flowering December to January. Dunetops, sandy
gullies and river beds. Locally common. Biome: Desert and
Succulent Karoo. Endemic. Domestic use (said to be plaited
into mats to cover huts).

Description: De Winter 1965 (331). Voucher: Coetzee
& Werger 1791a, Giess & Robinson 13233. PRECIS code
9902611-02700.

Stipagrostis schaeferi (Mez) De Winter

( =Aristida schaeferi Mez var.
biseriata Henr.) 1; ( =Aristida
schaeferi Mez var. schaeferi ) 1 .

Woolly leaved bushman grass.

Perennial; densely tufted
(cushion-forming), or rhizomat-
ous (rhizomes short, knotty); to
700 mm tall. Leaf blades to 100 mm long (basal leaves
shorter than cauline leaves); to 2 mm wide. Spikelets 12-15
mm long (excluding awns). Culm nodes glabrous; basal and
young sheaths usually woolly; leaves mainly basal; inflor-
escence narrow or open but pedicels always flexuous;
glumes firm, lower glume narrowly oblong to oblong, apex
obtuse to truncate, firm, glabrous or with rigid hairs; lemma
articulation near middle of lemma; column well developed,
scabrid; only the central awn plumose, hairs often
yellowish; callus 2 mm long, with a pungent, naked tip.

Flowering August to November, and March to June.
Gravelly soils in hollows in rocky outcrops, on gravel plains
and along dry watercourses. Locally common. Biome:
Nama-Karoo, Succulent Karoo and Desert. Endemic. Allied
to the more common and widespread S. ciliata var.
capensis, which has long spreading hairs at the nodes.

Description: De Winter 1965 (318). Voucher: Giess &
Van Vuuren 662. PRECIS code 9902611-02800.

Stipagrostis subacaulis (Nees) De Winter

(= Aristida subacaulis Nees
ex Steud.) 1.

Stemless bushman grass.

Annual; compactly tufted (to
prostrate and spreading); to 100
mm tall. Leaf blades to 30 mm
long; to 1 mm wide. Spikelets
15-16 mm long (excluding awns). Culms short, very poorly
developed; leaves mainly basal; inflorescence contracted,
dense, much branched, main axis not visible; glumes gla-
brous; lemma articulation between the apex of the lemma
and the base of the column; column very long, glabrous or
sometimes with a few scattered hairs shorter than 1.5 mm;
only central awn plumose; callus 0. 8-1.0 mm long, with a
naked, pungent tip.

Flowering January to November. Coarse sandy soils on
stony hillsides, and depressions on gravel flats, often in
soils with gypsum. Locally common. Biome: Succulent
Karoo and Desert. Northwards to southern Angola. Closely
allied to and often occuring with S. hermanii, which has the
column densely hairy on swollen area at branching point of
awns, and the callus 1.5 mm long.

Description: De Winter 1965 (373), Stapf 1898-1900
(568). Illustration: Chippindall 1955 (fig. 267). Voucher:
Giess 7844. PRECIS code 990261 1-02900.

Stipagrostis uniplumis (Licht.) De Winter var.

intermedia (Schweick.) De Winter

(-Aristida gracilior Pilg. var.
intermedia Schweick.) 1.

Annual; tufted; to 600 mm
tall. Leaf blades 100-200 mm
long; to 1 mm wide (flat or
folded). Spikelets 8-10 mm long
(excluding awns). Culms con-
spicuously striate; leaves main-
ly basal; inflorescence dense, branches long and flexuous;
glumes hairy, apex membranous; lemma densely
tuberculate around point of articulation, lemma articulation
between the apex of the lemma and the base of the column;
column well developed, long spreading hairs from
branching point down for a variable distance; central awn
distinctly plumose except in the lower 1/3; callus 0. 5-1.5
mm long, with long hairs at the junction between the lemma
and the callus and short hairs for the rest of its length, with
a naked, pungent tip.

Flowering March to June (rain dependent). Depressions
where water collects, on sandy and gravel plains. Infrequent
to common. Biome: Nama-Karoo and Desert. Endemic, or
possibly also in southern Angola. Somewhat intermediate
between S. uniplumis , which has a callus with short hairs
over the entire length, and S. hirtigluma , which has hairy
glumes and a distinct glabrous break between the short hairs
along the length of the callus and the long hairs at the
junction of the lemma and the callus. There are a number
of specimens that have been referred to as S. uniplumis X
5. hirtigluma , which are perennial, have hairy glumes and
a callus of the uniplumis type. De Winter (1963) refers to
these specimens as hybrids between S. uniplumis var. neesii
andS. hirtigluma subsp. patula.

Description: De Winter 1965 (359). Voucher: De Winter
& Hardy 8160, Giess 7848. PRECIS code 9902611-03000.

Stipagrostis uniplumis (Licht.) De Winter var. neesii

(Trin. & Rupr.) De Winter p| ^

(=Aristida uniplumis Licht.
var. neesii Trin. & Rupr.) 1 .

Perennial; laxly tufted (erect),
or rhizomatous (rhizomes short
and branched); to 900 mm tall.

Leaf blades to 160 mm long; set-
aceous. Spikelets 10.0-14.5 mm
long (excluding awns). Leaves

mainly basal; inflorescence narrow; spikelets few; glumes
10 mm or longer, glabrous; lemma tuberculate just below
the articulation point, articulation between the apex of the
lemma and the base of the column; column well developed
and plumed for the upper half of its length, with hairs longer
than 1.5 mm; only the central awn plumose, usually hairy
to the branching point but not the apex, diverging at right
angles from the column; callus 1 .0—1 .5 mm long, with short
hairs along the entire length and long hairs at the junction
of the lemma and the callus, with a naked, pungent tip.

Flowering December to May. Mainly gravel soils,
sometimes sandy soils on dolomite or over surface
limestone, and rocky slopes. Locally common. Biome: Sa-
vanna. Endemic. Pasture (good grazing). Grades into var.
uniplumis, which has more spikelets in the panicle, shorter
glumes and a straight central awn. Specimens with hairy
glumes have been refered to as hybrids between this var.
andS. hirtigluma subsp. patula.

Description: De Winter 1965 (358). Voucher: De Winter
793, Scheepers 1516. PRECIS code 990261 1-03100

328

Stipagrostis uniplumis (Licht.) De Winter var.

uniplumis

(=Aristida uniplumis Licht.
var .pearsonii Henr.) 1;

(=Aristida uniplumis Licht. var.
uniplumis) 1.

Silky bushman grass, blink-
aarboesmangras.

Perennial (to subperennial);
densely to laxly tufted (erect to geniculate); to 900 mm tall.
Leaf blades to 200 mm long; setaceous to 2 mm wide.
Spikelets 8-10 mm long (excluding awns). Culms simple
or branched well above the base; leaves mainly basal; in-
florescence narrow or effuse with many spikelets; glumes
usually shorter than 10 mm, glabrous; lemma tuberculate
around point of articulation, articulation between the apex
of the lemma and the base of the column; column well
developed, plumed in the upper half, with hairs longer than
1.5 mm; only the central awn plumose with hairs occuring
to 1/3 above or down to the branching point of the awns,
apex without hairs; callus 0.5-1 .2 mm long, with short hairs
along the entire length and long hairs at the junction of the
lemma and the callus, with a pungent, naked tip.

Flowering December to May. Usually on sandy soils and
disturbed areas and floodplains. Common. Biome: Savanna,
Nama-Karoo, and Desert. Zimbabwe, Angola, Uganda,
Somalia to Senegal. Valuable pasture. A variable taxon,
which grades into var. neesii, which has a panicle with
fewer spikelets. Hybridizes with S. hirtigluma (De Winter
5710).

Description: Chippindall & Crook 1976 (56), De Winter
1965 (359), Stapf 1898-1900 (569). Illustration: Muller
1985 (fig. 129). Voucher: Acocks 1938. PRECIS code
9902611-03200.

Fig. 206.

Stipagrostis zeyheri (Nees) De Winter subsp. barbata

(Stapf) De Winter

( =Aristida capensis Thunb.
var. barbata Stapf) 1 .

Cape bushman grass.

Robust perennial; densely
tufted (erect), or rhizomatous
(rhizomes short and knotty); to
900 mm tall. Leaf blades to 500
mm long; setaceous. Spikelets 16-18 mm long (excluding
awns). Culms 1. 2-5.0 mm wide; leaf blades flaccid, often
curling, almost overtopping the inflorescence; inflores-
cence contracted and dense; glumes glabrous, pallid, slight-
ly darker at the base; lemma articulation between the apex
of the lemma and the base of the column; column with long
hairs below the branching point of the awns; all three awns
plumose to the branching point and excurrent into a long
naked apex; callus 2. 0-2. 5 mm long, with a distinct naked
tip.

Flowering January to May (and July, October and
November). Coastal dunes. Locally common. Biome: Fyn-
bos and Savanna. Endemic. Not well differentiated from
subsp. zeyheri , which is apparently limited to the winter
rainfall area.

Description: De Winter 1965 (346), Stapf 1898-1900
(565). Voucher: Acocks 13588, Smook 1862. PRECIS code
9902611-03300.

Fig. 210. Stipagrostis zeyheri subsp. se^icans

329

Stipagrostis zeyheri (Nees) De Winter subsp. macropus

(Nees) De Winter

( =Aristida capensis Thunb.
var. genuina Henr.) 1;

(-Aristida capensis Thunb. var.
macropus (Nees) Trin. & Rupr.)

1; (=Aristida capensis) Thunb.,
non Stipagrostis capensis Nees) 1

Bushman grass.

Robust perennial; shortly rhizomatous; to 720 mm tall.
Leaf blades to 200 mm long; setaceous. Spikelets 17-18
mm long (excluding awns). Culms 1. 2-5.0 mm wide; leaves
mainly basal; inflorescence open, divaricate, with spikelets
at end of the branches; glumes usually dark, longer than 15
mm, glabrous; lemma articulation between the apex of the
lemma and the base of the column; column smooth, gla-
brous; all three awns plumose to the branching point and
excurrent into a naked apex, hairs dirty-white to golden;
callus 2. 0-2. 5 mm long, with a distinct, naked, pungent tip.

Flowering August to November. Sandy soils and old
cultivated lands. Locally common. Biome; Fynbos and Suc-
culent Karoo. Endemic. Reported to hybridize with S.
ciliata var. capensis (Acocks 14817). This hybrid was
assigned the name Aristida schlechteri by Henrard, but was
not accepted by De Winter.

Description: De Winter 1965 (346), Stapf 1898-1900
(565). Voucher; Davidse 3331 1, Van Breda 775. PRECIS
code 9902611-03400.

Stipagrostis zeyheri (Nees) De Winter subsp. sericans

(Hack.) De Winter

(=Aristida capensis Thunb.
var. dieterleniana Schweick.) 1;

( =Aristida sericans Hack, apud
Schinz ) 1 .

Robust perennial; densely
tufted (erect); to 750 mm tall.

Leaf blades to 300 mm long; set-
aceous, rolled. Spikelets to 14
mm long (excluding awns). Culms 1 .2-5.0 mm wide; inflor-
escence extending well above the leaves, usually narrow,
never effuse, with few spikelets; glumes pallid, long hairy;
lemma articulation at the apex of the lemma or absent;
column stout, with long hairs for some distance below the
branching point; all three awns plumose to the branching
point of the awns, excurrent into a naked apex; callus
1. 0-1.5 mm long, naked, with a pungent tip.

Flowering January to May. Sandy soils on rocky
outcrops and disturbed areas such as old lands. Infrequent
to locally common. Biome: Savanna and Grassland.
Endemic.

Description: De Winter 1965 (346 & 349). Voucher:
Smook 6358, Ferreira F213. PRECIS code 990261 1-03500.

Fig. 210.

Stipagrostis zeyheri (Nees) De Winter subsp. zeyheri

PI. 190.

(=Aristida capensis Thunb.
var. canescens Trin. & Rupr.) 1.

Robust perennial; shortly
rhizomatous (knotty), tufted
(erect); to 750 mm tall. Leaf
blades to 500 mm long; setace-
ous, folded. Spikelets 16—19 mm
long (excluding awns). Culms
1. 2-5.0 mm wide; leaves erect and rigid; inflorescence
open, divaricate, branches and pedicels flexuous with the
spikelets at the end of the branches; glumes glabrous; lem-
ma articulation between the apex of the lemma and the base
of the column; column long-hairy; all three awns plumose
to the branching point of the awns, excurrent into a naked

apex; callus 2 mm long, with a naked, pungent tip.

Flowering October to March (and May and July). Sandy
slopes and limestone hills, disturbed areas, especially on
recently burnt areas. Locally common. Biome: Fynbos.
Endemic. Not well differentiated from subsp. barbata,
which is common along the east coast of the Cape.

Description: Stapf 1898-1900 (563). Voucher: Taylor
5602. PRECIS code 990261 1-03600.

Streblochaete Pilg.

Koordersiochloa Merr., Pseudostreptogyne A. Camus.

Perennial; caespitose. Culms 300-1000 mm high; herba-
ceous; unbranched above. Sheath margins joined. Leaf
blades linear-lanceolate\ rolled. Ligule an unfringed
membrane.

Inflorescence paniculate; open; espatheate. Spikelet-
bearing axes persistent.

Spikelets 16-28 mm long; not noticeably compressed ;
disarticulating above the glumes. Glumes present; two; very
unequal; markedly shorter than the spikelets; awnless;
similar (narrow, membranous-herbaceous with hyaline
margins). Incomplete florets distal to the female-fertile
florets, merely underdeveloped (male or sterile); proximal
incomplete florets absent.

Female-fertile florets 2-6. Lemmas similar in texture to
the glumes to decidedly firmer than the glumes; 7 nerved;
incised (shortly so, to nearly entire); awned. Awns 1;
median; dorsal; geniculate (the very long awns intertwine,
so that the spikelet is dispersed as a unit); much longer than
the body of the lemma (distally filiform). Palea present;

Fig. 211. Streblochaete longiarista

330

conspicuous but relatively short (about half the lemma
length). Lodicules 2; membranous; ciliate, or glabrous.
Stamens 3. Ovary glabrous; hilum short; embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 10. Pooideae; Poodae; Meliceae. 1 species.
Tropical Africa, Java, Lombok, Phdippines. Montane.
Mesophytic; in shade; glycophytic. Natal and Cape
Province. 1 indigenous species.

References. 1. Clayton. 1970. FTEA.

Species treatment by G.E. Gibbs Russell.

Streblochaete longiarista (A. Rich.) Pilg.

Fig. 211. PI. 191.

Perennial; loosely tufted;

300-900 mm tall. Leaf blades to
250 mm long; 3-1 1 mm wide
(narrowed towards base). Leaf
sheaths tubular; callus sharp, 2-3
mm long, with dense short white
hairs; lemma with a fine twisted
awn, to 40 mm long, that tangles
with other awns in the panicle.

Flowering April to May. Open places in mountain
forests. Rare. Biome: Forest. Through east African
highlands to Ethiopia, also Reunion, Java and the
Phillipines. Apparently very dissimilar from its relative,
Melica, but classified in the same tribe because of lodicule
and chromosome similarities.

Description: Chippindall 1955 (82), Clayton et al.
1970-1982 (74). Illustration: Chippindall 1955 (fig. 53),
Clayton et al. 1970-1982 (fig. 25). Voucher: Chippindall
356. PRECIS code 9901971-00100.

Styppeiochloa De Winter

Sometimes included in Crinipes Hochst.

Perennial ; densely caespitose (the hard, fibrous basal
sheaths forming tough, fire-resistant mats). Culms 100-700
mm high; herbaceous; unbranched above (wiry, the nodes
hidden at the base). Leaf blades linear; to 1 mm wide\
setaceous (resembling the culms); rolled (convolute).
Ligule a fringe of hairs.

Inflorescence paniculate ; contracted (scanty, the
spikelets appressed to the panicle branches); espatheate.
Spikelet-bearing axes persistent.

Spikelets compressed laterally; disarticulating above the
glumes. Callus short. Hairy callus present. Glumes two;
very unequal, or more or less equal; markedly shorter than
the spikelets; short awned (or aristate from the excurrent
mid-nerve), or awnless; similar (lanceolate, apices 3-lobed
or acute/entire). Incomplete florets distal to the female-
fertile florets, merely underdeveloped, awned, or awnless;
proximal incomplete florets absent.

Female-fertile florets 2-5. Lemmas similar in texture to
the glumes (membranous); hairy (at margins, near base);
without a germination flap; 3-5 (-7) nerved; incised;
mucronate to awned (the three lobes with awns or mucros).
Awns, when present, 3\ median, or median and lateral (via
shortly excurrent nerves). The median awn similar in form
to the laterals (when laterals present); apical, non-genicu-
late; much shorter than the body of the lemma. Palea
present (narrowly lanceolate); relatively long (equalling the
lemma); 2-nerved. Stamens 3. Ovary glabrous. Fruit small
(about 2 mm long); fusiform; hilum long-linear; embryo
small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 2 species. South and southeastern tropical
African mountains. Helophytic to mesophytic; in open
habitats (where there is impeded drainage, in mountains);
glycophytic. Transvaal, Orange Free State, Natal, and Cape
Province. 1 indigenous species.

References. 1. De Winter. 1966. Bothalia 9: 134.

Species treatment by N.P. Barker.

Styppeiochloa gynoglossa (Goossens) De Winter

Fig. 212. PI. 192.

{=Crinipes gynoglossa
Goossens) 1.

Perennial; mat forming to tuft-
ed; 100-700 mm tall. Leaf blades
100-400 mm long; to 1.2 mm
wide. Spikelets 5-7 mm long.

Sheaths and leaves split with age,
forming a dense, fibrous base;
spikelets 2-5-flowered, the uppermost floret sterile or
reduced; glumes 1-3-nerved, unequal, 3 central nerves
produced into a short awn; lemmas basally pubescent with
trilobed apex, 3-5(-7)-nerved, with middle lobe extending
into a short awn.

331

Flowering September to January (and even later to the
north). Rock crevices and seepage areas over rocks; high
rainfall areas (800 mm or more) at high altitudes. Locally
dominant (in seepage areas). Biome: Afromontane.
Zimbabwe and southern Mozambique at altitudes as low as
610 m. Chippindall (1955) distinguishes a distinct variety
from the Natal Drakensberg. These specimens apparently
have larger spikelets and almost glabrous lemmas.

Description: De Winter 1966 (135), Goosens 1934 Kew
Bull. (200), Chippindall 1955 (123-124). Illustration:
Goosens 1934 Kew Bull. (200), Chippindall 1955 (fig. 96).
Voucher: Ellis 3288. PRECIS code 9903504-00100.

Tarigidia Stent

Perennial (glaucous); caespitose. Culms 800-1500 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear; flat (margins thickened). Ligule an
unf ringed membrane. Plants bisexual, with bisexual
spikelets. The spikelets of sexually distinct forms on the
same plant (in that some spikelets at the raceme bases may
be sterile), or all alike in sexuality.

Inflorescence paniculate ; contracted; espatheate .
Spikelet-bearing axes disarticulating (at least the lower
branches do so)', falling entire.

Spikelets clustered or in pairs on lower panicle branches.
Female fertile spikelets 4-4.5 mm long; abaxial; com-
pressed dorsiventrally; falling with the glumes. Glumes
two; very unequal (G1 sometimes much reduced), or more
or less equal (usually); awnless. Proximal incomplete
florets I ; epaleate; sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes; not becoming indurated; hairless; having the
margins lying flat and exposed on the palea; without a ger-

Fig. 213. Tarigidia aequiglumis

mination Hap; entire; awnless. Palea present; relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous.

Photosynthetic pathway. C4; XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Panicoideae; Pani-
codae; Paniceae. 1 species. Southern Africa. Mesophytic to
xerophytic; in open habitats (dry grassland); glycophytic.
Namibia, Transvaal, Orange Free State, and Cape Province.
1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA.

Species treatment by G.E. Gibbs Russell.

Tarigidia aequiglumis (Goossens) Stent

Fig. 213. PI. 193.

Perennial; densely tufted;

800-1500 mm tall. Leaf blades to
350 mm long; 3-5 mm wide.

Spikelets 4.0^4. 5 mm long. In-
florescence spike-like or narrowly
conical, with short branches
appressed or somewhat spreading
below; spikelets woolly; glumes
equal, about 2/3 of spikelet
length.

Flowering January to May. Open veld or among rocks.
Rare. Biome: Savanna and Grassland. Endemic. Similar in
general appearance to Anthephora , which does not have in-
florescence branches, and the spikelets are similar to
Digitaria, which has very small lower glumes and the upper
glumes less than 1/4 the spikelet length.

Description: Chippindall 1955 (422). Voucher: Dinter
5589. PRECIS code 9900891-00100.

Tetrachne Nees

Perennial; forming large tufts, the shoots crowded on a
short, oblique rhizome. Culms 300-1000 mm high; herba-
ceous; branched above to unbranched above. Leaf blades
linear; usually rolled. Ligule a fringe of hairs (dense).

Inflorescence of spike-like main branches ( the few to
many branches appressed, short, dense)', espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary; biseriate (on one side of rachis,
crowded); 4-6 mm long; compressed laterally; falling with
the glumes; not disarticulating between the florets. Glumes
two; more or less equal; decidedly shorter than the adjacent
lemmas; awnless; the keels somewhat winged; similar (thin,
acute, the lower smaller). Incomplete florets both distal and
proximal to the female-fertile florets; distal florets merely
underdeveloped, awnless; proximal incomplete florets 2.

Female-fertile florets 3-5. Lemmas similar in texture to
the glumes; without a germination flap; 5 nerved; entire;
awnless. Palea present; relatively long (equalling the
lemma). Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small (2.5 mm long); fusiform; hilum short;
pericarp fused, or loosely adherent (removable with
difficulty after soaking); embryo large (about 2/3 the length
of the fruit).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines fairly even. PCR cell
chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 1
species. South Africa and Pakistan. Mesophytic (often in
alluvial soil); in open habitats (in high altitude grassland);
glycophytic. Orange Free State, Lesotho, and Cape
Province. 1 indigenous species.

References. I. Chippindall. 1955. Gr. & Past.

Species treatment by M. Koekemoer.

332

Tetrachne dregei Nees

Fig. 214. PI. 194.

Robies cocksfoot, kropaar-
gras.

Perennial; rhizomatous and
tufted; 320-860 mm tall. Leaf
blades 50-125 mm long; to 1 mm
wide. Spikelets 4-6 mm long.

Base robust or woody; culms
branching freely; leaves curly;
spikes more than 4, stout, 10-40 mm long, not spreading
far from, the central axis, more or less their own length
apart.

Flowering November to March. Sandy soil on
riverbanks, rocky outcrops or mountain slopes, at altitudes
higher than 1250 m. Infrequent to locally common. Biome:
Grassland and Nama-Karoo. Pakistan. Pasture (mostly in
natural veld but also cultivated on small scale, semi-
procumbent and forming dense stands when grazed).

Description: Stapf 1898-1900 (710), Chippindall 1955
(188). Illustration: Chippindall 1955 (fig. 164). Voucher:
Smook & Gibbs Russell 2186. PRECIS code 9903270-
00100.

Tetrapogon Desf.

Codonachne Steud., Cryptochloris Benth.,
Lepidopironia A. Rich.

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 130-850 mm high; herbaceous. Leaf blades linear
(tapered); folded (usually). Ligule a fringed membrane
(very narrow ).

Inflorescence a single spike, or of spike-like main
branches (1-3 upright racemes or spikes, of which 2 may
be partly or completely fused along their backs)', digitate
or subdigitate, or non-digitate; espatheate. Spikelet-bearing
axes persistent.

Spikelets solitary, or in pairs; secund (the rachis one-
sided); biseriate; subsessile; 2.5-12 mm long (cuneate);
compressed laterally; disarticulating above the glumes (the
glumes persistent); not disarticulating between the florets,
or disarticulating between the florets (but under the fertile
florets only). Hairy callus present. Glumes two; relatively
large; more or less equal; long relative to the adjacent
lemmas; awned (awn-tipped), or awnless; similar
(lanceolate, long-pointed, subhyaline). Incomplete florets
distal to the female-fertile florets (1 — several, awned);
proximal incomplete florets absent.

Female-fertile florets 2-7 . Lemmas decidedly firmer
than the glumes (herbaceous, leathery, the margins
hyaline); 3-5 nerved; entire (truncate), or incised; awned.
Awns 1; median; dorsal ; non-geniculate; much longer than
the body of the lemma. Palea present; relatively long. Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small (1.5-3 mm); ellipsoid; hilum short; pericarp free;
embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x - 10. Chloridoideae; Chlorideae sensu lato. 5-6
species. Mediterranean to India, tropical and South Africa.
Helophytic to mesophytic; in shade, or in open habitats
(savanna); glycophytic. Namibia and Transvaal. 1
indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

Tetrapogon tenellus (Roxb.) Chiov.

Fig. 215. PI. 195.

Shortlived perennial, or an-
nual; loosely tufted (culms erect
or ascending); 130-750 mm tall.

Leaf blades 100-240 mm long;

2-4 mm wide. Spikelets 3. 5-5.0
mm long. Spikes solitary (seldom
paired); spikelets 5-6-flowered;
glumes persistent, 3-5 mm long;
lemma coriaceous, 4. 0-6. 5 mm
long, keeled; awns 3-1 1 mm long.

Flowering January to April. Usually in the shade in open
or dense bushveld, often in disturbed rocky soil and on
limestone. Infrequent. Biome: Savanna. Eastern Africa to
India.

Description: Chippindall 1955 (198), Clayton et al.
1970-1982 (348). Voucher: Van der Schijff 5204. PRECIS
code 9903220-00200.

333

Fig. 215. Tetrapogon tenellus

Thamnocalamus Munro

Himalayacalamus Keng f.

Perennial; caespitose. Culms 1000-5000 mm high;
woody and persistent. Culms reaching 20 mm in diameter;
culms branched above. Leaf blades pseudopetiolate. Ligule
a fringed membrane.

Inflorescence a single raceme, or paniculate ;
contracted; spatheate.

Spikelets solitary; 15-18 mm long; not noticeably com-
pressed. Glumes two; more or less equal; long relative to
the adjacent lemmas; awnless (upper glume pointed);
similar. Proximal incomplete florets absent.

Female-fertile florets 2-8. Lemmas entire; awnless, or
mucronate (?); 10-11 nerved. Palea present; relatively long;
with several nerves. Lodicules 3; membranous; ciliate.
Stamens 3. Ovary glabrous; stigmas 3.

Transverse section of leaf blade. Mesophyll with arm
cells; with fusoids. Midrib usually with one bundle only
(complex in T. afistatus).

Cytology, classification, distribution. Chromosome base
number, x = 12. Bambusoideae; Bambusodae; Bambuseae.
6 species. Eastern Asia, South Africa. Helophytic, or meso-
phytic; glycophytic. Orange Free State, Natal, Lesotho, and
Cape Province. 1 indigenous species.

References. 1. Soderstrom & Ellis. 1982. Bothalia 14:
53.

Species treatment by G.E. Gibbs Russell.

Fig. 216. Thamnocalamus tessellatus

Thamnocalamus tessellatus (Nees) Soderstrom & Ellis

Fig. 216. PI. 196.

( =Arundinaria tessellata
(Nees) Munro) 1 .

Bamboo; rhizomatous (rhi-
zomes stout, woody); 1000-5000
mm tall (culms to 20 mm in dia-
meter, profusely branched a-
bove). Leaf blades 50-150 mm
long; 8-15 mm wide. Spikelets
16-18 mm long. Leaf blades with conspicuous cross-veins.

Flowering in local populations after many years,
followed by death. Mountainsides, in wet places and

334

sheltered ravines, 1600-2700 m. Locally common. Biome:
Afromontane. Endemic.

Description: Chippindall 1955 (30), Soderstrom &
Ellis. 1982. (53). Illustration: Chippindall 1955 (fig. 1),
Soderstrom & Ellis. 1982. (fig. 1 & 2). Voucher:
Soderstrom & Du Toit 1610. PRECIS code 9904570-
00100.

Thelepogon Roth.

Rhiniachne Steud..

Annual (rather stout, erect or decumbent, often with
prop roots). Culms 100-1500 mm high: herbaceous;
branched above, or unbranched above. Leaf blades
lanceolate (from the amplexicaul base); cordate ; flat. Ligule
an unfringed membrane, or a fringed membrane. Plants
bisexual, with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant (hermaphrodite/sterile, but
the latter reduced to their pedicels).

Inflorescence of spike-like main branches (of long,
brittle golden ‘racemes' ); digitate or subdigitate (on a short
common axis); espatheate; not comprising ‘partial inflores-
cences’ and foliar organs. Spikelet-bearing axes ‘racemes’
(with numerous spikelets); clustered; with substantial
rachides; disarticulating at the joints. ‘Articles’ without a
basal callus-knob.

Spikelets in pairs (but the pedicellate one reduced to the
pedicel); consistently in ‘long-and-short’ combinations;
these pedicellate/sessile. Pedicels free of the rachis (the
pedicel and joint separated below, contiguous above). The
sessile spikelets hermaphrodite. The pedicellate spikelets
sterile (reduced to the pedicels). Female-fertile spikelets
5-13 mm long; compressed dorsiventrally; falling with the
glumes (deciduous with joint and sterile pedicel). Hairy
callus present (callus annular, short, ciliate). Glumes two;
more or less equal; awnless; very dissimilar. Lower glume
not two-keeled. Proximal incomplete florets 7; paleate,
palea fully developed; male.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); incised; awned. Awns 1; median; from
the sinus; geniculate; much longer than the body of the
lemma. Palea present; relatively long. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit small (about 3
mm long); ellipsoid; hilum short; embryo large (about half
the length of the fruit).

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Andropogoninae. 1
species. Tropical Africa, Asia. Helophytic to mesophytic;
in open habitats (seasonally wet, heavy soils and disturbed
ground); glycophytic. Namibia. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Thelepogon elegans Roem. & Schult.

Fig. 217. PI. 197.

Coarse annual; 100-1500 mm
tall. Leaf blades 40-200 mm
long; 5-30 mm wide (base
cordate, clasping). Spikelets (ses-
sile) 5-13 mm long (pedicellate
somewhat longer). Lower glume
of sessile spikelet prominently
rugose.

Flowering May. Black turf
soil. Biome: Savanna. Tropical Africa to Indonesia.

Description: Clayton et al. 1970-1982 (744).

Illustration: Clayton et al. 1970-1982 (fig. 174). Voucher:
Ellis 3742. PRECIS code 99001 10-00100.

Fig. 217. Thelepogon elegans

Themeda Forssk.

Androscepia Brong., Anthistiria L. f., Aristaria Jungh.,
Heterelytron Jungh., Perobachne Presl.

Annual, or perennial; caespitose (coarse, very rarely
stoloniferous). Culms 300-3100 mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear;
flat, or folded. Ligule an unfringed membrane to a fringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets of sexually distinct forms on the same plant (male
or neuter, and hermaphrodite ); overtly heteromorphic.

Inflorescence paniculate (leafy, comprising short
racemes in spatheate, hard-to-interpret clusters ); open;
spatheate; a complex of ‘partial inflorescences’ and inter-
vening foliar organs (composed of short racemes in
spatheate clusters: each cluster terminated by 1-3 pairs of
spikelets, one of each pair sessile and bisexual, the other
pedicellate and male-or-sterile (or a triplet of 1 terminal
sessile spikelet with 2 pedicellate ones), the whole
surrounded by a whorl of 4 male or sterile, sessile spikelets
constituting an involucre). Spikelet-bearing axes very much
reduced ; clustered (racemes solitary in their spatheoles,
these units in groups of three or more in short capituliform
glomerules); disarticulating at the joints (racemes
disarticulating at the level of the fertile spikelets).

Spikelets associated with bractiform involucres
(constituted by the four imperfect spikelets ); in pairs and in
triplets; consistently in ‘long-and-short’ combinations;
these pedicellate/sessile. Pedicels free of the rachis. The
sessile spikelets hermaphrodite. The pedicellate spikelets

335

Fig. 218. Themeda triandra

male-only, or sterile. Female-fertile spikelets not noticeably
compressed to compressed dorsiventrally; falling with the
glumes (clusters disarticulate immediately above
involucres). Glumes two; more or less equal; awnless;
leathery. Proximal incomplete florets 7; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline, stipitate beneath the awn); entire (usually);
awned. Awns 1; median; apical (usually); geniculate; much
shorter than the body of the lemma, to much longer than
the body of the lemma. Palea present, or absent; when
present conspicuous but relatively short, or very reduced.
Lodicules 2; fleshy; glabrous. Stamens 3. Ovary glabrous.
Fruit small; hilum short; embryo large.

Cytology, classification, distribution. Chromosome base
number, a: = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 18 species. Warm
Africa, Asia, Australia. Mesophytic; in open habitats
(savanna); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Swaziland, Natal, Lesotho, and Cape
Province. 1 indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA. 2.
PRE herbarium practice, following Smook & Gibbs
Russell.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

Themeda triandra Forssk.

Fig. 218. PI. 198. PI. 199.

(=7\ triandra Forssk. var.
burchellii (Hack.) Stapf) 1; (=7\
triandra Forssk. var. hispida
(Nees) Stapf) 1; (=7. triandra
Forssk. var. imberbis (Retz.) A.

Camus) 1; (= T . triandra Forssk.
var. trachyspathea Goossens) 1;

(=7’. triandra Forssk. var.
vulgaris auctt., non Hack.) 2.

Rooigras.

Perennial; rhizomatous; 300-1500 mm tall. Leaf blades
to 300 mm long; 1-8 mm wide. Spikelets (sessile) 5-7 mm
long (pedicellate equalling it or somewhat longer). Leaf
sheaths compressed; ligule a notched membrane; blade tips
abruptly or gradually tapering; spikelets awned, in drooping
triangular clusters with reddish spathes and bractlike sterile
spikelets.

Flowering September to June. Undisturbed veld. Widely
dominant. Biome: Fynbos, Savanna, Grassland, and Nama-
Karoo. Old World tropics and subtropics. Pasture (natural
veld). Although Themeda triandra is extremely variable,
the characters used to separate the traditional varieties are
poorly correlated with other attributes such as distribution,
habitat and chromosome number. Some forms may
vegetatively resemble Heteropogon contortus , which has an
undivided ligule and blunt often hooded leaf blades, or
Schizachyrium sanguineum , which has a strongly curved
undivided ligule, abruptly pointed leaf blades, and a reddish
colour.

Description: Chippindall 1955 (490), Clayton et al.
1970-1982 (829). Illustration: Chippindall 1955 (pi. 18),
Clayton et al. 1970-1982 (fig. 192), Flower. PI. Afr. (44:
1741). Voucher: De Winter 2748. PRECIS code 9900830-
00100.

336

Thinopyrum A. Loeve

Sometimes included in Elytrigia Desv. or Elymus L.

Perennial (rigid, erect); long-rhizomatous. Culms
250-700 mm high; herbaceous; unbranched above. The
shoots aromatic. Leaf blades linear (glaucous); rolled
(involute). Ligule an unfringed membrane.

Inflorescence a single spike (the spikelets usually
oppressed)', espatheate. Spikelet-bearing axes disarticu-
lating (fragile); disarticulating at the joints (the spikelets
falling with the internode below).

Fig. 219 .Thinopyrum distichum

Spikelets solitary; distichous; compressed laterally;
falling with the glumes. Glumes two; more or less equal
(subequal); markedly shorter than the spikelets; awnless;
similar. Incomplete florets distal to the female-fertile
florets, merely underdeveloped; proximal incomplete
florets absent.

Female-fertile florets 2-10. Lemmas similar in texture
to the glumes (leathery); 5 nerved; entire; awnless. Palea
present; relatively long. Lodicules 2; membranous; ciliate.
Stamens 3. Ovary hairy. Fruit medium sized; hilum long-
linear; embryo small.

Cytology, classification, distribution. Chromosome base
number, x - 7. Pooideae; Triticodae; Triticeae. 5 species.
Coasts of Europe. Xerophytic; in open habitats; maritime-
arenicolous. Cape Province. 1 indigenous species.

Intergeneric hybrids with Leymus and Elytrigia.

References. 1. Chippindall. 1955. Gr. & Past. 2. Dewey.
1984. Genomic classification in Gustafson, Gene
manipulation: 209.

Species treatment by M. Koekemoer.

Thinopyrum distichum (Thunb.) Loeve

Fig. 219. PI. 200.

(-Agropyron distichum
(Thunb.) Beauv.) 1 .

Strandkoringgras, coastal
wheat grass.

Hard, robust perennial; stolon-
iferous (underground parts thick,
creeping and profusely rooted at
the nodes); 400-600(-900) mm tall. Leaf blades
200^100(-500) mm long; 5-7 mm wide. Spikelets
( 1 5— )28— 40 mm long. Culms often branched below with
tufts of leaves from the nodes; leaves flat at first and then
rolled, rigid, sharp pointed; spike 60-250 mm long, rachis
breaking up easily; spikelets 5-1 1 -flowered, hard and
smooth, arranged alternately, appressed to rhachis.

Flowering October to January. On coastal sand dunes,
usually in areas exposed to seawinds and saltspray; it can
also tolerate inundation by spring tides. Locally common.
Endemic. Food and drink (culms chewed by people for
juicy sweet sap), or erosion control (efficient sand binder,
also used in reclamation work).

Description: Stapf 1898-1900 (743), Chippindall 1955
(69). Illustration: Chippindall 1955 (fig. 41). Voucher:
Moffett 3816. PRECIS code 9904348-00200.

Trachypogon Nees

Homopogon Stapf.

Perennial (very rarely annual); caespitose. Culms
300-2000 mm high; herbaceous (with slender culms); un-
branched above. Leaf blades linear; usually rolled, or flat
(sometimes). Ligule an unfringed membrane. Plants
bisexual , with bisexual spikelets. The spikelets of sexually
distinct forms on the same plant ( hermaphrodite and male
or neuter J; overtly heteromorphic (male or neuter spikelets
without callus, awnless); alt in heterogamous combina-
tions.

Inflorescence of a single raceme (or up to 5 racemes)',
digitate or subdigitate, or non-digitate; espatheate; not com-
prising ‘partial inflorescences’ and foliar organs. Spikelet-
bearing axes ‘racemes’ (long, terminating the culms);
solitary, or paired, or clustered (up to 5 ‘racemes’);
persistent (but joints articulated and usually shortly
bearded ).

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations (in which, however, the usual pattern of sexuality
is inverted); unequally pedicellate in each combination.
Pedicels free of the rachis. The short-pedicellate spikelets
male-only, or sterile, persistent, sometimes dorsally
flattened, without a callus, sometimes awnless, LI sterile.
The long-pedicellate spikelets hermaphrodite. Female-
fertile spikelets not noticeably compressed (cylindrical);
falling with the glumes (falling from pedicels). Glumes two;
relatively large; more or less equal; awnless; very dissimilar
(G1 firmer, convolute, 2-keeled; G2 thinner, channelled on
each side of the rounded keel). Proximal incomplete florets
1 ; epaleate; sterile.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline basally, but becoming stipitate-
cartilaginous above); entire; awned. Awns 1; median; api-
cal; geniculate; much longer than the body of the lemma.
Palea present, or absent; when present very reduced. Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous.
Embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5, or 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. About 13 species.
Tropical America and Africa, Madagascar. Mesophytic; in
open habitats (savanna); glycophytic. Namibia, Botswana,

337

Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell & M.
Koekemoer.

Trachypogon spicatus (L. f.) Kuntze

(=77 capensis (Thunb.)

Trin.) 1.

Giant spear grass, reuse pyl-
gras.

Perennial; rhizomatous and
tufted; 300-1200 mm tall. Leaf
blades 50-200 mm long; to 5 mm
wide. Spikelets (short-pedicel-
late) 6-8 mm long (long-pedi-
cellate slightly longer). Culm nodes with ring of hairs;

ligule membranous, splitting into three lobes; inflorescence
a single raceme (rarely up to 5 racemes) with velvety awns
throughout its length.

Flowering October to May. Bushveld and sourveld.
Common. Biome: Fynbos, Savanna, and Grassland. Africa
and tropical America. Urelytrum agropyroides and
Heteropogon contortus also have large, awned, single-
raceme inflorescences, but they lack the hairy culm nodes
of Trachypogon.

Description: Chippindall 1955 (494), Clayton et al.
1970-1982 (709). Illustration: Chippindall 1955 (PI. 20),
Clayton et al. 1970-1982 (fig. 163), Flower. PI. Afr. (38:
1512). Voucher: Ward 2778. PRECIS code 9900780-
00100.

Tragus Haller

Annual, or perennial; long-stoloniferous, or decumbent
(usually creeping). Culms 50-650 mm high; herbaceous.
Leaf blades flat. Ligule a fringed membrane (very narrow),
or a fringe of hairs. The spikelets of sexually distinct forms
on the same plant (with one or more members of the cluster
reduced), or all alike in sexuality.

Inflorescence a false spike, with clusters of spikelets on
reduced axes (a spicate raceme of crowded glomerules. the
latter very shortly- or rarely long- peduncled, with 2-5
spikelets ); espatheate. Spikelet-bearing axes disarticulating;
falling entire (the clusters falling whole).

Female-fertile spikelets 2-5 mm long; compressed dor-
siventrally, falling with the glumes (in the cluster). Hairy
callus absent. Glumes one per spikelet, or two; very
unequal, or not applicable (G 1 much reduced or absent);
long relative to the adjacent lemmas (G2 equalling the
spikelet); awnless; very dissimilar (lower tiny, scarious or
absent, upper large, hard, with 5 rows of hooked spines on
the back). All florets female-fertile; proximal incomplete
florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes (membranous); 3 nerved; entire; awnless. Palea
present. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small; hilum short; pericarp fused; embryo
large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
present. Maximum number of extension cells 1. PCR cell
chloroplasts with well developed grana; centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 7
species, 6 in warm Africa, 1 pantropical. Mesophytic to
xerophytic; in open habitats (often in disturbed ground);
glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 4 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Anton.
1981. Kew Bull. 36:55. 3. Clayton et al. 1974. FTEA.

Species treatment by G.E. Gibbs Russell.

1(0). Inflorescence appearing branched; peduncles of
spikelet clusters about same length as spikelets;
upper glume of lowest spikelet in each cluster
longer than 6 mm; glume hairs straight, never

curved or hooked at tip T. pedunculatus

Inflorescence appearing spike-like; peduncles of
spikelet clusters much shorter than spikelets; upper
glume of lowest spikelet in each cluster shorter than
5 mm; glume hairs curved or hooked at tip .... 2
2(1). Glume hairs curved, not hooked at tip; plant a
stoloniferous perennial; anthers 1.8-2.5 mm long

T. koelerioides

Glume hairs hooked at tips; plant annual, culms

Fig. 220. PI. 201.

338

sometimes rooting at lower nodes; anthers less than

1 mm long 3

3(2). Lowest glumes with 5 nerves; lowest spikelet in each
cluster 2. 0-3. 8 mm long; anthers 0.4-0. 6 mm long

T. berteronianus

Lowest glumes with 7 nerves; lowest spikelet in each
cluster 3. 5-5.0 mm long; anthers 0.6-0. 8 mm long
T. racemosus

Tragus berteronianus Schult.

Small carrot-seed grass,
kousklits.

Fig. 221. PI. 202.

Annual; loosely tufted; 50-
600 mm tall. Leaf blades 10-60
mm long; 2-5 mm wide. Spike-
lets (lowest in each cluster)

2. 0-3. 8 mm long. Inflorescence
narrowly spikelike; spikelets
clustered on peduncles that are much shorter than the spike-

lets; lowest glumes 5-nerved, spaces between nerves 2-5
times wider than the nerves (in the middle, tapering sharply
at their ends); glume hairs hooked at tips; anthers 0.4-0.6
mm long.

Flowering throughout the year (most commonly in
summer). Disturbed places. Common. Biome: Savanna,
Grassland, Nama-Karoo, and Desert. Throughout Africa
and in Arabia, Afghanistan, China and warm America.
Ruderal weed. May be confused with T. racemosus, which
has larger spikelets and anthers, and 7-nerved glumes.

Description: Chippindall 1955 (107), Clayton et al.
1970-1982 (400). Illustration: Chippindall 1955 (fig. 79),
Clayton et al. 1970-1982 (fig. 108). Voucher: Pott 5533.
PRECIS code 9902740-00100.

Tragus koelerioides Aschers.

Creeping carrot-seed grass,
kophaargras.

Perennial; rhizomatous and
stoloniferous; 120-650 mm tall.

Leaf blades 1 0— 50(— 80) mm long;
to 3 mm wide. Spikelets (lowest
in each cluster) 3. 5-4. 8 mm long.

Inflorescence narrowly spike-like;
spikelets clustered on peduncles that are much shorter than
the spikelets; glume hairs curved, not hooked at the tip;
anthers 1. 8-2.5 mm long.

Flowering October to May. Open veld, on a variety of
soil types. Infrequent. Biome: Savanna, Grassland, and
Nama-Karoo. Endemic. Ruderal weed (increases with
overgrazing).

Description: Chippindall 1955 (107). Illustration: Chip-
pindall 1955 (fig. 77). Voucher: Smook 3372. PRECIS code
9902740-00200.

Tragus pedunculatus Pilg.

Annual; culms branched
above, sometimes decumbent;

100^100 mm tall. Leaf blades
20-60 mm long; about 2 mm
wide. Spikelets (lowest in each
cluster) 6-10 mm long. Inflores-
cence appearing branched; spike-
lets clustered on peduncles about
as long as the spikelets; glume
hairs straight.

Flowering January to April. Shallow sand over
limestone. Conservation status not known. Biome: Savan-
na. Endemic.

Description: Chippindall 1955 (107). Voucher: Dinter
5698. PRECIS code 9902740-00300.

Tragus racemosus (L.) All.

Large carrot-seed grass,
klitsgras.

Annual; culms usually de-
cumbent; 1 1 0 — 400 mm tall. Leaf
blades 20-60 mm long; 2-\ mm
wide. Spikelets (lowest in each
cluster) 3. 5-5.0 mm long. Inflor-
escence loosely spike-like; spike-
lets clustered on peduncles that are much shorter than the
spikelets; lowest glume 7-nerved, spaces between nerves
about the same width as nerves; glume hairs hooked at tips;

anthers 0.6-0. 8 mm long.

Flowering November to May. Limestone and sandy
soils, often in moist places and disturbed areas. Common.
Biome: Savanna, Grassland, and Nama-Karoo.

Mediterranean region, Africa, SW Asia, introduced to
America. Ruderal weed. May be confused with T.
berteronianus , which has smaller spikelets and anthers, and
5-nerved glumes.

339

Description: Stapf 1898-1900 (577), Chippindall 1955
(107). Illustration: Chippindall 1955 (fig. 78). Voucher:
Smook 2774. PRECIS code 9902740-00400.

Tribolium Desv.

Brizopyrum Stapf, Lasiochloa Kunth, Plagiochloa
Adamson and Sprague.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose. Culms 20-600 mm high ; herbaceous;
branched above, or unbranched above. Plants unarmed.
Leaf blades flat, or rolled. Ligule a fringed membrane to a
fringe of hairs.

Inflorescence a single spike, or a single raceme, or pa-
niculate; contracted; espatheate. Spikelet-bearing axes
persistent.

Spikelets solitary; biseriate, or not two-ranked;
imbricate', 2-10 mm long; compressed laterally, or not
noticeably compressed; disarticulating above the glumes;
disarticulating between the florets (tardily). Callus short.
Glumes two; relatively large; very unequal, or more or less
equal; markedly shorter than the spikelets to much
exceeding the spikelets; awned (shortly), or awnless;
similar (naviculate, membranous to chartaceous). Lower
glume 5 nerved. Incomplete florets distal to the female-
fertile florets, merely underdeveloped, awnless; proximal
incomplete florets absent.

Female-fertile florets 2-9 ( occasionally to 14). Lemmas
less firm than the glumes to similar in texture to the glumes
(membranous to chartaceous); hairy (usually with clavate
hairs), or hairless; without a germination flap; 5-9 nerved;
entire', awnless to mucronate. Palea present; relatively long;
2-nerved. Lodicules 2; fleshy; ciliate, or glabrous. Stamens
3. Ovary glabrous. Fruit small (1-1.2 mm); hilum short;
pericarp fairly loosely adherent; embryo small.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Arundinoideae;
Danthonieae. 1 1 species. South Africa. Mesophytic to xero-
phytic (winter rainfall); in open habitats (Fynbos and
Karoo); glycophytic. Cape Province. 1 1 indigenous species.

References. 1. Renvoize. 1985. Kew Bull. 40: 795.

Species treatment by N.P. Barker.

1(0). Spikelets distichously arranged 2

Spikelets not distichously arranged 5

2(1). Spike clearly exserted from uppermost leaf 3

Spike partially enclosed by uppermost leaf 4

3(2). Glumes usually glabrous; spikelets up to 6 mm long;
spike up to 70 mm long; plant up to 600 mm tall

T. uniolae

Glumes pubescent; spikelets 4-5 mm long; spike up

to 25 mm long; plant up to 300 mm tall

T. brachystachyum

4(2). Glumes glabrous T. ampiexum

Glumes pubescent T. alternans

5(1). Plants annual 6

Plants perennial 8

6(5). Glumes covered in blunt, apically rounded hairs;
lemmas have two basal tufts of hairs on each

margin T. utriculosum

Glumes covered in slender, tapering hairs; lemma

margins fringed with hairs, but not tufted 7

7(6). Spikelets to 1.5 mm long; panicle up to 10 mm long

T. ciliare

Spikelets 3. 5-4.0 mm long; panicle up to 40 mm long

T. echinatum

8(5). Glumes densely pubescent, hairs up to 1.5 mm long

T. hispidum

Glumes glabrous or pubescent but not densely so and
then hairs shorter than 1 mm 9

9(8). Glumes glabrous 10

Glumes pubescent T. ohtusifolium

10(9). Hairs on lemmas club-shaped . . . T. acutiflorum
Hairs on lemmas not club-shaped . T. obliterum

Fig. 222. Tribolium uniolae

340

Tribolium acutiflorum (Nees) Renvoize

( =Plagiochloa acutiflora
(Nees) Adamson & Sprague) 1.

Perennial (sometimes weakly
so); slender, tufted; 100-300 mm
tall. Leaf blades 50(— 1 20) mm
long; to 3 mm wide. Spikelets 4-5
mm long. Panicle spike-like,

10-25 mm long, partly enclosed
in uppermost leaf; spikelets not distichous, 4— 6-flowered;
glumes glabrous, shorter than florets, acute, acuminate or
minutely awned, sometimes with a few hairs along the keel;
lemmas pubescent along keel and margins, hairs club-
shaped.

Flowering September to January. Sandy soils and dis-
turbed areas. Locally common. Biome: Fynbos and
Succulent Karoo. Endemic.

Description: Chippindall 1955 (115). Voucher: Davidse
33415A. PRECIS code 9904021-00100.

Tribolium alternans (Nees) Renvoize

( =Plagiochloa alternans
(Nees) Adamson & Sprague) 1.

Perennial; tufted; to 700 mm
tall. Leaf blades to 300 mm long;
to 5 mm wide. Spikelets 6— 7(— 1 0)
mm long; 4 mm wide. Inflores-
cence a loose spike, 40-60 mm
long, partly enclosed by upper-
most leaf; spikelets 4-8-flowered, barely overlapping each
other, distichous on a triangular rachis with a small, pointed
lobe or appendage opposite some or all spikelets; glumes
sparsely pubescent, hairs glassy, tuberculate; lower half of
lemma pubescent, hairs club-shaped.

Flowering October to December. River flats in sandy
soil. Infrequent. Biome: Fynbos. Endemic.

Description: Chippindall 1955 (115). Voucher:

Hanekom 2638. PRECIS code 9904021-00200.

Tribolium amplexum Renvoize

Perennial; tufted; 200-700
mm tall. Leaf blades 100-200
mm long; to 4 mm wide. Spike-
lets 4-7 mm long; 1.5-3. 5 mm
wide. Panicle spicate, 25-50 mm
long, 5-7 mm wide, partly
enclosed by uppermost leaf;
spikelets 4— 6-flowered, distich-
ous on a triangular, scabrid rachis; glumes glabrous; lower
half of lemma pubescent, hairs long, club-shaped.

Flowering September to December. Sandy soils in
disturbed areas. Infrequent. Biome; Fynbos. Endemic.

Description: Renvoize 1985 (797), Chippindall 1955
(115). Voucher: Oliver 4682. PRECIS code 9904021-
00300.

Tribolium brachystachyum (Nees) Renvoize

(-Plagiochloa brachystachya
(Nees) Adamson & Sprague) 1.

Perennial; prostrate to tufted;
to 300 mm tall. Leaf blades
10-120 mm long (rarely longer);
about 2.5 mm wide. Spikelets 4—5
mm long; to 3.5 mm wide. Inflor-
escence a compacted spike, to 25
mm long, exserted from the uppermost leaf; spikelets
(4-)5-6-flowered, distichous, overlapping for 3-4 or more
of their length; glumes pubescent, hairs glassy; lower half
of lemmas pubescent, hairs club-shaped; keels of paleas

may be slightly winged.

Flowering October to January. Sandy soils in mountains
and in disturbed areas. Locally common. Biome: Fynbos.
Endemic. May be confused with T. alternans , which has a
larger, less compacted spike and larger spikelets.

Description: Stapf 1898-1900 (707), Chippindall 1955
(115). Voucher: Ellis 2855. PRECIS code 9904021-00400.

Tribolium ciliare (Stapf) Renvoize

( =Plagiochloa ciliaris (Stapf)

Adamson & Sprague) 1.

Annual; weakly tufted; to 100
mm tall. Leaf blades to 20 mm
long; to 1 mm wide. Spikelets to
1.5 mm long; to 1 mm wide. Pani-
cle to 10 mm long, partly en-
closed in the uppermost leaf
sheath; spikelets not distichous; glumes pubescent, hairs
glassy, tubercle-based, tapering apically; lemma backs
glabrous but lower margins fringed with acicular hairs and
upper margins fringed with a few tubercle-based hairs.

Flowering September to October. On limestone
outcrops. Locally common (in the Bredasdorp district).
Biome: Fynbos. Endemic.

Description: Stapf 1899 Hook. Icon. PI. Plate. 27 t2602.
Voucher: Davidse 33525. PRECIS code 9904021-00500.

Tribolium echinatum (Thunb.) Renvoize

( -Lasiochloa echinata
(Thunb.) Adamson) 1.

Annual; prostrate to tufted; to
250 mm tall. Leaf blades to 150
mm long; to 5 mm wide. Spike-
lets 3. 5-4.0 mm long; 1.0-1. 5
mm wide. Leaves pubescent,
sometimes densely so; panicle to
40 mm long, partly enclosed in uppermost leaf; spikelets
(2-)3(-4)-flowered, not distichous; glumes long-acuminate,
pubescent, hairs long, glassy, tuberculate, tapering apically;
lemmas glabrous except for a fringe of hairs on the lower
margins.

Flowering August to November. Sandy soils and road-
sides. Locally common. Biome: Fynbos and Succulent
Karoo. Endemic.

Description: Chippindall 1955 (116). Voucher: Davidse
33250. PRECIS code 9904021-00600.

Tribolium hispidum (Thunb.) Renvoize

(-Lasiochloa longifolia
(Schrad.) Kunth) 1.

Perennial; tufted; to 400 mm
tall. Leaf blades to 250 mm long;
to 4 mm wide. Spikelets 3-4 mm
long; to 1.5 mm wide. Panicle
1 0— 50(— 70) mm long; spikelets
(2-)3(-4)-flowered, not distich-
ous; glumes densely pubescent, hairs 1.0-1. 5 mm long,
tubercle-based; lower margin of lemma fringed with hairs.

Flowering August to February. Sandy soils. Locally
common (Cedarburg-Clanwilliam area and in Renosterbos-
veld). Biome: Fynbos, Grassland, and Succulent Karoo.
Endemic.

Description: Chippindall 1955 (1 16). Illustration: Chip-
pindall 1955 (fig. 85). Voucher: Smook 3633. PRECIS code
9904021-00700.

341

Tribolium obliterum (Hemsl.) Renvoize

(-Plagiochloa oblitera
(Hemsl.) Adamson &

Sprague) 1.

Perennial; stoloniferous, pro-
strate or tufted; 100-350 mm tall.

Leaf blades to 150 mm long; to 2
mm wide. Spikelets 4-5 mm
long; to 2 mm wide. Panicle
15-25 mm long, often partly enclosed in the uppermost
sheath; spikelets 3^t(-6)-flowered, not distichous; glumes
glabrous, as long as florets; lemma keels and/or margins
fringed with hairs which are not club-shaped.

Flowering September to December. Disturbed areas
such as cultivated fields and roadsides. Locally common.
Biome: Fynbos. Endemic.

Description: Chippindall 1955 (115). Voucher: Davidse
33840. PRECIS code 9904021-00800.

Tribolium obtusifolium (Nees) Renvoize

(= Lasiochloa obtusifolia
Nees) 1.

Perennial; stoloniferous; to
250 mm tall. Leaf blades to 200
mm long; to 1 mm wide. Spike-
lets 3-4 mm long; 1. 5-2.0 mm
wide. Panicle 20-30 mm long;
spikelets 2-3-flowered, not dis-
tichous; glumes pubescent, but not densely so, hairs taper-
ing and glassy, especially along the keel; lemma margins
fringed with hairs which are not club-shaped.

Flowering September to November. Sandy areas. Infre-
quent. Biome: Fynbos. Endemic.

Description: Chippindall 1955 (116). Voucher: Duthie
1761a. PRECIS code 9904021-00900.

Tribolium uniolae (L.f.) Renvoize

Fig. 222. PI. 203.

( =Plagiochloa uniolae (L. f.)

Adamson & Sprague var.
uniolae 1; ( =Plagiochloa
uniolae (L. f.) Adamson &

Sprague var. villosa (Stapf)

Adamson) 1.

Perennial; tufted; 100-600
mm tall. Leaf blades to 200 mm
long; to 3 mm wide. Spikelets to 6 mm long; to 4 mm wide.
Inflorescence a spike, 8-70 mm long, often branched at the
base, exserted from uppermost leaf; spikelets 5-9-flowered,
distichous; glumes usually glabrous; lower half of lemma
backs pubescent, hairs club-shaped, margins fringed with
stout hairs.

Flowering September to December. Disturbed areas
such as roadsides and fields. Locally common. Biome: Fyn-
bos. Endemic.

Description: Stapf 1898-1900 (705). Illustration: Chip-
pindall 1955 (fig. 84 (inflorescence only)). Voucher:
Davidse 34149. PRECIS code 9904021-01000.

Tribolium utriculosum (Nees) Renvoize

(=Lasiochloa utriculosa
Nees) 1 .

Annual; tufted; to 90 mm tall.

Leaf blades 10-100 mm long; to
2.5 mm wide. Spikelets 2-3 mm
long; to 2 mm wide. Leaf blades
sparsely pubescent; panicle
10-20 mm long, usually partly
enclosed in uppermost leaf; spikelets not distichous; glumes

pubescent, hairs short, glassy, tuberculate, apically blunt
and rounded; lemma margins with two tufts of club-shaped
hairs.

Flowering August to October. Sandy alluvial soils and
disturbed areas. Locally common. Biome: Succulent
Karoo. Endemic.

Description: Chippindall 1955 (116). Illustration: Chip-
pindall 1955 (fig. 86). Voucher: Thompson & Le Roux 99.
PRECIS code 9904021-01 100.

Tricholaena Schrad.

Annual (rarely), or perennial; caespitose, or decumbent.
Culms 100-1200 mm high; herbaceous; unbranched above.
Leaf blades often glaucous-inrolled, rigid. Ligule a fringed
membrane (very narrow ), or a fringe of hairs. Plants with
hermaphrodite florets.

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets 2-3.5 mm long; compressed laterally. Glumes
present; one per spikelet, or two; very unequal; awnless (the
upper sometimes mucronate); very dissimilar (the lower
often reduced to a tiny scale, hairy or glabrous), or similar
(rarely). Lower glume 0-1 nerved. Proximal incomplete
florets 7; paleate, palea fully developed; florets male.
Proximal lemmas awnless.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes (sub-crustaceous); smooth; hairless (shiny);
having the margins lying flat and exposed on the palea; 3-5
nerved; entire to incised; awnless. Palea present; relatively
long. Lodicules 2; glabrous. Stamens 3. Ovary glabrous.
Hilum short; embryo large.

Photosynthetic pathway. C4; XyMS+. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 9. Panicoideae; Panicodae; Paniceae
(Melinideae). 12 species. Africa, Madagascar, Canaries,
Mediterranean. Xerophytic; in open habitats (sandy and
stony soil, sometimes ruderal); glycophytic. Namibia,
Botswana, Transvaal, Orange Free State, Swaziland, Natal,
and Cape Province. 2 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Launert.
1970. FSWA. 3. Anderson. 1961 . Kirkia 1 : 103. 4. Clayton
& Renvoize. 1982. FTEA. 5. Zizka. 1988. Bibliotheca
Botanica, 138.

Species treatment by H.M. Anderson.

1(0). Culms rarely branching from lower or middle nodes;
leaves sometimes hairy; spikelets glabrous or very

rarely hairy T. monachne

Culms often branching from lower and middle nodes;

leaves and spikelets hairy 2

2(1). Lower glume 1-3 mm long; spikelets with hairs 1-3

mm long T. capensis subsp. capensis

Lower glume a scale 0. 1 — 0.4(— 2.0) mm long; spikelets

with hairs 0.5-2. 0 mm long

T. capensis subsp. arenaria

Tricholaena capensis (Licht. ex Roem. & Schult.) Nees
subsp. arenaria (Nees) Zizka

(: -T . arenaria Nees) 2; (-T.
arenaria Nees var. glauca
(Hack.) Stapf) 2.

Perennial; tufted; 200-600
mm tall. Leaf blades 30-70 mm
long; 2.0-3. 5 mm wide. Spikelets
2-3 mm long; 1 mpn wide. Culms
branching from lower nodes;

342

culm nodes, leaves andspikelets hairy; lower glume nearly
always a scale 0. 1— 0.4(— 2.0) mm long; glumes and lemmas
covered sparingly with short hairs 0.2-2. 0 mm long.

Flowering January to April. Sandy, dry areas. Locally
common. Biome: Nama-Karoo. Endemic. This subspecies
is restricted to Namibia and is sympatric near the Orange
River with subsp. capensis, which has more hairy spikelets

and a larger lower glume.

Description: Zizka 1988 (49). Voucher: Giess 10250.
PRECIS code 9901330-00050.

Tricholaena capensis (Licht. ex Roem. & Schult.) Nees
subsp. capensis

Fig. 223. Tricholaena monachne

Perennial; tufted; 200-600
mm tall. Leaf blades 30-70 mm
long; 2. 0-3. 5 mm wide. Spikelets
2-3 mm long; 1 mm wide. Culms
branching from lower nodes;
culms, leaves and spikelets hairy;
lower glume 1-3 mm long;
glumes and lemmas covered with
dense hairs 1-3 mm long.

Flowering January to June. Sandy, dry soil. Locally
common. Biome: Nama-Karoo and Succulent Karoo.
Endemic. See note under T. capensis subsp. arenaria.

Description: Zizka 1988 (48). Chippindall 1955 (435).
Illustration: Chippindall 1955 (fig. 361). Voucher: Davidse
6209. PRECIS code 9901330-00100.

Tricholaena monachne (Trin.) Stapf & C.E. Hubb.

Fig. 223. PI. 204.

Blousaadgras.

Perennial, or annual; 200-
1000 mm tall. Leaf blades 30-70
mm long; 2.0-3. 5 mm wide.

Spikelets 2-3 mm long; 1 mm
wide. Culm glabrous; leaves
sometimes hairy; spikelets usual-
ly glabrous or very rarely hairy.

Flowering November to March. Favours sandy soil, also
occurs as a ruderal. Common. Biome: Savanna and Grass-
land. Tropical Africa. Natural pasture. An annual form oc-
curs in Namibia that is less than 300 mm tall with a softer
appearance than the hardy, wiry, drought resistant perennial
form. The spikelets in this species are usually glabrous, but
individuals with hairy spikelets can be separated by their
glabrous culms from T. capensis , which has hairy culms.
T. monachne is distinguished from Panicum species which
have distinct lower glumes never reduced to a scale, and
Eriochloa meyeriana which has a scale-like lower glume,
but it is broadly ovate and clasps the base of the spikelet.

Description: Chippindall 1955 (434), Zizka 1988 (38).
Illustration: Chippindall 1955 (fig. 360). Voucher: Smook
4234. PRECIS code 9901330-00300.

Trichoneura Anderss.

Crossotropis Stapf.

Annual, or perennial; xeromorphic. Culms 120-1000
mm high; herbaceous; branched above (often), or unbranch-
ed above. Leaf blades linear (pointed); usually flat. Ligule
an unfringed membrane .

Inflorescence of spike-like main branches ( the racemes
scattered along a central axis)-, open; espatheate. Spikelet-
bearing axes persistent.

Spikelets solitary; 5.3-14 mm long; compressed
laterally; disarticulating above the glumes; disarticulating
between the florets. Glumes two; more or less equal; about
equalling the spikelets to much exceeding the spikelets ;
awned, or awnless (tapered into a mucro or short awn);
similar (narrowly lanceolate, membranous, persistent). In-
complete florets distal to the female-fertile florets; proximal
incomplete florets absent.

Female-fertile florets 2-8. Lemmas similar in texture to
the glumes (membranous); non-carinate ( rounded on the
back)-, 3 nerved; incised; mucronate, or awned. Awns when
present 1; from the sinus; non-geniculate; much shorter than

343

Fig. 224. Trichoneura grandiglumis

the body of the lemma. Palea present; relatively long. Lodi-
cules 2; fleshy; glabrous. Stamens 3. Ovary glabrous. Fruit
small; hilum short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
XyMS+. PCR cell chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 1
species. America, tropical Africa. Xerophytic; in open
habitats (in sandy or stony soil). Namibia, Botswana,
Transvaal, Orange Free State, Swaziland, Natal, Lesotho,
and Cape Province. 3 indigenous species.

References. 1. Clayton et al. 1974. FTEA.

Species treatment by M. Koekemoer.

1(0). Plants perennial; inflorescences broadly pyramidal,
70-320 mm long; primary branches stiff and
straight, usually longer than 1/2 the central axis,
often spreading horizontally; glumes as long as, or
to twice as long as the spikelets . T. grandiglumis
Plants annual; inflorescences broadly lanceolate,
40-190 mm long; primary branches firm, often
slightly curved, shorter than 1/2 the central axis,
somewhat contracted but never spreading
horizontally; glumes as long as the spikelets ... 2
2(1). Culms decumbent or ascending, 90-350 mm tall;
panicle usually 40-90(-130) mm long; from

Namibia T. eleusinoides

Culms erect, 340-660 mm tall, panicle 120-190 mm
long; from the Zoutpansberg district of Transvaal
T. sp. (=Codd 5325)

Trichoneura eleusinoides (Rendle) Ekman

Annual; tufted; 90-350 mm
tall. Leaf blades 20^)5 mm long;

2-3 mm wide. Spikelets 3-4 mm
long. Inflorescence up to 90 mm
long, 50 mm wide, contracted
with side branches stiff, shorter
than 30 mm and not spreading
more than 45 degrees.

Flowering January to May.

Rocky outcrops and granite mountain slopes. Infrequent to
locally common. Biome: Savanna, Nama-Karoo, and Des-
ert. To east Africa.

Description: Chippindall 1955 (129). Voucher: Dinter
7053. PRECIS code 9903530-00100.

Trichoneura grandiglumis (Nees) Ekman

Rolling grass, waaigras.

Perennial; tufted (culms slen-
der; erect or ascending); 220-630
mm tall. Leaf blades 30-200 mm
long; 3-7 mm wide. Spikelets
5-14 mm long. Inflorescence
branches spreading horizontally;
spikelets 4-9-flowered; glumes
5.0-13.5 mm long.

Flowering November to April. On sandy soils on
hillsides or open floodplains and in bushveld, sometimes in
disturbed areas. Common. Biome: Savanna and Grassland.
Africa south of the Congo Basin. The length of the glumes
and the spikelets vary considerably. Previously two
varieties, var. grandiglumis and var. minor , were
distinguished on spikelet length (5.3-14.0 mm and 5- 3—8.2
mm respectively), and the ratio of the awns (longer than and
shorter than the floret respectively). No clear separation of
specimens are given by these two characters and pending
further investigation, these varieties are not upheld.

Description: Chippindall 1955 (128), Clayton et al.
1970-1982 (299). Voucher: Huntley 967, Strey R.S.B. 53.
PRECIS code 9903530-00200.

Fig. 224. PI. 205.

344

Trichoneura sp. (=Codd 5325)

Annual; tufted (slender with
leaves mostly cauline); 340-660
mm tall. Leaf blades 50-150 mm
long; 2^1 mm wide. Spikelets
5-1 1 mm long. Panicle up to 190
mm long with branches not
spreading more than 45 degrees;
spikelets more or less their own
length apart.

Flowering January to April. Sandy soil in rocky areas.
Infrequent to locally common. Biome: Savanna. This
species resembles T. eleusinoides but Chippindall (1955)
regards it as an undescribed annual species. Specimens of
this species have mistakenly been identified as T.
schlechteri , a perennial known from Lourenco Marques and
not recorded for the FSA area.

Description: Chippindall 1955 (129). Voucher: Codd
5325. PRECIS code 9903530-99999.

Trichopteryx Nees

Annual, or perennial (with slender culms); caespitose,
or decumbent. Culms 20-900 mm high; herbaceous;
branched above, or unbranched above. Leaf blades linear-
lanceolate to lanceolate. Ligule a fringe of hairs. Plants
with hermaphrodite florets .

Inflorescence paniculate ; open, or contracted; espathe-
ate. Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs', consistently in Tong-and-
short’ combinations, or not in distinct ‘long-and-short
combinations. Spikelets 2.5-6 mm long; compressed
laterally to not noticeably compressed; disarticulating
above the glumes (disarticulating readily between LI and
L2, less readily between G2 and LI). Hairy callus present.
Glumes two; relatively large; very unequal (G 1 one third
to one half spikelet length); awnless (though the G1 can be
aristulate and the G2 acuminate); similar (membranous or
papery, G1 narrower). Proximal incomplete florets 1\
paleate, palea fully developed (two keeled, thin); male, or
sterile.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (membranous,
hardening to leathery); hairy (with a sub-marginal tuft of
erect hairs, in the middle on each side); the margins tucked
in onto the palea (the palea embraced and almost enclosed);
without a germination flap; 5-7 nerved; incised; awned.
Awns 1 (from the sinus), or 3; median, or median and lateral
(with the lobes terminating in awns, in addition to the
median awn). The median awn different in form from the
laterals (when laterals present); from the sinus; geniculate
(the lateral awns, when present, straight); much longer than
the body of the lemma. Palea present; relatively long. Lodi-
cules 2; fleshy. Stamens 2. Ovary glabrous. Hilum long-
linear; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional, or unconventional. Organization
of PCR tissue when unconventional Arundinella type.
XyMS-. PCR cell chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, jc = 12 (?). Panicoideae; Panicodae; Arundinelleae.
5 species. Southern and tropical Africa, Madagascar. Helo-
phytic, or mesophytic; in shade, or in open habitats
(streambanks, grasslands and forest margins); glycophytic.
Namibia, Transvaal, Swaziland and Cape Province
(Transkei). 1 indigenous species.

References. 1. Clayton et al. 1972. FTEA.

Species treatment by H.M. Anderson.

Trichopteryx dregeana Nees

Fig. 225. PI. 206.

Vleigras.

Perennial; tufted; to 900 mm
tall. Leaf blades to 500 mm long
(light green); 4 mm wide. Spike-
lets 4-7 mm long. Grows in a
tangled mass, leaves spreading
and reflexed; panicle open, up to
140 mm long; spikelets with a
whorl of hairs at the base; glumes and lower lemma bright
brown and tips transparent; female-fertile (upper) lemma
with two conspicuous tufts of white hairs, side awns 2—3
mm long, central awns slender, 4-7 mm long.

Flowering December to May. Vleis and wet places,
shady crevices among rocks on hillsides. Locally common.
Biome: Savanna and Grassland. Southern tropical Africa.
Habit similar to Eragrostis volkensii, which can be dis-
tinguished vegetatively by the blades being blue-green,
wider (up to 8 mm) and stiffer.

Description: Chippindall 1955 (287). Illustration. Chip-
pindall 1955 (fig. 257). Voucher: Kluge 1694. PRECIS
code 9901750-00100.

345

Tripogon Roem. & Schult.

Archangelina Kuntze, Kralikia Coss. & Dur.,
Kralikiella Batt. & Trab., Plagiolytrum Nees.

Annual, or perennial; caespitose. Culms 40-650 mm
high; herbaceous; unbranched above. Leaves without
auricles. Leaf blades linear (often filiform). Ligule an
unfringed membrane to a fringe of hairs.

Inflorescence a single spike ( slender ); espatheate.
Spikelet-bearing axes persistent.

Spikelets solitary; alternately distichous', 3-25 mm long;
compressed laterally; disarticulating above the glumes; dis-
articulating between the florets. Glumes two; very unequal,
or more or less equal; markedly shorter than the spikelets;
dorsiventral to the rachis\ awnless; very dissimilar, or
similar (membranous, narrow, G1 often asymmetric). In-
complete florets distal to the female-fertile florets, merely
underdeveloped; proximal incomplete florets absent
(rarely, LI also neuter).

Female-fertile florets 3-20. Lemmas 1-3 nerved;
mucronate, or awned. Awns when present 1, or 3, or 5
(usually awned or mucronate from a median sinus or behind
the apex, the lobes sometimes awned or mucronate);
median, or median and lateral (via mucronate to awned
lobes). The median awn similar in form to the laterals
(when laterals present); from the sinus, or apical; non-gen-
iculate; much shorter than the body of the lemma, to much
longer than the body of the lemma. Palea present. Lodicules

Fig. 226. Tripogon minimus

2; fleshy; glabrous. Stamens 2, or 3. Ovary glabrous. Fruit
small (0.8-2. 2 mm); hilum short; pericarp fused; embryo
large, or small ( 1/3 the length of the fruit or somewhat less).

Photosynthetic pathway and related features. C4;
XyMS+. PCR sheath outlines even. PCR sheath extensions
present. Maximum number of extension cells 1. PCR cell
chloroplasts centripetal.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato.
About 30 species. Tropical Africa, Asia, Australia. Helo-
phytic to xerophytic; in open habitats; glycophytic.
Namibia, Botswana, Transvaal and Natal. 1 indigenous
species.

References. 1. Clayton. 1970. FTEA. 2. Clayton et al.
1974. FTEA.

Species treatment by M. Koekemoer.

Tripogon minimus (A. Rich.) Steud.

Fig. 226. PI. 207.

( -T . abyssinicus sensu
Chippind., non Nees) 1.

Perennial; tufted; 80-220 mm
tall. Leaf blades 10-90 mm long,
filiform; to 0.5 mm wide. Spike-
lets 2. 6-8.0 mm long. Old leaf
sheaths divide into coarse fibres;
spikes 20-80 mm long, slender,
erect; spikelets 5-10-flowered; lemma tip emarginate and
mucronate.

Flowering December to May. Mostly in shallow soil on
rocky outcrops but also in waterlogged sand and seasonal
pans. Infrequent. Biome: Savanna. Tropical east Africa and
Madagascar.

Description: Phillips & Launert 1971 Kew Bull. 25,2
(301), Clayton et al. 1970-1982 (289). Illustration: Kew
Bull. 25,2 (fig. 1). Voucher: Killick & Leistner 3371.
PRECIS code 9903180-00100.

Triraphis R.Br.

Annual, or perennial; caespitose (mostly small
xeromorphs). Culms (10-)40-1400 mm high; herbaceous.
Leaf blades flat, or rolled (or junciform). Ligule a fringe
of hairs.

Inflorescence paniculate ; open, or contracted (rarely
spiciform); espatheate. Spikelet-bearing axes persistent.

Spikelets compressed laterally; disarticulating above the
glumes; disarticulating between the florets. Glumes two;
relatively large; very unequal (rarely), or more or less
equal; markedly shorter than the spikelets ; awned (or
mucronate, from the sinus), or awnless; similar (narrow,
persistent). Incomplete florets distal to the female-fertile
florets, merely underdeveloped; proximal incomplete
florets absent.

Female-fertile florets 5-10. Lemmas 3 nerved; incised;
deeply cleft ; awned. Awns 3, or 5; median and lateral (the
lateral lobes setiform-awned or mucronate). The median
awn similar in form to the laterals (setiform); from the sinus
(of the central lobe); non-geniculate. Palea present; shorter
than the lemma. Lodicules 2; fleshy, or membranous;
glabrous. Stamens 3. Ovary glabrous. Fruit small; linear;
hilum short; pericarp fused; embryo large.

Photosynthetic pathway and related features. C4;
NAD-ME (mollis)-, XyMS+. PCR sheath outlines uneven.
PCR sheath extensions present. Maximum number of
extension cells 3. PCR cell chloroplasts ovoid; with well
developed grana; centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10. Chloridoideae; Chlorideae sensu lato. 7
species. Tropical and southern Africa, Australia. Meso-
phytic to xerophytic; in open habitats (savanna, in sandy or

346

rocky soil); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Natal, and Cape Province. 5 indigenous
species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton.
1970. FTEA. 3. Launert. 1970. FSWA.

Species treatment by M. Koekemoer.

1(0). Plants annual; leaves mostly basal 2

Plants perennial; leaves mostly cauline 3

2(1). Plants shorter than 250 mm; panicle dense, widely
elliptic, 10-30 mm long; anthers shorter than 0.5

mm T. pumilio

Plants to 770 mm tall; panicle open to dense, widely
ovate, longer than 30 mm; anthers 1 .2-2.0 mm long

T. purpurea

3(1). Culms profusely branched, yellowish; plants tufted;
spikelets 4-15 mm long; central awn of the lemmas

longer than the lemmas T. ramosissima

Culms unbranched, dark brown to reddish; plants
rhizomatous or tufted; spikelets to 10 mm long;
central awn of the lemmas shorter or longer than

the lemmas 4

4(3). Central awn of lemmas shorter than the lemmas;
rhizomes long and very well developed; panicles
dense, 120-300 mm long . T. andropogonoides
Central awn of lemmas longer than the lemmas;
rhizomes short; panicles sparse, to 400 mm long
T. schinzii

Triraphis andropogonoides (Steud.) Phill.

Besemgras.

Perennial; rhizomatous (long
creeping rhizome); 380-1220mm
tall. Leaf blades 200^100 mm
long; 2-6 mm wide. Spikelets
6—1 0(— 1 5) mm long. Plant base
dark brown to reddish; rootstock
very well developed; tillers very
loosely grouped; panicle dense, 120-300 mm long;
spikelets 5-15-flowered, central awn shorter than the
lemma; anthers 1.2-2. 3 mm long.

Flowering October to May. Well-drained soil on rocky
slopes or in deep sand in open grassland. Common. Biome:
Fynbos, Savanna, Grassland, and Nama-Karoo. Endemic.
Similar to T. schinzii , which has a central lemma awn longer
than the lemma and short rhizomes. Some interesting
specimens were collected in the Bathurst and Alexandria
districts. They resemble T. andropogonoides in all aspects,
except that the culms are fasciculately branched at the
nodes.

Description: Chippindall 1955 (125). Illustration: Chip-
pindall 1955 (fig. 98). Voucher: Van der Schijff 5321.
PRECIS code 9903500-00100.

Triraphis pumilio R. Br.

Annual; tufted; 40-220 mm
tall. Leaf blades 50-120 mm
long; 2 mm wide. Spikelets 2-\
mm long. Panicle dense, ovoid,

5-30 mm long; spikelets 3—1 1 -
flowered; lemma 3-nerved;
central awn about as long as the
lemma; anthers 0. 2-0.4 mm long.

Flowering January to May. In
riverbeds or moist depressions in sand. Locally common.
Biome: Desert. Northern Africa through Mauritania to
Arabia.

Description: Launert 1970 (160:211), Stapf 1898-1900
(653), Chippindall 1955 (127). Illustration: Chippindall

Fig. 227. PI. 208.

1955 (fig. 99). Voucher: Oliver & Muller 6661. PRECIS
code 9903500-00400.

Fig. 227. Triraphis andropogonoides

Triraphis purpurea Hack.

(-T.fleckii Hack.) 2.

Red honey grass.

Annual; tufted; 90-770 mm
tall. Leaf blades 35-60 mm long;

1-2 mm wide. Spikelets 6-10 mm
long. Panicle longer than 30 mm,
open or dense; spikelets 5-11
(-24)-flowered; anthers 1. 2-2.0 mm long.

Flowering January to June. Often in moist patches in the
shade on red sand or rocky calcareous soils. Common.
Biome: Savanna, Nama-Karoo, and Desert. Endemic. At the
moment this taxon contains all annual specimens that do not
match T. pumilio. Launert (1970) recognizes three groups
within this species, based on the types of T. purpurea , T.
fleckii Hack, and T. welwitschii Rendle, but is reluctant to
assign any taxonomic ranks until a proper revision can be
done.

Description: Launert 1970 (160:212), Muller 1984
(262), Stapf 1898-1900 (653), Chippindall 1955 (127).
Illustration: Muller 1984 (fig. 132). Voucher: Van Vuuren
& Giess 1095. PRECIS code 9903500-00500.

347

Triraphis ramosissima Hack.

( -T . elliottii Rendle) 2.

Berggras.

Bushy perennial; rhizomatous
and tufted; 250-8 10 mm tall. Leaf
blades 60-120 mm long; 1 mm
wide. Spikelets 4-15 mm long.

Culms yellowish and woody,
branching profusely; spikelets 4-19-flowered, central awn
longer than the lemma.

Flowering February to June. Rocky hillslopes, on
floodplains, in dry watercourses, often in sand or calcareous
soil. Common. Biome: Savanna, Nama-Karoo, and Succu-
lent Karoo. Endemic. Distinguished from other Triraphis
species by the profusely branched culms.

Description: Muller 1984 (264), Stapf 1898-1900 (651),
Chippindall 1955 (125). IllustratiomMuller 1984 (fig. 133),
Chippindall 1955 (fig. 97). Voucher: De Winter 2618.
PRECIS code 9903500-00600.

lobes, or not in tufts); the margins tucked in onto the palea
(palea enclosed, save at its summit); with a clear germin-
ation flap; 5-7 nerved; incised; awned. Awns 1; median;
from the sinus (from between the lobes); geniculate; much
longer than the body of the lemma. Palea present. Lodicules
2; fleshy (narrowly cuneate). Stamens 3 (usually?). Ovary
hairy. Hilum long-linear; embryo large.

Photosynthetic pathway. C4; XyMS-. PCR cell
chloroplasts centrifugal/peripheral.

Cytology, classification, distribution. Chromosome base
number, x = 10 and 12. Panicoideae; Panicodae; Arun-
dinelleae. About 20 species. Tropical and southern Africa,
Madagascar, tropical America. Helophytic to xerophytic; in
shade and in open habitats (grassland and savanna,
woodland and floodplains, wet to dry soils); glycophytic.
Namibia, Botswana, Transvaal, Orange Free State,
Swaziland, Natal, Lesotho, and Cape Province. 6 indige-
nous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
et al. 1972. FTEA.

Species treatment by H.M. Anderson.

Triraphis schinzii Hack.

(=T. schlechteri Pilg. ex
Stent) 2.

Perennial; short-rhizomatous
and tufted; 700-1400 mm tall.

Leaf blades 250-500 mm long;

2-5 mm wide. Spikelets 6-1 1 mm
long. Plant base dark brown to
reddish; panicle open, 200^100
mm long; central awn longer than the lemma.

Flowering November to April. Sandy grassland or
bushveld, deep sand on dunes or riverbanks and on forest
margins. Common. Biome: Savanna and Grassland.
Tanganyika. Closely related to T. andropogonoides , which
has a very well developed rhizome and the central awn of
the lemma shorter than the lemma.

Description: Muller 1984 (266), Chippindall 1955 (125),
Clayton et al. 1970-1982 (128). Illustration: Muller 1984
(fig. 134). Voucher: Story 6398. PRECIS code 9903500-
00700.

Tristachya Nees

Apochaete (C. E. Hubbard) Phipps, Dolichochaete
Phipps, Loudetia A. Bv.,Monopogon Presl, Muantijamvella
Phipps, Veseyochloa Phipps.

Annual (rarely), or perennial; caespitose. Culms
150-2700 mm high; herbaceous. Leaf blades flat, or rolled
(then involute or convolute, often rigid). Ligule a fringe of
hairs. Plants with hermaphrodite florets.

Inflorescence a single raceme, or paniculate ; open;
espatheate. Spikelet-bearing axes persistent.

Spikelets in triplets (the triads terminating the panicle
branches)-, 10-45 mm long-, compressed laterally to not
noticeably compressed (?); disarticulating above the
glumes. Glumes two; more or less equal; awnless (obtuse,
or lanceolate to acuminate, or rostrate); similar ; the lower
glume exceeding the female-fertile lemma. Proximal in-
complete florets 1 ; paleate, palea fully developed (narrow,
two keeled); male.

Female-fertile florets 1 . Lemmas similar in texture to the
glumes to decidedly firmer than the glumes (leathery to
cartilaginous); not becoming indurated; usually hairy, or
hairless (hairs in tufts, rarely with tufts at the bases of the

Fig. 228. Tristachya leucothrix

348

1(0). Pedicels free 2

Pedicels connate 3

2(1). Awns 40-120 mm long; culms markedly bulbous at

base T. superba

Awns 15-35 mm long; culms partly swollen at base

T. lualabaensis

3(1). Culms robust, 2-3-noded, 600-2000 mm long ....

T. nodiglumis

Culms slender, 1— 2-noded, 150—900 mm long ... 4
4(3). Upper lemma side-awns 3-5 mm long; tubercle-based

hairs common on glumes and lower lemma

T. leucothrix

Upper lemma side-awns longer than 10 mm; tubercle-

based hairs on glumes only 5

5(4). Upper lemma side-awns 10-14 mm long; tubercle-

based hairs along margins of glumes only

T. biseriata

Upper lemma side-awns 18-24 mm long; tubercle-

based hairs usually absent on glumes

T. rehmannii

Tristachya biseriata Stapf

Perennial; tufted; 300 - 900
mm tall. Leaf blades 300-400
mm long; to 2 mm wide. Spike-
lets 20-25 mm long. Leaf blades
filiform; pedicels connate;
glumes with tubercle-based hairs
along the margins; female-fertile
(upper) lemma side-awns (10-) 12
(-14) mm long, central awns
30-50 mm long.

Flowering October to March. Shallow stony soils on
hillsides and rocky outcrops. Locally common. Biome;
Grassland.

Description: Chippindall 1955 (277). Illustration: Chip-
pindall 1955 (fig. 248). Voucher: Smook 4853. PRECIS
code 9901740-00100.

Tristachya leucothrix Nees

Fig. 228. PI. 209.

( =Apochaete hispida (L. f.)

J.B. Phipps) 2; ( -T . hispida (L.
f.) K. Schum.) 2.

Rooisaadgras, trident grass.

Perennial; tufted; 150 -900
mm tall. Leaf blades 50^100 mm
long; 2-6 mm wide. Spikelets
24—45 mm long. Basal leaf sheaths covered with dense
brown hairs at the base; pedicels connate; glumes and lower
lemma with many tubercle-based hairs; female-fertile
(upper) lemma side-awns 3-5 mm long, central awns
50-100 mm long.

Flowering October to March. Marshy grassland,
mountain sourveld and on hillsides. Locally dominant
(highland sourveld). Biome; Fynbos, Savanna and
Grassland. Tropical Africa. Natural pasture (for sheep).

Description: Chippindall 1955 (276). Illustration: Chip-
pindall 1955 (fig. 249). Voucher: Smook 1699. PRECIS
code 9901740-00450.

Tristachya lualabaensis (De Wild.) J.B. Phipps

(=T. hitchcockii (C.E. Hubb.)

Conert) 2.

Perennial; tufted; 700-1400
mm tall. Leaf blades 60-300 mm
long; 2-6 mm wide. Spikelets
10-20 mm long. Culms partly
swollen at the base but not bulb-
ous; spikelets in triads, rarely in
pairs, pedicels 5 mm and 10 mm long respectively; female-

fertile (upper) lemma side-awns2-5 mm long, central awns
15-35 mm long.

Flowering January to March. Alluvial soils subject to
flooding. Locally common (river floodplains). Biome: Sa-
vanna. Tropical Africa. Allied to T. superba , which has a
bulbous base and much larger spikelets.

Description: Clayton et al. 1970-1982 (423). Voucher:
Curson 669. PRECIS code 9901740-00540.

Tristachya nodiglumis K. Schum.

( =T . eylesii Stent &

Rattray) 2.

Robust perennial; tufted;

600-2000 mm tall. Leaf blades
150-600 mm long; 3-13 mm
wide. Spikelets 18-30 mm long.

Panicle with 8-70 triads; pedicels
connate; lower glume glabrous or
with tubercle-based hairs; female-fertile (upper) lemma
side-awns 10-20 mm long, central awns 30-60 mm long.

Flowering December to March. Floodplain grassland on
sandy soil. Infrequent. Biome: Savanna. Tropical Africa. A
variable species, which intergrades with T. rehmannii,
which has tubercled hairs, and T. longispiculata, which has
longer spikelets.

Description: Clayton et al. 1970-1982 (426).

Illustration: Clayton et al. 1970-1982 (427). Voucher:
Smith 2230. PRECIS code 9901740-00550.

Tristachya rehmannii Hack.

(=Dolichochaete rehmannii
(Hack.) J.B. Phipps) 2.

Besemgras, broom trident
grass.

Perennial; tufted; 200 - 900
mm tall. Leaf blades to 200 mm
long; 1-3 mm wide. Spikelets
20-30 mm long. Leaf blades curl-
ing when old; pedicels connate; glumes and lower lemma
glabrous or with occassional tubercle-based hairs; female-
fertile (upper) lemma side-awns ( 1 8— )22(— 24) mm long,
central awns 50-100 mm long.

Flowering November to March. Shallow stony soils.
Locally common. Biome: Savanna and Grassland.

Widespread in tropical Africa. Domestic use (brooms).

Description: Chippindall & Crook 1976, Chippindall
1955 (279). Illustration: Chippindall 1955 (fig. 250).
Voucher: Liebenberg 8574. PRECIS code 9901740-00600.

Tristachya superba (De Not.) Schweinf. & Aschers.

(=Loudetia superba De
Not.) 2.

Giant trident grass.

Perennial; tufted; 1200-2700
mm tall. Leaf blades to 600 mm
long; 8-20 mm wide. Spikelets
25-35 mm long. Culms hard and
bulbous at the base; spikelets in triads, rarely in pairs, pedi-
cels unequal, 2-7 mm and 10-25 mm long respectively; fe-
male-fertile (upper) lemma side-awns 3-5 mm long, central
awns 40-120 mm long.

Flowering February to August. Granite sandveld and
Kalahari sands. Locally common (sandy areas, widespread).
Biome: Savanna. Tropical Africa. Domestic use (culms us-
ed as drinking straws by Bushmen), or pasture (roots eaten
by warthogs).

Description: Chippindall 1955 (281). Illustration:
Chippindall & Crook 1976 (76). Voucher: Ellis 2747.
PRECIS code 9901740-00700.

349

Urelytrum Hack.

Annual (rarely), or perennial; caespitose. Culms
600-2500 mm high; herbaceous (erect); unbranched above.
Leaves auriculate ( the auricles from the sheaths , glabrous
or hairy). Leaf blades linear; flat, or rolled (convolute).
Ligule an unfringed membrane. Plants bisexual, with
bisexual spikelets. The spike lets of sexually distinct forms
on the same plant ; overtly heteromorphic (the pedicellate
spikelet usually with a long-awned Gl).

Inflorescence of one to many long, rigid spike-like main
branches', digitate or subdigitate, or non-digitate; espathe-
ate; not comprising ‘partial inflorescences’ and foliar
organs. Spikelet-bearing axes ‘racemes’ (these long, many-
noded); solitary to clustered; with substantial rachides; dis-
articulating at the joints.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets with two florets, these male-only (or very rarely
with one floret hermaphrodite ), or sterile and reduced to
the glumes; the lower glume conspicuously long-awned , the
awns 5-10 mm long or longer ; Female-fertile spikelets 5-10
mm long; compressed dorsiventrally; falling with the
glumes (and with the adjacent joint and pedicel). Glumes
two; more or less equal; awned (Gl occasionally bi-
aristulate), or awnless; very dissimilar (Gl leathery,
dorsally flattened, 2-keeled, G2 thinner, naviculate-keeled).
Proximal incomplete florets 1; paleate, palea fully
developed; male.

Female-fertile florets 1. Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present; relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous. Fruit small (about 3^1 mm long); ellipsoid; hilum
short; embryo large.

Cytology, classification, distribution. Chromosome base
number,* = 10. Panicoideae; Andropogonodae; Andropo-
goneae; Rottboelliinae. 7 species. South and tropical Africa,
Madagascar. Mesophytic; in open habitats (savanna
grassland); glycophytic. Namibia, Botswana, Transvaal,
Orange Free State, Natal, and Cape Province. 1 indigenous
species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Urelytrum agropyroides (Hack.) Hack.

( -U . squarrosum Hack.) 1.

Centipede grass, kinagras,
quinine grass, varkstertgras.

Coarse perennial; tufted;

600-1600 mm tall. Leaf blades to
400 mm long; 1-6 mm wide
(rolled when young and later
when old). Spikelets (sessile) 7-8 mm long (pedicellate
smaller except for awn). Inflorescence usually a solitary
raceme; lower glume of pedicellate spikelets with a long
rough recurved awn.

Flowering October to June. Open grassland and stony
hillsides. Common. Biome: Savanna and Grassland.
Tropical Africa and Madagascar. The large, awned single-
raceme inflorcence resembles Trachypogon spicatus, which
has hairy culm nodes, and Heteropogon contortus, which
has awns from only the upper half of the inflorescence.
Also, both these species have velvety awns and lack the
bitter taste of Urelytrum.

Description: Chippindall 1955 (516), Clayton et al.
1970-1982 (833). Illustration: Chippindall 1955 (pi. 26),
Flower. PI. Afr. (47: 1841). Voucher: De Winter & Marais
4819. PRECIS code 9900170-00100.

Fig. 229. PI. 210.

\

Fig. 229 . Urelytrum agropyroides

350

Urochlaena Nees

Annual ; caespitose. Culms 70-200 mm high; herbaceous
(glabrous); much branched from the base. The uppermost
sheath blade-bearing, broadly winged from the margins in
the upper half and clasping the inflorescence. Leaf blades
linear; flat, or rolled. Ligule a fringed membrane. The
spikelets of sexually distinct forms on the same plant ;
overtly heteromorphic (those at the bases of the lower
branches 1 -flowered, or consisting of 2-A empty glumes).

Inflorescence paniculate ; deciduous in its entirety as a
‘tumbleweed’ (the culm disarticulates at the uppermost
node, complete with inflorescence and uppermost leaf);
contracted (to 25 mm long).

Female-fertile spikelets solitary; 4 mm long; com-
pressed laterally (slightly); falling with the glumes (and
with the whole inflorescence, the adjacent node and its
leaf). Glumes two; relatively large; more or less equal;
markedly shorter than the spikelets; awned (acuminate into
scabrid 8-13 mm awns); similar (ovate-oblong, acuminate,
membranous). Incomplete florets distal to the female-fertile
florets, merely underdeveloped, awned; proximal incom-
plete florets absent.

Female-fertile florets 3-7. Lemmas similar in texture to
the glumes; hairy (with fine tubercle-based marginal hairs
above, and club-shaped hairs on the mid-nerve); without a
germination flap; 7-9 nerved; entire; awned (tapering into
the awn). Awns 1; median; apical; non-geniculate (curved);
much shorter than the body of the lemma, to much longer
than the body of the lemma (but shorter than the glume
awns). Palea present (linear-oblong); relatively long
(equalling the lemma); 2-nerved. Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit small (1-2 mm);
hilum short (but relatively large); pericarp free; embryo
large.

Photosynthetic pathway. C3; XyMS+.

Cytology, classification, distribution. Chromosome base
number, ;c = 7. Arundinoideae; Danthonieae (?). 1 species.
South Africa. Xerophytic; in open habitats (in Succulent
Karoo); glycophytic. Cape Province. 1 indigenous species.

References. 1. Chippindall. 1955. Gr. & Past.

Species treatment by N.P. Barker.

Fig. 230. Urochlaena pusilla

Urochlaena pusilla Nees

Fig. 230. PI. 211.

Annual; tufted; to 200 mm
tall. Leaf blades to 30 mm long;
to 1.5 mm wide. Spikelets to 6
mm long (including awns).

Leaves expanded, soft, pubescent
or glabrous; inflorescence a
dense, spike-like panicle, 5-20
mm long and almost as wide,
partially enclosed in the up-
permost, modified leaf; spikelets 3-7-flowered; lemmas
with tubercle-based, glassy hairs on upper half and club-
shaped hairs along the central nerve and margins of the low-
er half.

Flowering September and October. Dry sandy areas and
disturbed places such as roadsides. Locally common (near
Nieuwoudtville). Biome: Succulent Karoo. Endemic. The
entire inflorescence and uppermost leaf sheath disartuculate
and are dispersed as a tumbleweed by wind.

Description; Chippindall 1955 (117). Illustration: Chip-
pindall 1955 (fig. 87 (inflorescence only)). Voucher:
Davidse 33398. PRECIS code 9903680-00100.

Urochloa P. Beauv.

Annual, or perennial; long-rhizomatous, or long-stolon-
iferous, or caespitose, or decumbent. Culms 200-1700 mm
high; herbaceous; branched above, or unbranched above.
Leaf blades linear to lanceolate; flat, or rolled. Ligule a
fringed membrane to a fringe of hairs. Plants bisexual, with
bisexual spikelets. The spikelets of sexually distinct forms
on the same plant (some spikelets reduced to disc-tipped
pedicels), or all alike in sexuality.

Inflorescence of spike -like main branches (these sessile
or subsessile)', digitate or subdigitate, or non-digitate;
espatheate. Spikelet-bearing axes persistent.

Spikelets solitary, or in pairs (or in fascicles of 3 to 4).
Female-fertile spikelets abaxial (when orientation
ascertainable)', compressed dorsiventrally; falling with the
glumes. Glumes two; very unequal, or more or less equal
(rarely); awnless; very dissimilar, or similar (membranous,
the lower sometimes tiny). Proximal incomplete florets 7;
paleate, or epaleate, palea when present fully developed to
reduced; male, or sterile.

Female-fertile florets 1 . Lemmas decidedly firmer than
the glumes; rugose', becoming indurated (crustaceous);
hairless; having the margins tucked in onto the palea; with
a clear germination flap; 5-7 nerved; entire; usually awned
(or at least strongly mucronate). Awns 1; median; apical;
non-geniculate; much shorter than the body of the lemma.
Palea present; relatively long. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small, ellipsoid to
subglobose; hilum short; embryo large.

Photosynthetic pathway. C4. The anatomical
organization conventional. Biochemical type PCK (4
species); XyMS+. PCR cell chloroplasts centrifugal/
peripheral.

Cytology, classification, distribution. Chromosome base
number, x - 7, 9, and 15. Panicoideae; Panicodae; Paniceae.
1 1 species. Tropical Africa, Asia. Mesophytic; in shade, or
in open habitats (usually: savanna, often weedy);

glycophytic. Namibia, Botswana, Transvaal, Orange Free
State, Swaziland, Natal, Lesotho, and Cape Province. 6 in-
digenous species.

References. 1. Chippindall. 1955. Gr. & Past. 2. Clayton
& Renvoize. 1982. FTEA.

Species treatment by M. Koekemoer.

351

1(0). Lower glume l/3(-l/2) the spikelet length; plants
annual; spikelets usually glabrous . U. panicoides
Lower glume 2/3 to as long as the spikelet; plants
annual or perennial; spikelets glabrous or pubescent

2(1). Plants annual 3

Plants perennial 4

3(2). Lower glume 3-nerved, the middle nerve with 1-5
stiff hairs on the back, the tip broadly rounded or

truncate U. trichopus

Lower glume 5-nerved, without stiff hairs on the back,

the tip narrowly rounded U. brachyura

4(2). Lower glume 5-nerved; spikelets lanceolate; basal
sheaths densely hairy, old sheaths splitting into

fibres; plants rhizomatous U. oligotrieha

Lower glume 3-nerved; spikelets ovate to broadly
lanceolate; basal sheaths glabrous to densely hairy,
rarely splitting into fibres; plants stoloniferOus . 5
5(4). Awn on upper lemma well developed, 0.5-1. 2 mm
long; plants 200-1500 mm tall; spikelets neatly
arranged in two rows on the rachis; lower glume

with 1-3 stiff hairs on the back

U. mosambicensis

Awn on upper lemma reduced, less than 0.5 mm long;
plants to 300 mm tall; spikelets usually untidily
arranged on the rachis; lower glume without stiff
hairs on the back U. stolonifera

Fig. 231 . Urochloa mosambicensis

Urochloa brachyura (Hack.) Stapf

Annual; coarsely tufted
(culms erect or geniculately
ascending); 200-1200 mm tall.

Leaf blades 30-300 mm long;

3-16 mm wide. Spikelets 3.5-6
mm long. Racemes (2— )5— 6(— 10),

10-60 mm long; spikelets
narrowly ovate; lower glume 2/3
the spikelet length, 5-nerved, tip
narrowly rounded and without stiff hairs on the back; upper
lemma shortly mucronate, mucro about 1 mm long.

Flowering October to April. Usually on black turf and
clayey soils in woodlands or grassveld, often in the shade.
Common. Biome: Savanna, Grassland, and Nama-Karoo.
Tropical east Africa. Closely related to U. trichopus , which
has 3-nerved lower glumes that have stiff hairs on the back.

Description: Stapf 1920 (592), Chippindall 1955 (384),
Clayton et al. 1970-1982 (606). Voucher: Tinley 1308.
PRECIS code 9901 100-00200.

Urochloa mosambicensis (Hack.) Dandy

Fig. 231. PI. 212.

(-U. pullulans Stapf) 2; (=(/.
rhodesiensis Stent) 2.

Perennial; stoloniferous and
tufted (sometimes rooting and
branching from the lower nodes);

200-1500 mm tall. Leaf blades
20-300 mm long; 3-20 mm wide.

Spikelets 3-5 mm long. Basal
sheaths glabrous or hairy, usually not splitting into fibres;
racemes (2— )3— 1 5, 20-80 mm long; lower glume 3-nerved
with 1-3 stiff hairs on the back; awn of upper lemma well
developed, 0.5- 1.2 mm long.

Flowering October to May. On a variety of soil types,
usually in sheltered disturbed places. Common. Biome: Sa-
vanna and Grassland. Tropical east Africa. Pasture
(introduced forage crop in tropical countries). Closely
related to U. stolonifera , which is a smaller plant with
spikelets untidily arranged and the upper lemma very
shortly awned. Not always clearly distinguished from U.
oligotrieha , because of the presence of intermediates.

Description: Chippindall & Crook 1976 (234), Chippin-
dall 1955 (382), Clayton et al. 1970-1982 (603).
Illustration: Chippindall 1955 (fig. 327). Voucher: Smook
5389. PRECIS code 9901 100-00400.

Urochloa oligotrieha (Fig. & De Not.) Henr.

(=(J. bolbodes (Steud.)

Stapf) 2.

Perennial; rhizomatous (rhi-
zomes stout, sometimes shortly
creeping); 600-1000 mm tall.

Leaf blades 50-100 mm long;

6-12 mm wide. Spikelets 3-5 mm
long. Basal sheaths very densely
hairy, old sheaths splitting into fibres; racemes 5-20,
30-100 mm long; spikelets lanceolate; lower glume 5-
nerved; upper lemma with mucro 0.3-0. 5 mm long.

Flowering December to May. Wooded grassland,
roadsides and old farmland, often in wet areas on clay or
loam. Locally common. Biome: Savanna and Grassland.
Tropical east Africa to Ethiopia. Natural pasture and weed
(in disturbed areas). Distinguished from U. mosambicensis
and U. stolonifera by 5-nerved lower glumes, lanceolate
spikelets and fibrous old leaf sheaths; however,
intermediates with U. mosambicensis are present.

Description: Stapf 1920 FTA (593), Chippindall &
Crook 1976 (234), Chippindall 1955 (384), Clayton et al.
1970-1982 (606). Voucher: Giess 7784. PRECIS code
9901 100-00450.

352

Urochloa panicoides Beauv.

(=(/. ruschii sensu Chippind.,
non Pilg.) 2.

Annual; tufted (erect or
prostrate; often spreading
cartweel-like); 100-900 mm tall.

Leaf blades 20-250 mm long;

5-18 mm wide. Spikelets
(2.5— )3.5— 4.5(— 5.5) mm long. In-
florescence of 2— 7(— 10) racemes. 10-90 mm long; lower
glume less than 1/2 the spikelet length; cross-veins often
present on upper glume and lemmas; awn on upper lemma
0. 3—1.0 mm long.

Flowering October to May. Weedy or overgrazed places
and in gardens and cultivation. Common. Biome: Savanna,
Grassland, and Nama-Karoo. Northwards to Sudan and
Yemen and in India. Introduced to Australia. Weed
(widespread in gardens and in cultivation). Easily
distinguished from other annual Urochloa species by its
shorter lower glume and glabrous spikelets.

Description: Chippindall & Crook 1976 (235), Chippin-
dall 1955 (385), Clayton et al. 1970-1982 (602).
Illustration: Chippindall 1955 (fig. 328). Voucher: Smook
4619, Smook & Gibbs Russell 2486. PRECIS code
9901100-00500.

Urochloa stolonifera (Goossens) Chippind.

Perennial; rhizomatous (root-
stock almost woody), or stolon-
iferous and tufted (with basal
nodes swollen); 100-300 mm tall.

Leaf blades 40-1 30 mm long; 2-9
mm wide. Spikelets 2. 5-3.0 mm
long. Racemes 2-6, 10-40 mm
long; spikelets untidily arranged
on rachis; lower glume 3-nerved,
without stiff hairs on the back; awn of upper lemma
reduced, less than 0.5 mm long.

Flowering December to April. On sandy or calcareous
soils near rivers or pans, often in disturbed places. Infre-
quent. Biome: Savanna. Closely related to U.

mosambicensis, which is a larger plant with the spikelets
neatly arranged in two rows and has a longer awn on the
upper lemma.

Description: Chippindall 1955 (381). Illustration: Chip-
pindall 1955 (fig. 326). Voucher: Zwanziger 520. PRECIS
code 9901100-00700.

Urochloa trichopus (Hochst.) Stapf

(=(/. engleri Pilg.) 2.

Annual; coarsely tufted
(usually erect with few flowering
culms); 200-1700 mm tall. Leaf
blades 50-300 mm long; 5-20
mm wide. Spikelets 2. 5-5. 5 mm
long. Racemes 3-20, 10-140 mm
long; spikelets ovate; lower

glume 2/3 the spikelet length, 3-nerved, tip broadly rounded
or truncate with a tuft of 1-5 stiff hairs on the middle nerve.

1/3 from the tip; upper lemma with mucro 0.5-1 .0 mm long.

Flowering December to April. Usually on sandy soils in
wooded grassland or on floodplains and riverbanks, often
in cultivated lands. Locally common. Biome: Savanna.
Eastern tropical Africa to Yemen. Closely related to U.
brachyura, which has 5-nerved lower glumes that lack stiff
hairs on the back.

Description: Stapf 1920 (589), Chippindall 1955 (384),
Clayton et al. 1970-1982 (604). Illustration: Clayton et al.
1970-1982 (fig. 141). Voucher: De Winter & Wiss 4163.
PRECIS code 9901 100-00800.

Vetiveria Bory

Mandelorna Steud .. Lenormandia Steud.

Perennial (with aromatic roots); forming large clumps
from stout rhizomes. Culms 500-3000 mm high; herba-
ceous; unbranched above. Leaf blades linear. Ligule a
fringed membrane to a fringe of hairs. Plants bisexual, with
bisexual spikelets; with hermaphrodite florets. The
spikelets of sexually distinct forms on the same plant
(hermaphrodite and male or neuter ); homomorphic.

Inflorescence of spike-like main branches , or paniculate
(a panicle with slender, whorled, simple or rarely
compound racemes)-, open; espatheate; not comprising
‘partial inflorescences’ and foliar organs. Spikelet-bearing
axes 'racemes’ (these with many spikelet pairs)-, with very
slender rachides; disarticulating at the joints.

Spikelets in pairs; consistently in ‘long-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets male-only, or sterile, similar to the sessile ones,
or slightly smaller. Female-fertile spikelets 4.5-10 mm
long; compressed laterally, falling with the glumes (and
with the joint and pedicel). Glumes two; more or less equal;
awned (G2, sometimes), or awnless; very dissimilar (lower
rounded on back, upper naviculate). Proximal incomplete
florets 7; epaleate; sterile.

Fig. 232. Vetiveria nigritana

353

Female -fertile florets 1. Lemmas less firm than the
glumes (hyaline); incised; awnless, or mucronate, or awned.
Awns when present 1; from the sinus; geniculate; much
shorter than the body of the lemma, to much longer than
the body of the lemma. Palea present, or absent; when
present very reduced. Lodicules 2; fleshy; glabrous.
Stamens 3. Ovary glabrous. Fruit small; hilum short;
embryo large.

Cytology, classification, distribution. Chromosome base
number, x = 5 and 10. Panicoideae; Andropogonodae;
Andropogoneae; Andropogoninae. 10 species. Tropical
Africa, Asia, Australia. Helophytic; floodplains and
streambanks; glycophytic. Namibia and Botswana. 1
indigenous species.

References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Vetiveria nigritana (Benth.) Stapf

Fig. 232. PI. 213.

Perennial; tufted; to 3000 mm
tall. Leaf blades to 90 mm long;
to 7 mm wide (margins cutting).

Spikelets (sessile) 5. 5-7.0 mm
long (pedicellate slightly shorter).

Glumes dark purple, short-spiny,
tips rounded.

Flowering July to June. Wet
places, often on black turf soil.

Infrequent. Biome: Savanna. Tropical Africa, sporadic east
to the Phillipines. V. zizanioides (L.) Nash, a native of Asia,
was formerly cultivated in the Transvaal for its scented
roots, and is grown elsewhere commercially to yield vetiver
oil.

Description: Chippindall 1955 (469), Clayton et al.
1970-1982 (739). Illustration: Chippindall 1955 (fig. 385).
Voucher: De Winter & Wiss 4125. PRECIS code
9900490-00100.

Vossia Wall. & Griff.

Perennial; long-rhizomatous. Culms 1000-2000 mm
high (above the water — from floating culms up to 7 m
long); herbaceous (aquatic, often floating, propagating from
stem fragments). Leaf blades broad\ flat. Ligule a fringed
membrane. Plants bisexual, with bisexual spikelets. The
spikelets of sexually distinct forms on the same plant, or
all alike in sexuality (the pedicellate spikelets
hermaphrodite or male); homomorphic.

Inflorescence of spike-like main branches (rarely a
single 'raceme' ); digitate or subdigitate ( usually ); espathe-
ate; not comprising ‘partial inflorescences’ and foliar
organs. Spikelet-bearing axes ‘racemes’ (spiciform,
subcylindrical or flattened, with 12 or more internodes);
clustered; with substantial rachides; disarticulating at the
joints (but rachis not very fragile).

Spikelets in pairs; consistently in Tong-and-short’ com-
binations; these pedicellate/sessile. Pedicels free of the
rachis. The sessile spikelets hermaphrodite. The pedicellate
spikelets hermaphrodite, or male-only. Female-fertile
spikelets 6-8 mm long; compressed dorsiventrally; falling
with the glumes (falling with the adjacent joint and
pedicel). Glumes two; very unequal; awned (or at least, G1
long-caudate, the tail flat); very dissimilar (G1 leathery,
flat-backed, caudate-acuminate, 2-keeled, G2 thinner,
naviculate). Proximal incomplete florets 7; paleate, palea
fully developed; male. The proximal lemmas 2 nerved.

Female-fertile florets 1 . Lemmas less firm than the
glumes (hyaline); entire; awnless. Palea present; relatively
long. Lodicules 2; fleshy; glabrous. Stamens 3. Ovary
glabrous.

Cytology, classification, distribution. Panicoideae;
Andropogonodae; Andropogoneae; Rottboelliinae. 1
species. Tropical Africa and Asia. Hydrophytic, or helo-
phytic; open habitats (swamps and river margins);
glycophytic. Namibia and Botswana. 1 indigenous species.
References. 1. Clayton & Renvoize. 1982. FTEA.

Species treatment by G.E. Gibbs Russell.

Fig. 233. Vossia cuspidata

Vossia cuspidata (Roxb.) Griff.

Fig. 233. PI. 214.

Hippo grass.

Perennial; hydrophyte; 1000-
2000 mm tall (above water, sub-
merged culms to 5000 mm long).

Leaf blades 300-1000 mm long;

6-18 mm wide. Spikelets (ses-
sile) 20-40 mm long (pedicellate
a little smaller). Lower glume
of sessile spikelets with a long flattened awn-like tail.

Flowering August to May. In permanent rivers and
lakes. Rare, but locally dominant (at riverbanks). Biome:
Savanna. Throughout tropical Africa, India. Weed.

Description: Clayton et al. 1970—1982 (832).

Illustration; Clayton et al. 1970-1982 (fig. 193). Voucher:
Gibbs Russell 2807. PRECIS code 9900160-00100.

354

Vulpia C. Gmelin

Chloammia, Distomomischus Dulac, Festucaria Link,
Loretia Duval-Jouve, Mygalurus Link, Narduretia Villar,
Nardurus (Bluff, Nees & Schauer) Reichenb., Prosphysis
Dulac, Zerna Panzer.

Annual, or perennial (rarely); caespitose. Culms 50-900
mm high; herbaceous; unbranched above. Leaf blades
linear; flat, or rolled (convolute when dry). Ligule an
unfringed membrane. Plants without hidden cleistogenes
(but all the spikelets often cleistogamous).

Inflorescence a single raceme (rarely), or paniculate',
open, or contracted; espatheate. Spikelet-bearing axes
persistent.

Spikelets secund (usually, more or less)', 5-16 mm long;
compressed laterally; disarticulating above the glumes
(also, sometimes at the base of the pedicel). Glumes two;
very unequal', markedly shorter than the spikelets; awned
(G2, sometimes), or awnless; very dissimilar (usually — G1
often minute, G2 acute to acuminate ). Incomplete florets
distal to the female-fertile florets; proximal incomplete
florets absent.

Female-fertile florets 2-15 (rarely only 1 ). Lemmas de-
cidedly firmer than the glumes (chartaceous, with thin
margins); 3-5 nerved; entire; awned. Awns 1; median; api-

cal; non-geniculate; much shorter than the body of the
lemma, to much longer than the body of the lemma. Palea
present; relatively long. Lodicules 2; membranous;
glabrous. Stamens 1-2 (rarely 3). Ovary glabrous, or hairy.
Fruit small, or medium sized, or large; hilum long-linear;
embryo small.

Cytology, classification, distribution. Chromosome base
number, x = 7. Pooideae; Poodae; Poeae. 23 species.
Temperate. Mesophytic, or xerophytic; in open habitats;
maritime-arenicolous (sometimes), or glycophytic.
Transvaal, Orange Free State, Natal, Lesotho, and Cape
Province. 4 naturalized species.

Intergeneric hybrids with Festuca — X Festulpia
Melderis ex Stace & R. Cotton (several species involved).

References. 1. Chippindall. 1955. Gr. & Past. 2. Linder.
Unpubl. ms, FSA.

Species treatment by M. Koekemoer.

1(0). Upper glume awned, 12-16 mm long (excluding

awn); awn 10-20 mm long; callus pointed

V. fasciculata

Upper glume acute or very shortly awned, 3-10 mm
long (excluding awn); awn to 2 mm long when

present; callus rounded 2

2(1). Inflorescences partially enclosed in the uppermost
leaf sheaths; lower glume often scale-like or to
■ nearly 1/2 as long as the upper glume • V. myuros
Inflorescences well exserted from the leaf sheaths;
lower glume 1/4 — 3/4 as long as the upper glume

3

3(2). Lower glume 1/2-3/4 as long as the upper glume;
spikelets usually secund and often almost
perpendicular to the central axis . . V. bromoides
Lower glume 1/4-1/2 as long as the upper glume;

spikelets mostly appressed to central axis

V. muralis

Vulpia bromoides (L.) S.F. Gray

Squirreltail fescue.

Annual; culms solitary or
numerous and then loosely tufted;

50-600 mm tall. Leaf blades
100-200 mm long; 0. 5-3.0 mm
wide. Spikelets 7-14 mm long
(excluding awns). Inflorescence
20- 1 20 mm long, to 1 5 mm wide,
well exserted from the uppermost leaf sheath; spikelets
usually secund and often almost perpendicular to the central
axis; lower glume 1/2-3/4 as long as upper; upper glume
3-10 mm long, acute or shortly awned; callus of lemma
rounded.

Flowering August to January. In weedy and disturbed
rocky places such as roadsides and along streams. Locally
common. Naturalized from Europe. Biome: Fynbos,
Savanna and Nama-Karoo. Naturalized worldwide. There
are many overlapping characters between this species, V.
muralis and V. myuros, but V. bromoides can be
distinguished by its longer lower glumes and spikelets that
are often almost perpendicular to the central axis.

Description: Linder (26), Stapf 1898-1900 (725), Chip-
pindall 1955 (60), Clayton et al. 1970-1982 (64).
Illustration: Chippindall 1955 (fig. 31), Clayton et al.
1970-1982 (fig. 22). Voucher: Smook 3675. PRECIS code
9904180-00100.

355

Vulpia fasciculata (Forssk.) Samp.

Annual; culms solitary or
loosely tufted; 100-450 mm tall.

Leaf blades 30-250 mm long; 2-5
mm wide. Spikelets 10-20 mm
long (excluding awns). Inflores-
cence partially exserted from the
uppermost leaf sheath-; glumes
unequal, lower glume 0. 5-2.0
mm long; upper glume 12-16 mm
long, awned, awn 1 0-20 mm long; callus of lemma pointed.

Flowering October to November. In weedy and
disturbed places such as gardens and roadsides, also in
coastal dunes with other alien plants. Locally common.
Biome: Fynbos. Naturalized from the coastal areas of
southern and western Europe. The long-awned upper glume
uniquely distinguishes this species from other Vulpia
species in southern Africa.

Description: Linder (25). Voucher: Smook 3714.
PRECIS code 9904180-00150.

Vulpia muralis (Kunth) Nees

Annual; culms solitary or
loosely tufted; 60-700 mm tall.

Leaf blades 1-3 mm wide. Spike-
lets 5-10 mm long (excluding
awns). Inflorescence 20-160 mm
long, well exserted from the
uppermost leaf sheath; spikelets
usually appressed to the central
axis; lower glume 1/4-1/2 as long
as upper; upper glume 3-10 mm long, acute; callus of
lemma rounded.

Flowering September to December. Generally in dry
habitats on calcareous or limestone soils and in disturbed
areas such as road verges. Locally common. Naturalized
from Europe. Biome: Fynbos and Grassland. Introduced to
the Mediterranean and the New World. Very similar to V.
bromoides and V. myuros, with which it shares many
overlapping characters. Distinguished only by the key
characters.

Description: Bor 1985 (1732), Linder (27). Voucher:
Smook 3693. PRECIS code 9904180-00200.

Vulpia myuros (L.) C. Gmel.

Ratstail fescue, langbaard-
swenkgras.

Annual; tufted (culms usually
densely fascicled); 50-700 mm
tall. Leaf blades 20-150 mm
long; 0. 5-3.0 mm wide. Spikelets
6-10 mm long (excluding awns).

Inflorescence 50-120 mm long,
partially enclosed in uppermost leaf sheath; spikelets
usually appressed to the central axis; lower glume 0. 5-2.0
mm long, often scale-like but to nearly 1/2 as long as upper;
upper glume 3-6 mm long, acute; callus of lemma rounded.

Flowering September to November. Disturbed places in
wet or damp areas but extending also to the more arid
regions. Locally common. Naturalized from western,
central and southern Europe. Biome: Fynbos, Grassland,
and Succulent Karoo. Introduced worldwide in temperate
regions. Distinguished from V. bromoides and V. muralis
by the key characters only.

Description: Bor 1985 (1733), Linder (28), Stapf
1898-1900 (724), Chippindall 1955 (61), Clayton et al.
1970-1982 (64). Illustration: Chippindall 1955 (fig. 32).
Voucher: Acocks 16522. PRECIS code 9904180-00300.

Fig. 234. PI. 215.

Willkommia Hack.

Willbleibia Herter.

Annual, or perennial; long-stoloniferous, or caespitose.
Culms 200-400 mm high; herbaceous; unbranched above.
Leaf blades linear; flat. Ligule a fringe of hairs.

Inflorescence of spike-like main branches', non-digitate
(spikes scattered along a central axis); espatheate. Spikelet-
bearing axes persistent.

Spikelets biseriate; about 4 mm long', compressed dorsi-
ventrally, disarticulating above the glumes. Glumes two;
very unequal (G1 about two-thirds length of G2); long rela-
tive to the adjacent lemmas (i.e., the upper glumes, which
slightly exceed the spikelet); awnless; similar (thin: G1
flimsier). All florets female-fertile; proximal incomplete
florets absent.

Female-fertile florets 1. Lemmas less firm than the
glumes to similar in texture to the glumes; without a ger-
mination flap; 3 nerved; entire; awnless (but acuminate), or
awned. Awns when present 1; apical; non-geniculate; much
shorter than the body of the lemma. Palea present; relatively
long (glabrous or silky-hairy). Lodicules 2; fleshy;
glabrous. Stamens 3. Ovary glabrous. Fruit ellipsoid; hilum
short; embryo large.

Fig. 235. Willkommia sarmentosa

356

Photosynthetic pathway and related features. C4;
XyMS+.

Cytology, classification, distribution. Chloridoideae;
Chlorideae sensu lato. 4 species, 1 in southern U.S.A., 3 in
southern Africa. Xerophytic; sandy savanna. Usually
halophytic. Namibia and Botswana. 3 indigenous species.

References. 1. Launert. 1970. FSWA. 2. Clayton &
Renvoize. 1986. Gen. Gram.

Species treatment by L. Smook.

1(0). Racemes several, closely arranged on the central axis;

spikelets elliptic W. newtonii

Racemes few (occasionally several), distant from one
another on the central axis; spikelets narrowly

elliptic 2

2(1). Plants annual W. annua

Plants perennial W. sarmentosa

Willkommia annua Hack.

Willkommia newtonii Hack.

Perennial (subperennial); sto-
loniferous and tufted (geniculate
at base); to 500 mm tall. Leaf
blades to 20 mm long; 3. 0-3. 5
mm wide. Spikelets 2. 5-3.0 mm
long. Leaf margins with long cilia
close together; inflorescence with
several racemes closely associ-
ated on the central axis so that the
axis is not easily visible between each raceme; spikelets el-
liptic, sometimes flushed purple, with long cilia on the cen-
tral nerve of the upper glume or if these absent with large
prickles near the apex, occasionally with hairs on the upper
glume.

Flowering March to April. Sandy soils in clearings
between tall trees. Rare. Infrequent. Biome: Savanna.
Angola. Barely distinguishable from W. sarmentosa, which
has fewer racemes arranged well apart on the central axis
and narrowly elliptic spikelets. Intermediates have been
found. The genus is in need of revision.

Description: Hackel 1896 Bull. Herb. Boiss. Ser 1,10
(810). Voucher: Giess 9305. PRECIS code 9903100-
00200.

Annual; tufted; to 600 mm
tall. Leaf blades to 30 mm long;

1.5-2. 5 mm wide. Spikelets 4—5
mm long. Leaf margins thicken-
ed, cilia present on margins, far a-
part; inflorescence with a few ra-
cemes well apart from each other
on the central axis; spikelets nar-
rowly elliptic, green; upper glume
scaberulous, especially at the apex.

Flowering January. Moist, sandy, often halophytic soils.
Infrequent. Biome: Savanna. Possibly in Angola. Barely
distinguished from the perennial W. sarmentosa.

Description: Hackel 1888 in Verh. Bot. Ver. Brand. (30:
146). Voucher: Barnard 16495. PRECIS code 9903100-
00100.

Willkommia sarmentosa Hack.

Fig. 235.

( =Craspedorhachis
sarmentosa (Hack.) Pilg.) 2.

Perennial; stoloniferous and
tufted (mat-forming); to 800 mm
tall. Leaf blades to 1 10 mm long
(usually shorter); to 5 mm wide.

Spikelets 4-5 mm long; 0.5-0. 9
mm wide. Leaves usually glau-
cous, leaf margins thickened and with long cilia that are far
apart or only occasional; inflorescence usually with only a
few racemes distant from each other; spikelets narrowly el-
liptic, green; upper glume scaberulous with minute prickles
especially near the apex.

Flowering November to March (and July). Moist sandy,
often halophytic soils along edges of pans and marshes, or
seasonally waterlogged areas. Locally common. Biome: Sa-
vanna. Zimbabwe, Zambia. W. annua is can be recognized
as an annual form, but W. newtonii is barely distinguishable
and intermediates are found. The genus is in need of
revision.

Description: Launert 1970 (160:49), Chippindall 1955
(204). Illustration: Chippindall 1955 (fig. 181). Voucher:
Giess & Muller 13953. PRECIS code 9903100-00300.

SPIKELET PHOTOGRAPHS

scale bar = 1 mm

357

PI. 1 . Acrachne racemosa,
6-9 mm

PI. 4. Aira cupaniana,
2-3 mm

PI. 7. Ammophila arenaria ,
10-15 mm

PI. 2. Acroceras macrum,
4-5 mm

PI. 3. Agrostis eriantha ,
3. 5-5.0 mm

PI. 5. Alloteropsis semialata
subsp. eckloniana, 5-8 mm
(side view)

PI. 8. Andropogon chinensis,
5-7 mm (spikelet pair)

PI. 6. Alloteropsis semialata
subsp. eckloniana , 5-8 mm
(abaxial view)

PI. 9. Anthephora ramosa,
6-7 mm (spikelet cluster)

358

scale bar = 1 mm

PI. 10. Anthephora schinzii ,
to 10 mm (spikelet cluster)

PI. 11. Anthoxanthum ecklonii,
6-8 mm

PI. 12. Aristida adscensionis ,
10-40 mm (incl. awns)

PI. 13. Arrhenatherum elatius,
7-1 1 mm

PI. 14. Arthraxon lanceolatus,
5.0-6. 5 mm (spikelet pair)

PI. 15. Arundinella nepalensis ,
4-6 mm

scale bar = 1 mm

359

PI. 20. Bewsia biflora ,
5. 5-9.0 mm

PI. 23. Brachiaria deflexa ,
2.0-3.4 mm (spikelet pair)

PI. 26. Brachychloa

schiemanniana , 4-7 mm

PI. 21. Bothriochloa insculpta,
4. 5-5.0 mm (spikelet pair)

PI. 24. Brachiaria serrata,
2. 3-4. 5 mm

PI. 27. Brachypodium flexum,
12-44 mm

PI. 19 . Bambusa balcooa ,
7-16 mm

PI. 22. Brachiaria brizantha ,
4-6 mm

PI. 25. Brachyachne

patentiflora, 3. 0-4. 4 mm

360

scale bar = 1 mm

PI. 28. Briza maxima,
8-25 mm

PI. 29. Briza subaristatum ,
4-5 mm

PI. 30. Bromus catharticus ,
20-35 mm

PI. 31. Calamagrostis epigeios
var. capensis, 5. 5-8.0 mm

PI. 32. Catalepis gracilis,
4-5 mm

PI. 33. Catapodium rigidum,
5-7 mm

PI. 34. Cenchrus ciliaris,

4-5 mm (spikelet cluster)

PI. 35. Centropodia glauca,
7.5-10.0 mm

PI. 36. Chaetobromus
dregeanus, 12-18 mm

scale bar = 1 mm

361

PI. 37. Chloris virgata,
3.0-3. 5 mm

PI. 40. Cleistachne sorghoides,
4-5 mm

PI. 43. Coix lacryma-jobi ,
25-35 mm (partial
inflorescence)

PI. 38. Chrysopogon

serrulatus, 5-8 mm (triad
of spikelets)

PI. 41. Coelachyrum
yemenicum , 5-10 mm

PI. 44. Colpodium hedbergii ,
2. 5-4.0 mm (three
spikelets)

PI. 39. Clador aphis spinosa,
6-18 mm

PI. 42. Coelorhachis capensis,
4. 5-5.0 mm (spikelet pair)

PI. 45. Cortaderia selloana,
to 15 mm

362

scale bar = 1 mm

PI. 46. Corynephorus
fasciculatus, to 3 mm

PI. 49. Cymbopogon

marginatus , 5.0-6. 5 mm
(spikelet pair)

PI. 52. Dactylis glomerata,
5-9 mm

PI. 47. Craspedorhachis
africana, 3-4 mm (several
spikelets)

PI. 50. Cynodon dactylon,
2. 0-2. 5 mm (several
spikelets)

PI. 53. Dactyloctenium
giganteum, 4.0-6.2 mm

PI. 48. Ctenium concinnum,
5-7 mm

PI. 51. Cynosurus color atus,
10-25 mm

PI. 54. Danthoniopsis dinteri,
14-20 mm

scale bar = 1 mm

363

PI. 55. Deschampsia cespitosa,
3. 5-6.0 mm

PI. 58. Digitaria eriantha,

2. 2-4.0 mm (two spikelets)

PI. 61. Dinebra retroflexa,
5.7-9. 0 mm (several
spikelets)

PI. 56. Diandrochloa
namaquensis , 2-3 mm
(several spikelets)

PI. 59. Digitaria monodactyla ,
2. 8-3. 2 mm (abaxial view)

PI. 62. Diplachne fusca,
6-14 mm

PI. 57. Dichanthium

annulatum, 2. 5-5.0 mm
(spikelet pair)

PI. 60. Diheteropogon
amplectens, 7-9 mm
(spikelet pair)

PI. 63. Dregeochloa pumila,
9-13 mm

364

PI. 64. Echinochloa crus-galli,
3-7 mm

PI. 67. Ehrharta longiflora ,
10-25 mm (incl. awns)

PI. 70. Elymandra grallata,
6.5-12.0 mm (spikelet pair)

scale bar = 1 mm

PI. 65. Ehrharta calycina ,
4. 0-8. 5 mm

PI. 66. Ehrharta capensis,
8-12 mm

PI. 68. Eleusine coracana
subsp. africana , 5-8 mm

PI. 69. Elionurus muticus ,
6-14 mm (spikelet pair)

365

scale bar = 1 mm

PI. 73. Enneapogon
cenchroides , 3-5 mm

PI. 74. Enteropogon

macrostachyus , 8-10 mm

PI. 75. Entolasia imbricata,
4. 5-6. 5 mm

PI. 76. Entoplocamia
aristulata, 9-17 mm

PI. 77. Eragrostis capensis ,
3.5-15.0 mm

PI. 78. Eragrostis curvula,
4-10 mm

PI. 79. Eragrostis racemosa,
3-10 mm

PI. 80. Eragrostis superba ,
6-16 mm

PI. 81. Eriochloa meyeriana
subsp. meyeriana
2. 5-3. 5 mm

366

scale bar = 1 mm

PI. 82. Eriochrysis pallida,

3. 5-5.0 mm (spikelet pair)

PI. 85. Festuca costata,
10-20 mm

PI. 88. Hackelochloa
granularis, 1.0- 1.5 mm
(two spikelet pairs)

PI. 83. Eulalia villosa ,

5-7 mm (spikelet pair)

PI. 86. Fingerhuthia africana,
4.0-5. 5 mm

PI. 89. Hainardia cylindrica,
5-8 mm (several spikelets)

PI. 84. Eustachvs paspaloides ,
1 .5-2.5 mm

PI. 87. Gastridium phleoides,
5-7 mm (two spikelets)

PI. 90. Harpochloa falx,
6-9 mm

PI. 91. Helictotrichon
turgidulum,

10-12 mm

PI. 94. Holcus lanatus,
3-4 mm

scale bar = 1 mm

367

PI. 92. Hemarthria altissima,
5-7 mm (two spikelet pairs)

PI. 93. Heteropogon contortus,
5. 5-7.0 mm (spikelet pair)

PI. 95. Hordeum murinum,
20-35 mm (incl. awns
and sterile spikelets)

PI. 98. Imperata cylindrica,
3-6 mm (spikelet pair)

PI. 96. Hyparrhenia hirta,

4. 0-6. 5 mm (spikelet pair)

PI. 99. Ischaemum afrum ,
5-8 mm (spikelet pair)

368

scale bar = 1 mm

PI. 100. Kaokochloa
nigrirostris , to 7 mm

PI. 101. Karroochloa purpurea ,
5-7 mm

PI. 102 . Koeleria capensis ,
3. 5-4.0 mm

PI. 103. Lagurus ovatus ,

7-10 mm (several spikelets)

PI. 104. Lamarckia aurea,
6-9 mm

PI. 105. Leersia hexandra ,
3.4-4. 8 mm

PI. 106. Leptocarydion
vulpiastrum , 5-11 mm

PI. 107 . Leptochloa panicea,
1.9-2. 5 mm

PI. 108. Lepturus repens,
10-14 mm (incl. awns)

scale bar - 1 mm

369

PI. 109 . Leucophrys
mesocoma, to 7 mm

PI. 110. Lintonia nutans,
6-10 mm

PI. 111. Lolium multiflorum ,
8-20 mm

PI. 112 . Lophachme digitata,
5-6 mm

PI. 113. Lophochloa pumila,
2. 5-4.0 mm

PI. 114. Loudetia simplex,
7-13 mm

PI. 115. Megaloprotachne
albescens, 4. 0-4. 5 mm (two
spikelets)

PI. 116 . Megastachya
mucronata , 7-15 mm

PI. 1 17. Melica racemosa,
5-9 mm

370

scale bar = 1 mm

PI. 118. Melinis minutiflora,
1. 5-2.0 mm (several
spikelets)

PI. 121 . Merxmuellera stricta,
to 23 mm

PI. 124. Microlaena stipoides,
9-1 1 mm (excl. awns)

PI. 1 19. Melinis repens

subsp. repens , 2. 2-4.0 mm

PI. 122. Microchloa caffra ,
3. 0-5. 5 mm (several
spikelets)

PI. 125. Microstegium nudum ,
3. 5-4.5 mm (terminal
spikelets)

PI. 120. Merxmuellera
arundinacea ,
13.5-16.5 mm

PI. 123. Microlaena stipoides,
20-30 mm (incl. awns)

PI. 126 . Miscanthus capensis,
4-6 mm

scale bar = 1 mm

371

PI. 127 . Monelytrum
luederitzianum ,

3-4 mm (spikelet cluster)

PI. 130. Nassella trichotoma ,
6. 0-8. 5 mm

PI. 128 .Monocymbium
ceresiiforme,

3. 5-4.0 mm (spikelet pair)

PI. 131. Odontelytrum
abyssinicum, to 12 mm

PI. 129 .Mosdenia

leptostachys , 2. 5-3. 7 mm

PI. 132. Odyssea paucinervis,
5-9 mm

PI. 133 . Olyra latifolia,
7-10 mm

PI. 134 . Oplismenus hirtellus,
2-4 mm

PI. 135. Oropetium capense,
2.5-4. 0 mm (rachis and
several spikelets)

372

scale bar - 1 mm

PI. 136. Oryza longistaminata,
7-9 mm

PI. 139 . Panicum maximum,
2. 5-3.0 mm

PI. 142. Paratheria prostrata,
to 9 mm

PI. 137. Oryzidium barnardii,
8-10 mm

PI. 140. Panicum natalense,

1.7-2. 2 mm (several spikelets)

PI. 138. Oxyrhachis
gracillima, 3-6 mm
(spikelet pair)

PI. 141. Parapholis incurva,

4. 5-5. 5 mm (several spikelets)

PI. 143 . Paspalidium

obtusifolium, 3.0-3. 5 mm

PI. 144. Paspalum dilatatum,
3-4 mm

scale bar = 1 mm

373

PI. 145. Pennisetum setaceum ,
4.0-6. 5 mm (spikelet
cluster)

PI. 148. Pentaschistis galpinii,
4-6 mm

PI. 146. Pentameris thuarii,
16-22 mm

PI. 149. Pentaschistis pusilla,
2-4 mm

PI. 147 . Pentaschistis
curvifolia ,

1 1-14 mm

PI. 150 . Perotis patens,
1 .2-2.7 mm

PI. 151. Phacelurus franksae,
6-8 mm (spikelet pair)

PI. 152. Phalaris aquatica,
4. 5-7. 5 mm

PI. 153 . Phragmites australis,
12-18 mm

374

scale bar - 1 mm

PI. 154. Poa annua ,
4-6 mm

PI. 157 . Polypogon
monspeliensis,

2-3 mm (two spikelets)

PI. 160. Pseudechinolaena
polystachya, 3. 5-5.0 mm

PI. 155 . Pogonarthria
squarrosa, 3. 3-7. 8 mm

PI. 158 . Prionanthium
pholiuroides, 3-7 mm

PI. 161 . Pseudopentameris
macrantha , 30-40 mm

PI. 156. Polevansia rigida ,
3. 5-4. 5 mm

PI. 159 . Prosphytochloa
prehensilis, 6-9 mm

PI. 162 . Puccinellia
acroxantha, 3-5 mm

scale bar = 1 mm

375

PI. 163 . Rendlia altera ,
4.0-5. 5 mm (several
spikelets)

PI. 166. Sacciolepis typhura ,
1.7-2. 5 mm

PI. 164 . Rhytachne

rottboellioides, 3-5 mm
(spikelet pair)

PI. 167 . Sartidia angolensis,
90-120 mm (incl. awns)

PI. 165 . Rottboellia
cochinchinensis,

4-7 mm (spikelet pair)

PI. 168 . Schismus barbatus,
4-7 mm

PI. 169. Schizachyrium
sanguineum , 6-9 mm
(spikelet pair)

PI. 170. Schmidtia

pappophoroides, 8-15 mm

PI. 171. Schoenefeldia
transiens, 3. 5 -5.0 mm

376

scale bar = 1 mm

PI. 172. Secale africanum,
10-15 mm

PI. 175. Setaria sphacelata
var. torta , 2. 5-3.0 mm

PI. 178. Sorghum bicolor
subsp. arundinaceum,
5-7 mm (spikelet pair)

PI. 173. Sehima galpinii,
12-15 mm (spikelet pair)

PI. 176. Setaria verticillata,
1 .5-2.5 mm

PI. 174. Setaria
lindenbergiana ,
2.0-3. 5 mm

PI. 177 . Sorghastrum friesii,
5-7 mm

PI. 181 . Sporobolus

centrifugus , 2. 5-4. 2 mm

PI. 184. Stenotaphrum

secundatum, 4-5 mm (two
spikelets)

PI. \%1 . Stipa dregeana
var. elongata , 5-7 mm

scale bar - 1 mm

PI. 182. Sporobolus

discosporus, 1 .0-1.7 mm
(several spikelets)

PI. 185 . Stereochlaena
cameronii, 2. 0-3. 5 mm
(two spikelets)

PI. 188. Stipagrostis anomala,
11-14 mm

377

PI. 183. Sporobolus
fimbriatus, 1.4-2. 2 mm
(several spikelets)

PI. 186. Stiburus

alopecuroides , 2. 7-4.0 mm

PI. 189. Stipagrostis uniplumis
var. neesii, 10.0-14.5 mm

378

scale bar = 1 mm

PI. 190. Stipagrostis zeyheri
subsp. zeyheri , 16-19 mm

PI. 193 .Tarigidia

aequiglumis , 4. 0-4. 5 mm

PI. 196. Thamnocalamus
tessellatus, 16-18 mm

PI. 191 . Strehlochaete
longiarista, 15-25 mm

PI. 194 .Tetrachne dregei,
4-6 mm

PI. 197. Thelepogon elegans ,
5-13 mm (spikelet pair)

PI. 192. Styppeiochloa
gynoglossa , 5-7 mm

PI. 195. Tetrapogon tenellus ,
3. 5-5.0 mm

PI. 198. Themeda triandra ,
about 60 mm (spikelet
cluster)

scale bar = 1 mm

379

PI. 199 . Themeda triandra,
5-7 mm (spikelet pair)

PI. 202. Tragus berteronianus ,
2. 0-3. 8 mm (spikelet
cluster)

PI. 205. Trichoneura

grandiglumis , 5-14 mm

PI. 200. Thinopyrum
distichum , 28-40 mm

PI. 203. Tribolium uniolae,
to 6 mm

PI. 201. Trachypogon spicatus,
4-8 mm (spikelet pair)

PI. 204. Tricholaena
monachne, 2-3 mm

PI. 207. Tripogon minimus,
2. 6-8.0 mm

380

scale bar = 1 mm

PI. 208. Trir aphis
andropogonoides ,
6-10 mm

PI. 21 1. Urochlaena pusilla ,
to 6 mm

PI. 214. Vossia cuspidata ,
20-40 mm (spikelet pair)

PI. 209. Tristachya leucothrix,
24-45 mm (triad of spikelets)

PI. 212. Urochloa

mosambicensis, 3-5 mm

PI. 215. Vulpia myuros ,
6-10 mm

PI. 210. Urelytrum
agropyroides ,

7-8 mm (spikelet pair)

PI. 213. Vetiveria nigritana ,
5. 5-7.0 mm (spikelet pair)

PI. 216. Willkommia annua ,
4-5 mm

381

APPENDIX 1:

SUMMARIZED CLASSIFICATION OF SOUTHERN AFRICAN GRASSES

There are few absolute criteria for reliably referring grasses
to subtribes, tribes, supertribes and subfamilies. These higher
levels of classification are recognisable only in terms of correlat-
ed tendencies among suites of characters. It will be apparent,
therefore, that the classification provided here has no identifica-
tory role. On the contrary, this illustrates an important general
principle of taxonomy: identification is most effectively pursued
at the lowest available hierarchical level. When identification has
been satisfactorily achieved at the level of species or genus, using
a printed key or a microcomputer cross-referenced with printed
descriptions, the relationships of the organism are readily ascer-
tained.

Detailed classificatory information, including group descrip-
tions and diagnostic characters, are readily obtainable from the
automated generic data by application of the program INTKEY.
It seemed more appropriate here to present a summarized classifi-
cation of the southern African grasses, with relatively brief de-
scriptions, because the comprehensive and fully comparative de-
scriptions derived from the complete character list are so complex
as to be quite intimidating. Exclusion of genera not represented
in southern Africa has permitted some simplification of the group
descriptions, through omission of ‘exceptional' taxa, but we have
tried to ensure that the descriptions remain reasonably representa-
tive of the groups as world entities.

The characters printed in boldface approximate a diagnostic
description for each group. It is not possible to be strictly diag-
nostic at these higher levels because of the paucity of absolute
criteria separating the groups, as mentioned above.

The classification is intended to fulfil an introductory, educa-
tional role, and to provide guidance for defining appropriate
samples to use in experimental work. Therefore, both ‘exoteric’
characters, of the kind acceptable in general purpose keys, and
‘esoteric’ characters (anatomy, physiology, cytology etc.) which
constitute essential components of modern taxonomic classifica-
tion are included. We also wished to illustrate the diversity of in-
formation that can be applied to taxonomic group-making, and
to hint at the range of biological disciplines with which grass tax-
onomy usefully exchanges information. For this reason the de-
scriptions summarize taxonomic patterns observable in 2c DNA
values, chromosome numbers and susceptibilities to pathogens,
notwithstanding that most of the available information derives
from observations on species not represented in southern Africa.

Circumscriptions of the subfamilies and supertribes employed
here have been discussed in detail elsewhere (Watson et al. 1985,
Watson 1987). Apart from the placement of some controversial
genera and small groups (most of which are not represented in
southern Africa), these groupings are widely accepted as a rea-
sonable taxonomic interpretation of the facts (cf. Clayton &
Renvoize 1986, Soderstrom et al. 1987). The Centotheceae (rep-
resented in southern Africa only by Megastachya) are not very
convincingly bambusoid, but a better location has yet to be agre-
ed upon. The Arundinoideae constitute an unsatisfactory subfam-
ily which is not amenable to anything approaching a diagnostic
description, and which is probably polyphyletic (see Watson et
al. 1985 and Kellogg & Campbell 1987 for detailed analyses in
phenetic and phylogenetic terms, respectively). However, the re-
lationships of the individual arundinoid tribes with other sub-
families remain obscure. Even when their relationships are satis-
factorily resolved, difficulties of practical implementation will
remain. Including arundinoid tribes in other subfamilies (e.g.
returning the Stipeae to the Pooideae (Clayton & Renvoize 1986))
extends the diversity of the latter so as to undermine practical
usefulness; on the other hand, promoting the tribes to subfamilies
amounts to a kind of nomenclatural inflation which contributes
nothing to the portrayal of inter-subfamilial relationships.

The tribes used here are also fairly conventional. However,
they have been subjected to original, critical (but as yet incom-
plete) analyses in terms of the generic descriptions (Macfarlane
& Watson 1982; Watson unpublished), and those which have not
proved amenable to adequate definition in terms of tangible cor-
relations of characters have been rejected. Detailed analyses of
the world data on the Andropogoneae detected ill-defined, low-

level groupings similar to those set out by Clayton (1972, 1973),
which scarcely seem to merit nomenclatural recognition. How-
ever, they also revealed the "awned Andropogoneae" and the
"awnless Andropogoneae" of informal parlance as rather clearly
defined series, whose recognition here as subtribes reflects a
genuinely informative distinction.

Pooideae

Culms 2-200 cm high; herbaceous; unbranched above;
usually with hollow internodes. Leaf sheaths occasionally with
joined margins. Leaf blades linear to linear-lanceolate; not
pseudopetiolate; without readily visible transverse veins; not
disarticulating. Adaxial ligule an unfringed membrane.
Abaxial ligule absent. Inflorescence determinate; usually
paniculate, occasionally a raceme, a spike or spicate with clusters
of spikelets, but never comprising spikelike main branches;
espatheate. Spikelets hardly ever in distinct long-and-short
combinations. Female-fertile spikelets nearly always laterally
compressed or terete (dorsiventrally compressed in Hainardia);
with or without an apically prolonged rachilla. Glumes present;
usually 2 (one in Hainardia, Lolium); usually similar (dissimilar
in Vulpia). Lower glume 1— 5(— 1 1) nerved. Upper glume 1 — 7(— 1 2)
nerved. Spikelets with female-fertile florets only, or having
incomplete florets. Incomplete florets when present usually distal
(proximal in Anthoxanthum , Phalaris). Female-fertile florets
1-30. Lemmas entire or incised (not deeply cleft); hairs when
present neither in tufts nor in transverse rows; without a
germination flap; ( 1— )3— 7(— 15) nerved; often awned. Lemma
awns when present 1 or 3, the median (or only) awn apical, from
a sinus or dorsal; geniculate or non-geniculate. Palea present,
usually 2-keeled (rarely keel-less); usually apically notched.
Lodicules present; 2; usually membranous and free (exception
Melica); glabrous or ciliate. Ovary glabrous or hairy. Styles 2;
nearly always free to their bases. Stigmas 2; white. Fruit a
caryopsis, often longitudinally grooved. Hilum short or long-
linear. Embryo small; usually with an epiblast, with neither
mesocotyl internode or scutellar tail, the embryonic leaf
margins meeting.

Abaxial leaf blade epidermis. Microhairs absent. Mid-
intercostal long-cells more or less rectangular or (almost as often)
fusiform; their walls markedly sinuous or (about as commonly)
more or less straight. Costal silica bodies variously crescentic,
tall-and-narrow or rounded, but more often ‘pooid-type’ (i.e.
elongated, sinuous or crenate) or elongated-smooth, and hardly
ever ‘panicoid-type’ or saddle-shaped. Costal short-cells only
infrequently in long rows, usually solitary, in short rows and/
or pairs. Stomatal guard-cells nearly always overlapped by the
interstomatals (exception: Vulpia); subsidiaries parallel-sided or
dome-shaped. Transverse section of leaf blade, physiology. C}.
XyMS+. Blade usually adaxially ribbed, much less frequently
flat; the ribs usually more or less constant in size. Mesophyll not
traversed by colourless columns; without arm-cells; without
fusoids. Midrib conspicuous or inconspicuous; nearly always
with a single bundle , rarely a simple arc; without adaxial
colourless tissue. Bulliforms often present, usually as simple fans
(very rarely combining with colourless cells to form deeply-
penetrating fans). Only infrequently exhibiting small vascular
bundles unaccompanied by sclerenchyma. Very rarely exhibiting
sclerenchyma additional to that associated with the vascular
bundles.

Chromosome base number usually x = 7 (rarely 2, 5, 9-10,
13, 19). Mean diploid 2c DNA value 2.3-17.7, group mean 8.9.

Rusts: Puccinia species. Smuts: species of Entyloma , Tilletia,
Urocystis, Ustilago (only questionable records for
Sphacelotheca , none for Sorosporium).

382

Triticodae

Leaves often auriculate. Inflorescence usually a distichous
spike or spicate with clusters of spikelets, or a raceme, rarely
a panicle. Inflorescence axes persistent or (commonly)
disarticulating at the joints. Spikelets often large (5-70 mm
long). Female-fertile spikelets with the rachilla prolonged beyond
the uppermost female-fertile floret, with distal incomplete florets.
Glumes often lateral to the rachis or displaced to the front
of the spikelet, sometimes joined, sometimes subulate. Female-
fertile florets 1-30. Lemmas often awned (awns 1, rarely 3), the
(median) awn non-geniculate, usually apical or from a sinus,
usually entered by several veins. Ovary apex hairy, lodicules
often ciliate. Caryopsis longitudinally grooved, with a ling-linear
hilum. Endosperm hard, without lipid, containing only simple
starch grains. Embryo sometimes without an epiblast.

Abaxiai leaf blade epidermis. Crown cells often present.
Stomata sometimes very large (up to 84 microns), the apparatus
usually conspicuously sunken.

Often xerophytic

Mean diploid 2c DNA value 3.7-16.8 pg, group mean 10.63.

Triticeae : Elytrigia, Hordeum. Secale, Thinopyrum.

Culms 5-170 cm high; herbaceous; unbranched above. Culm
internodes solid or hollow. Leaves often auriculate. Adaxial
ligule an unfringed membrane. Inflorescence a single spike, or
a false spike with clusters of spikelets; the axes disarticulating
at the joints, or persistent. Female-fertile spikelets 7-23 mm long;
compressed laterally to terete; disarticulating above the glumes,
falling with them, or (in cultivated forms) not disarticulating;
with the rachilla prolonged apically. Glumes two; lateral to the
rachis or displaced; similar; 1—12 nerved. Spikelets with or
without incomplete florets, these when present distal. Female-
fertile florets 1-10. Lemmas awnless to awned, the awn when
present from a sinus or apical, non-geniculate. Lemmas keeled
or not; without a germination flap; 5(-l 1 ) nerved. Palea relatively
long; 2-nerved. Lodicules 2; membranous; ciliate. Ovary hairy.
Stigmas white. Fruit, embryo. Hilum long-linear. Endosperm
hard, starch grains simple. Embryo large or (more often) small;
with or without an epiblast; with neither scutellar tail nor
mesocotyl internode, the embryonic leaf margins meeting.
Abaxiai leaf blade epidermis. Crown cells often present.

Basic chromosome number, x = 7. 2n = 14-84.

Brachypodieae : Brachypodium. (See genus for description.)

Bromeae: Bromus. (See genus for description.)

Poodae

Leaves rarely auriculate (exceptions in Poeae). Inflorescence
nearly always a panicle (a spike in Lolium, Hainardia), the axes
persistent (except Hainardia). Spikelets often small (l-25(-45)
mm long). Female-fertile spikelets with or without the rachilla
prolonged beyond the uppermost female-fertile floret, with or
without incomplete florets, incomplete florets when present
usually distal, occasionally proximal (Anthoxanthum, Phalaris).
Female-fertile florets 1-22. Lemmas awnless, mucronate or
awned; awns 1 or 3, the (median) awn from a sinus, apical or
dorsal, geniculate or non-geniculate, usually entered by only one
vein. Ovary apex occasionally hairy (e.g. Festuca), usually
glabrous; lodicules nearly always glabrous (exceptions
Ammophila, Festuca). Caryopsis longitudinally grooved or not,
the hilum short or (less often) long-linear. Endosperm hard or
liquid, with lipid or (less often) without, usually containing
compound starch grains. Embryo nearly always with an
epiblast.

Abaxiai leaf blade epidermis. Crown cells absent. Stomata
21-54 microns long, mean 57 microns.

Helophytic, mesophytic or xerophytic.

Mean diploid 2c DNA value 2.3-17.7 pg, group mean 7.9.

Aveneae (including Agrostideae, Phalarideae): Agrostis, Aira,
Ammophila, Anthoxanthum, Arrhenatherum, Avena,
Calamagrostis, Corynephorus, Deschampsia, Gastridium,
Helictotrichon, Holcus, Koeleria, Lagurus, Lophochloa,
Periballia, Phalaris, Polypogon

Culms 2-200 cm high; herbaceous; unbranched above; with
hollow internodes. Leaves non-auriculate, sheath margins free.
Adaxial ligule an unfringed membrane. Inflorescence nearly

always a panicle, rarely a raceme; the axes persistent. Spikelets
not secund. Female-fertile spikelets 0.8^45 mm long; compressed
laterally; usually disarticulating above the glumes, occasionally
falling with them or not disarticulating; with or without an
apically prolonged rachilla. Hairy callus commonly present.
Glumes 2; similar; sometimes carinate; 1-3 (-17) nerved; the
upper nearly always long relative to the adjacent lemmas.
Spikelets with or without incomplete florets, these when present
proximal, distal or both distal and proximal. Proximal incomplete
florets when present 1 or 2, paleate or epaleate, male or sterile.
Female-fertile florets l-2(-7). Female-fertile lemmas only
infrequently carinate, awnless or 1-, 3- or 5-awned. The
(median) awn from a sinus or dorsal, non-geniculate or more
often geniculate. Lemmas without a germination flap;
( 1— )3— 7(— 9) nerved. Palea usually relatively long, occasionally
reduced and very short; (l-)2(-several) nerved, rarely nerveless.
Lodicules nearly always present, nearly always membranous and
glabrous. Ovary usually glabrous, rarely hairy. Fruit, embryo.
Hilum short or long-linear. Endosperm hard or liquid, usually
with lipid. Starch grains usually compound.

Basic chromosome number,* = 4—13 (usually 7). 2 n = 8-147.

Meliceae : Melica, Streblochaete .

Leaves with sheath margins joined. Inflorescence a raceme
or panicle. Female-fertile spikelets laterally compressed or terete,
with distal incomplete florets; with an apically prolonged
rachilla; disarticulating above the glumes. Hairy callus absent.
Glumes non-carinate. Female-fertile florets 1-7. Lemmas not
carinate. Lodicules joined or free, fleshy or membranous,
glabrous or ciliate. Ovary glabrous. Endosperm hard.

Basic chromosome number, * = 9 or 10.

Poeae (including Hainardieae, Monermeae): Briza, Catapodium,
Colpodium. Cynosurus, Dactylis, Festuca, Hainardia,
Lamarckia, Lolium, Parapholis, Poa, Puccinellia, Sphenopus,
Vulpia.

Culms 2-200 cm high; herbaceous; rarely branched above.
Nodes glabrous, intemodes nearly always hollow. Leaves
sometimes auriculate, sheath margins occasionally joined.
Adaxial ligule an unfringed membrane. Inflorescence
occasionally a spike or a raceme, usually a panicle; the axes
occasionally disarticulating, usually persistent; the spikelets
sometimes secund, occasionally two-ranked and distichous.
Female-fertile spikelets 1.5-26 mm long; nearly always
compressed laterally, rarely terete or dorsiventrally compressed;
usually disarticulating above the glumes, rarely falling with them;
with the rachilla prolonged or not. Hairy callus very rarely
present. Glumes usually two and similar, rarely only one.;
commonly short relative to the adjacent lemmas; carinate or
non-carinate. Lower glume (0— ) 1— 3(— 1 5) nerved; upper glume
1-7(8-15) nerved. Spikelets with or without incomplete florets,
these when present distal. Female-fertile florets 1-22. Lemmas
awnless or 1-awned from a sinus, apically or dorsally; the awn
when present usually non-geniculate. Lemmas carinate or not,
( 1 — )3— 7(— 1 5 nerved. Palea usually relatively long, occasionally
reduced and very short; 2-nerved. Lodicules 2, membranous,
free, nearly always glabrous. Ovary usually glabrous, sometimes
hairy but without an apical appendage. Fruit, embryo. Hilum
short or long-linear. Endosperm liquid or hard, with or without
lipid. Starch grains usually compound.

Basic chromosome number,* usually = 7 (occasionally 2, 5-6,
9, 13, 19); 2 n = 4-1 17.

Bambusoideae

Mostly perennial, culms woody or herbaceous; often hut not
always overtly ‘bambusoid’ in appearance. Leaves rarely
basally aggregated, sometimes auriculate and often with auricular
setae. Leaf blades linear to elliptic (i.e often relatively broad),
often pseudopetiolate, often with readily visible transverse
veins, commonly disarticulating. Abaxiai ligules common.
Inflorescence sometimes indeterminate (sometimes with
‘pseudospikelets’), usually paniculate, the axes usually
persistent, often spatheate. Glumes 1-several, usually similar,
sometimes minute or lacking. Spikelets frequently with
incomplete florets, these proximal, distal or both proximal and
distal to the female-fertile florets. Proximal incomplete florets
when present usually more than one (except Olyra). Female-

383

fertile florets 1-30. Lemmas usually entire, awned (with a single,
apical non-geniculate awn) or awnless; hairy (the hairs not in
tufts or horizontal rows); occasionally with hairy (the hairs not
in tufts or horizontal rows), occasionally with a germination flap.
Palea present, usually relatively long; nerves 1 , 2 or several, keel-
less, 1-keeled or 2-keeled. Lodicules usually present, l-5(-10),
often 3\ ciliate or glabrous, often heavily vascularized. Stamens
variable in number, often more than 3. Ovary apex glabrous or
hairy. Stigmas l^f, often 3. Fruit sometimes with a free pericarp,
this sometimes thick and hard or fleshy; longitudinally grooved
or not. Hilum occasionally short, usually long-linear. Embryo
usually small; with an epiblast; usually with a scutellar tail
and overlapping embryonic leaf margins. Endosperm without
lipid.

Abaxial leaf blade epidermis. Microhairs usually present;
panicoid-type. Mid-intercostal long-cells rectangular or
fusiform, with markedly sinuous walls. Papillae often present,
often overarching the stomata, usually several or many per
long-cell. Costal silica bodies often panicoid-type, oryzoid or
saddle-shaped, hardly ever pooid-type, elongated-smooth or
rounded. Stomatal guard-cells overlapping the interstomatals,
flush with or overlapped by them; the subsidiaries usually
trianguar or dome-shaped, but occasionally parallel-sided.
Transverse section of leaf blade, physiology. C3. XyMS+. Blade
commonly adaxially flat. Mesophyll not traversed by colourless
columns; often with arm-cells and/or fusoids. Midrib usually
conspicuous; with one bundle, a conventional arc or (often) with
complex vascularization. Bulliforms usually present, usually on
simple fans, very rarely associated with colourless cells to form
deeply-penetrating fans. All the vascular bundles accompanied
by sclerenchyma. Hardly ever exhibiting sclerenchyma additional
to that directly associated with the vascular bundles (other than
in midribs).

Chromosome base number, x = 10, 11, 12, 15 or 19 (nearly
always 10, 11 or 12).

Hydrophytic to mesophytic, rarely xerophytic. Often shade
plants.

Rusts: Dasturella , Physopella , Stereostratum and Puccinia.
Smuts: Entyloma, Tilletia, Sorosporium, Tolyposporium and
Ustilago.

Oryzodae

Mostly perennial, but a few annuals; ‘grasses’, or to varying
degrees ‘bambusoid’ in appearance. Culms to 1000 cm high
or scandent, woody or (mostly) herbaceous; branched or
(commonly) unbranched above. Leaves only occasionally with
auricular setae. Leaf blades linear to ovate; often not
pseudopetiolate;sometimes disarticulating, but more often
persistent. Inflorescence without pseudospikelets, determinate
except sometimes in Olyreae ; sometimes spatheate, more often
espatheate; of various forms. Female-fertile spikelets sometimes
with the rachilla prolonged, more often not so. Glumes usually
two and relatively large, but sometimes minute and not
infrequently absent. Often without incomplete florets; these when
present proximal, distal (or both), proximal incomplete florets
1-several. Female-fertile florets 1-17. Palea present, usually
relatively large; nerves 0, 1, 2 or several; with equal frequency
1— or 2-keeled, less often keel-less; entire. Lodicules usually
present, more often 2 than 3; rarely ciliate. Ovary nearly always
glabrous and unappendaged. Stigmas usually 2.

Abaxial leaf blade epidermis. Papillae present or absent with
about equal frequency, sometimes present on the stomatal
subsidiaries. Stomatal guard-cells more often flush-to-
overlapping the interstomatals than overlapped by them.
Mesophyll sometimes with arm cells and/or fusoids, but often
without either.

Chromosome base numbers mostly x = 10, 11 or 12; mostly
diploid. Mean diploid 2 c DNA value 1 .7-4.4 pg, group mean
3.05.

Rusts Physopella, Puccinia. Smuts Entyloma. Tilletia,
Sorosporium, Tolyposporium, Ustilago.

Oryzeae\ Leersia, Oryza, Prosphytochloa.

Culms herbaceous. Leaf blades without readily visible
transverse veins. Adaxial ligule an unfringed membrane;
sometimes very long. Inflorescence paniculate; espatheate; the
spikelet-bearing axes persistent. Spikelets all alike; laterally
compressed; disarticulating above the glumes, or above the

vestiges representing them; without an apically prolonged
rachilla. Glumes absent, or reduced to a 2-lobed cupule.
Incomplete florets present or absent; when present proximal only,
represented by a single sterile lemma exceeded by the female-
fertile one. Female-fertile florets 1. Lemma carinate, without a
germination flap. Palea relatively long, with several nerves.
Ovary apex glabrous, unappendaged. Stigmas white. Hilum long-
linear. Embryo small; with an epiblast; without a mesocotyl
internode; embryonic leaf margins overlapping.

Abaxial leaf blade epidermis. Microhairs present; panicoid-
type. Mid-intercostal long-cells rectangular, with markedly
sinuous walls. Papillae present. Costal silica bodies ‘oryzoid-
type’. Stomata with triangular subsidiaries. Costal short-cells
conspicuously in long rows. Transverse section of leaf blade,
physiology. C3; XyMS+. Mesophyll with or without arm-cells;
with or without fusoids.

Chromosome base number,.* =12. 2 n = 24-60.

Olyreae. Olyra. (See genus for description.)

Centotheceae: Megastachya. (See genus for description.)

Ehrharteae : Ehrharta, Microlaena.

Culms 15-200cm high, woody and persistent or herbaceous.
Leaf blades linear to linear-lanceolate; not pseudopetiolate;
without readily visible transverse veins. Adaxial ligule a fringed
or unfringed membrane, or a fringe of hairs. Infloresce
paniculate, or a single raceme; espatheate; with persistent axes.
Female-fertile spikelets compressed laterally to terete;
disarticulating above the glumes; with or without an apically
prolonged rachilla (Ehrharta sometimes exhibiting a minute
prolongation). Glumes minute (Microlaena) or relatively large.
Incomplete florets present; proximal only; 2; epaleate; sterile.
Female-fertile florets 1. Lemma decidedly firmer than the
glumes; carinate; without a germination flap; 5-7 nerved. Palea
present; variable in relative size; nerveless, with 1 nerve, or with
several; (0— ) 1 (—2) keeled. Stamens 2-6. Ovary glabrous. Stigmas
2. Caryopsis compressed laterally, not grooved. Hilum long-
linear. Embryo small.

Abaxial leaf blade epidermis. Microhairs present or absent;
when present panicoid-type. Costal silica-bodies "panicoid-type’.

Transverse section of the leaf blade, physiology. C3; XyMS-.
Arm-cells occasionally present in Ehrharta.

Chromosome base number, x = 10 or 12. 2 n = 24 or 48.

Bambusodae

Woody bamboos, with branching, robust culms. Leaves
pseudopetiolate, commonly with auricular setae; the blades
lanceolate to ovate, with or without readily visible transverse
veins, disarticulating from the sheaths. Inflorescence
indeterminate or determinate, with or without pseudospikelets;
structure variable, but usually spatheate; axes persistent.
Female-fertile spikelets usually large (group mean 30 mm long);
usually disarticulating above the glumes and (when applicable)
between the florets. Rachilla usually prolonged beyond the
uppermost female-fertile floret. Glumes occasionally 1, usually
2 or (not infrequently) several; usually relatively large, but
decidedly shorter than the adjacent lemmas; similar. Incomplete
florets usually present; occasionally proximal only, more often
distal or both proximal and distal. Proximal incomplete florets
when present 1-several. Female-fertile florets 1-30. Palea
present, relatively large; usually with several nerves; 2-keeled
or keel-less; notched or entire. Lodicules usually present;
occasionally 1 or 2, usually 3 or more; usually ciliate. Stamens
usually more than 3. Anthers sometimes with an apically
prolonged connective. Ovary glabrous or hairy; often with a
conspicuous apical appendage; styles usually joined, at least
below; stigmas 2 or (more often) 3 or more.

Abaxial leaf blade epidermis. Papillae nearly always present
and very conspicuous, usually over-arching the stomata; absent
from the subsidiaries. Stomatal guard-cells often overlapped by
the interstomatals. Transverse section of leaf blade. Lamina often
distinctly asymmetrical about the midrib. Mesophyll nearly
always with both arm-cells and fusoids.

Chromosome base number, x = 12 (very rarely 11); rarely
diploid, usually tetra- or hexaploid.

Rusts: Dasturella, Stereostratum, Puccinia. Smuts: Tilletia,
Ustilago (very few recorded).

Bambuseae. Bambusa, Oxytenanthera, Thamnocalamus.

384

Arundinoideae

Mostly perennial herbs, often caespitose with mainly basal
leaves, but occasionally more or less ‘bambusoid’ in habit. Culm
internodes solid or hollow. Leaf blades mostly linear or linear-
lanceolate, hardly ever pseudopetiolate, but not infrequently
disarticulating; hardly ever exhibiting conspicuous transverse
veins. Adaxial ligule present, sometimes an unfringed membrane
but more often a fringed membrane or a fringe of hairs. Abaxial
ligule sometimes present. Inflorescence determinate, without
pseudospikelets; occasionally of very few spikelets; usually
paniculate or reduced to a raceme, occasionally a spike or spicate;
espatheate; the axes persistent. Spikelets not in distinct long-and-
short combinations. Female-fertile spikelets usually compressed
laterally or terete, occasionally compressed dorsiventrally; nearly
always disarticulating above the glumes and (when applicable)
between the florets. Rachilla prolonged apically or not. Glumes
2, relatively large, very unequal or (more often) more or less
equal, carinate or not, nearly always similar. Lower glume
( 1— 3— 7(— 11) nerved; upper glume 1 — 7(— 11) nerved. Incomplete
florets present or absent, nearly always distal only, very
occasionally both distal and proximal (e.g. Phragmites).
Proximal incomplete florets when present 1 only. Female-fertile
florets 1-10. Lemmas sometimes entire, but usually incised and
often deeply cleft; sometimes muticous or mucronate, usually
awned. Awns 1 or 3, (the median) from a sinus or occasionally
apical, geniculate or non-geniculate (with a peculiar awn
configuration characterizing Aristideae: q.v.). Lemmas usually
hairy, the hairs sometimes conspicuously arranged in tufts and/
or transverse rows; occasionally carinate, more often rounded on
the back; ( 1— )3— 9(— 11) nerved. Palea present, nearly always 2
nerved and usually 2 keeled or keel-less; entire, notched or
occasionally deeply cleft. Lodicules usually present, usually 2
(occasionally 3, usually 3 in Stipeae); fleshy or membranous;
ciliate or glabrous. Stamens (l-)3. Ovary apex glabrous or hairy.
Styles occasionally joined below, usually free. Stigmas nearly
always 2; white, red-pigmented or brown. Fruit usually a
caryopsis, but occasionally with a free pericarp, the latter
sometimes thick and hard; longitudinally grooved or not. Hilum
long-linear to short. Embryo large or small, sometimes waisted.
Endosperm hard; without lipid; usually with compound starch
grains (Prionanthium exceptional). Embryonic leaf margins
usually meeting, the other embryo anatomical features variable.

Abaxial leaf blade epidermis. Microhairs present or absent
(but usually present somewhere on the plant); panicoid-type or
‘stipoid’ (in Stipeae) . Papillae nearly always absent. Costal
silica-bodies of various forms, but hardly ever of the ‘pooid’
(horizontally elongated-crenate or sinuous) type. Stomatal guard-
cells usually overlapping to flush with the interstomatals;
subsidiaries triangular, dome-shaped or occasionally parallel-
sided. Transverse section of leaf blade , physiology. C4 or (more
often) C3. XyMS+ or XyMS-. Where biochemically typed,
consistently NADP-ME (i.e. even where XyMS+). The blade
usually ribbed adaxially, the ribs rather frequently of different
size orders. Mesophyll without arm-cells (except Phragmites)-,
without fusoids; hardly ever traversed by colourless columns.
Midrib conspicuous or (more often) not readily distinguishable
from the other main veins; usually with one bundle only, rarely
a simple arc of bundles. Bulliforms present or absent; when
present usually simple fans, hardly ever associated with
colourless cells to for deeply-penetrating fans. Smallest vascular
bundles usually accompanied by sclerenchyma. Rather frequently
exhibiting sclerenchyma additional to that directly associated
with the vascular bundles (e.g. with a continuous abaxial
hypodermal layer).

Chromosome base numbers very variable (6, 7, 9, 11, 12, 13
etc.).

Helophytic, mesophytic or xerophytic; in open habitats.

Rusts: Dasturella, Puccinia. Smuts: Neovossia, Tilletia,
Urocystis, Sorosporium, Sphacelotheca, Tolyposporium,
Ustilago.

Stipeae: Nassella, Stipa.

Culms 10-250 cm high; nearly always herbaceous. Leaf
blades narrow; not pseudopetiolate; without readily visible
transverse veins. Adaxial ligule a fringed or unfringed membrane.
Abaxial ligule sometimes present. Inflorescence determinate;
paniculate; espatheate; the axes persistent. Female-fertile
spikelets compressed laterally; disarticulating above the glumes;

without an apically prolonged rachilla; with a hairy callus.
Glumes 2; more or less equal; about equalling or exceeding the
spikelets. Lower glume 1-4 nerved; upper glume 3-6 nerved.
Without incomplete florets. Female-fertile florets 1. Lemma
often convolute; decidedly firmer than the glumes, becoming
indurated; without a germination flap; conspicuously awned.
The single awn apical, from a small sinus or dorsally from near
the top; geniculate; entered by several veins. Palea well
developed to very reduced; keel-less. Lodicules present; 2 or 3;
membranous; glabrous. Stamens 3 (sometimes with penicillate
anthers). Ovary glabrous. Stigmas 2 or 3-4. Caryopsis not
grooved. Hilum long-linear. Embryo large or small.

Abaxial leaf blade epidermis. Microhairs absent (but a
peculiar form occasionally seen abaxially). Costal silica-bodies
variable (often ‘panicoid-type’, crescentic or rounded), but not
‘pooid-type’.

Transverse section of leaf blade, physiology. C3; XyMS+.
Mesophyll with neither arm-cells nor fusoids. All vascular
bundles accompanied by sclerenchyma.

Chromosome base number, x = 9-12 or 22. 2 n = 22-96.

Arundineae: Arundo, Phragmites.

Reeds. Culms 80-600 cm high; woody and persistent or
herbaceous; branched or unbranched above. Leaf blades 6-60
mm wide; lanceolate to linear-lanceolate; not pseudopetiolate;
without readily visible transverse veins; disarticulating from
the sheaths. Adaxial ligule a fringed membrane or a fringe of
hairs. Abaxial ligule absent. Inflorescence a large, plumose,
open panicle; the axes persistent. Female-fertile spikelets
compressed laterally; disarticulating above the glumes and
between the florets; with an apically prolonged rachilla. Glumes
similar, both 3-5 nerved. Incomplete florets present or absent; if
present distal or both distal and proximal. Female-fertile florets
2-10. Lemmas entire to incised but not deeply cleft; less firm
than the glumes to resembling them in texture; awned (with a
median, non-geniculate awn apically or from a sinus); hairless,
or hairy but lacking tufts and transverse rows of hairs; rounded
on the back. Palea 2-nerved. Lodicules 2; fleshy; ciliate or
glabrous. Stamens 3. Ovary glabrous. Hilum short. Embryo large.

Abaxial leaf blade epidermis. Microhairs present (panicoid-
type), or absent. Papillae absent. Costal short-cell arrangements
and silica-body forms variable. Transverse section of the leaf
blade, physiology. C3; XyMS+. Mesophyll tightly packed; with
(. Phragmites ) or without arm-cells; without fusoids.

Chromosome base number, x = 12. 2 n - 36-112.

Danthonieae: Centropodia. Chaetobromus, Cortaderia,

Dregeochloa. Elytrophorus, Karroochloa, Merxmuellera,
Pentameris, Pentaschistis, Prionanthium, Pseudopentameris,
Schismus. Styppeiochloa, Tribolium, Urochlaena.

Culms 2-300 cm high (mostly less than 250 cm); herbaceous;
usually caespitose; branching or unbranched above. Leaves
usually basally aggregated. Leaf blades 0.3-15 mm wide; usually
linear; without readily visible transverse veins; rarely
disarticulating. Adaxial ligule nearly always a fringe of hairs
or less often a fringed membrane, rarely an unfringed
membrane. Abaxial ligule sometimes present. Inflorescence
usually a panicle, occasionally a raceme, rarely a spike
(. Tribolium ) or falsely spicate with spikelet clusters
( Elytrophorus ); axes ending in spikelets; nearly always
persistent. Female-fertile spikelets 1-55 mm long; usually
disarticulating above the glumes and between the florets; with an
apically prolonged rachilla (except sometimes in Pentaschistis).
Glumes usually more or less equal; similar; markedly shorter than
to much exceeding the spikelets. Incomplete florets usually
present, distal (proximal incomplete florets absent). Female-
fertile florets 2-10 (1 in Poagrostis). Lemmas usually incised
and sometimes deeply cleft, occasionally entire; awnless,
mucronate or 1 (occasionally 3 or 5) -awned apically or (more
often) from the sinus. The (median) awn often geniculate.
Lemmas sometimes with conspicuous tufts and/or transverse
rows of hair's; only infrequently carinate; without a germination
flap. Palea usually well developed, usually apically notched but
occasionally entire or deeply bifid; 2-nerved and 2-keeled, the
keels sometimes winged. Lodicules 2; usually fleshy; ciliate or
glabrous. Ovary apex usually glabrous (hairy in Dregeochloa and
Pentameris). Hilum shape and embryo size variable.

Abaxial leaf blade epidermis very variable. Papillae absent.

Transverse section of the leaf blade, physiology. Nearly all C3
0 Centropodia C4); XyMS+. Mesophyll without arm cells; without
fusoids. Midrib usually inconspicuous, usually with one bundle
only. Chromosome base numbers, x = 6,1, 9, 12, 13.

385

Aristideae : Aristida, Sartidia, Stipagrostis.

Culms 15-200 cm high; herbaceous; caespitose. Leaf blades
mostly not disarticulating; linear . Adaxial ligule a fringed
membrane or a fringe of hairs. Inflorescence an espatheate
panicle with persistent axes. Female-fertile spikelets laterally
compressed or terete; disarticulating above the glumes; without
incomplete florets; without an apically prolonged rachilla.
Female-fertile floret 1. Lemma narrow, often convolute; with or
without a germination flap; awned apically. The awn of
characteristic form, with a basal column and trifid above (or
evidently a derivative of this). Palea relatively short to very
reduced; 0-2 nerved. Lodicules present or absent; membranous;
glabrous. Embryo without an epiblast; with an elongated
mesophyll internode intemode; the embryonic leaf margins
meeting.

Abaxial leaf blade epidermis. Microhairs present; panicoid
type. Papillae absent. Costal silica bodies variable in form; costal
short-cells variable in arrangement. Transverse section of leaf
blade, physiology. C4 ( Aristida , Stipagrostis) or C3 {Sartidia).
XyMS+ or XyMS-. Mesophyll with neither arm-cells nor fusoids.
Midrib conspicuous to inconspicuous, with one bundle only.

Chromosome base number, x = 11 or (occasionally) 12. 2 n =
22-66.

Chloridoideae

Culms 5-250(-300) cm high, nearly always herbaceous;
branching above, or unbranched. Young shoots nearly always
intravaginal. Leaves non-auriculate, without auricular setae.
Blades nearly always linear or linear-lanceolate ; not
pseudopetiolate; without readily visible transverse veins; hardly
ever disarticulating. Adaxial ligule nearly always a fringed
membrane or a fringe of hairs, very rarely an unfringed
membrane {Bewsia, Diandrochloa, Lintonia). Abaxial ligule very
rarely present. Inflorescence determinate; commonly of spikelike
main branches (sometimes digitate) or paniculate, but sometimes
a raceme, a spike, or falsely spicate with clusters of spikelets;
espatheate; axes usually persistent, occasionally
disarticulating. Spikelets often secund (then often biseriate), or
non-secund (occasionally distichous); hardly ever in distinct
long-and-short combinations. Female-fertile spikelets usually
compressed laterally or terete, but sometimes compressed
dorsiventrally ( Craspedorhachis , Diplachne, Microchloa,
Monelytrum etc.); falling with the glumes or (more commonly)
disarticulating above them; when applicable, disarticulating
between the florets or (often) not so. Rachilla prolonged beyond
the uppermost female-fertile floret or (less frequently) not.
Glumes present; 2; relatively large; equal or unequal; sometimes
awned; similar to very dissimilar. Lower glume (O-)l(-several)
nerved; upper glume 1 — 3(— 5 or more) nerved. With or without
incomplete florets, these when present usually distal,
occasionally both distal and proximal, very rarely proximal
only (then 1-several). Female-fertile florets 1-20. Lemmas
entire or incised and sometimes (then sometimes deeply cleft);
muticous, mucronate or awned; hairy (the hairs not conspicuously
in tufts and/or transverse rows) or hairless; carinate or not;
without a germination flap; nerves (l-)3(-ll). Awns when
present 1-several (then the median similar in form to the laterals);
(the median) apical or from a sinus, non-geniculate . Palea
nearly always 2-nerved and 2-keeled, the keels often winged;
entire or notched. Lodicules occasionally absent; when present
2, fleshy, nearly always glabrous. Stamens (1-3. Ovary apex
glabrous. Styles occasionally joined at the base, usually free.
Stigmas 2; white, red or brown. Fruit rarely longitudinally
grooved; a caryopsis or (quite frequently) the pericarp free or
loose. Hilum short. Endosperm hard, without lipid, usually but
not always containing compound starch grains. Embryo large;
usually with an epiblast; scutellar tail and elongated
mesocotyl internode present, embryonic leaf margins usually
meeting.

Abaxial leaf blade epidermis. Microhairs nearly always
present; occasionally panicoid-type (e.g. Eragrostis and allies),
sometimes ‘ Enneapogon type’, usually chloridoid-type. Mid-
intercostal long-cells rectangular, nearly always with markedly
sinuous walls. Papillae present or absent. Costal silica bodies
diverse, but mostly ‘panicoid-type’ or saddle-shaped (never
‘pooid-type’, hardly ever elongated-smooth). Stomatal guard-
cells nearly always flush with to overlapping the
interstomatals; subsidiaries triangular or dome-shaped,

hardly ever parallel-sided. Costal short-cells usually in long
rows, but occasionally predominantly paired, solitary or in short
rows. Transverse leaf blade section, physiology . C4, with the sole
known exception of Eragrostis walteri. XyMS +. Biochemical
types PCK and NAD-ME. Mesophyll often traversed by
colourless columns; rarely exhibiting arm-cells; without fusoids.
Blade very frequently adaxially flat, ribs when present usually
constant in size. Midrib conspicuous or not readily
distinguishable; usually with a single bundle, occasionally with
a simple arc; sometimes with colourless tissue. Bulliforms
usually present; commonly combined with colourless cells to
form deeply-penetrating fans, or comprising simple fans each
with a deeply-penetrating median cell. Smallest bundles
usually accompanied by sclerenchyma. Rarely exhibiting
sclerenchyma other than that directly associated with the bundles.

Chromosome base number, x - 10 (infrequently 7, 8, 9, 12).
Mean diploid 2 c DNA value 0. 7-1.4 pg, group mean 1.05;

Mostly mesophytic to xerophytic, occasionally helophytic. In
open habitats; rather frequently maritime or halophytic.

Rusts: Physopella, Puccinia. Smuts: Entyloma,

Melanotaenium, Tilletia, Sorosporium, Sphacelotheca ,

Tolyposporella , Ustilago.

Pappophoreae: Enneapogon, Kaokochloa, Schmidtia.

Culms 5-100 cm high; herbaceous. Nodes often hairy,
internodes hollow. Leaf blades linear to linear-lanceolate.
Adaxial ligule a fringe of hairs. Inflorescence an open or
contracted (sometimes very contracted) panicle; espatheate;
the axes persistent. Female-fertile spikelets disarticulating above
the glumes but not between the florets; the rachilla hairy,
prolonged apically. Glumes about equalling the spikelets;
similar; (5— )7— 1 1(— 21 ) nerved. Incomplete florets present, distal
only. Female-fertile florets 1-9. Lemmas firmer than the glumes;
incised into 4, 6 or 9 lobes; hairy; not carinate; without a
germination flap; 9 nerved; 2-3, 5 or 9 awned, the awns lateral
only or median and lateral, all similar and non-geniculate.
Palea well developed, 2 nerved and 2 keeled. Lodicules fleshy
or membranous, ciliate or glabrous. Ovary glabrous; stigmas
white. Pericarp fused. Hilum short. Embryo large; with epiblast,
scutellar tail and mesocotyl internode, the embryonic leaf
margins overlapping.

Abaxial leaf blade epidermis. Microhairs present;
Enneapogon type. Mid-intercostal long-cells rectangular.
Papillae absent. Costal short-cells in long rows, the costal silica
bodies panicoid-type. Transverse section of the leaf blade,
physiology. C4; XyMS+. Midrib of one bundle only. All the
vascular bundles accompanied by sclerenchyma.

Chromosome base number, x = 9, 10.

Chlorideae {including Cynodonteae, Eragrosteae, Sporoboleae,
Aeluropodeae, Lappagineae, Leptureae, Trageae, Spartineae):
Acrachne, Bewsia, Brachyachne, Brachychloa, Catalepis,
Chloris, Cladoraphis, Coelachyrum, Craspedorhachis, Ctenium,
Cynodon, Dactyloctenium, Diandrochloa, Dinebra, Diplachne,
Eleusine, Enleropogon, Entoplocamia, Eragrostis, Eustachys,
Fingerhuthia, Harpochloa, Leptocarydion, Leptochloa,
Lepturus, Lintonia, Lophachme, Microchloa, Monelytrum,
Mosdenia, Odyssea, Oropetium, Perotis, Pogonarthria,
Polevansia, Rendlia, Schoenefeldia, Spartina, Sporobolus,
Stiburus, Tetrachne, Tetrapogon, Tragus, Trichoneura,
Tripogon, Triraphis, Willkommia.

Culms ( 1— ) 1 0— 250(— 300) cm high; herbaceous; with glabrous
nodes. Internodes solid or hollow. Leaf blades usually linear or
linear-lanceolate. Adaxial ligule usually a fringed membrane
or a fringe of hairs, occasionally an unfringed membrane (e.g.
Bewsia, Diandrochloa). Inflorescence variously a single spike or
a raceme, of spikelike main branches (sometimes digitate),
falsely spicate with clusters of spikelets, or a panicle; espatheate.
Inflorescence axes persistent, less often disarticulating (then
disarticulating at the joints, or the reduced axes falling as clusters
of spikelets). Spikelets secund (often biseriate), or non-secund;
sessile, subsessile or pedicellate, but not in distinct long-and-
short combinations. Female-fertile spikelets usually compressed
laterally, (exceptions Diplachne, Enteropogon, Lepturus,
Microchloa, Monelytrum)', usually adaxial in forms with
discernable orientation; usually disarticulating above the glumes
(or between them), sometimes falling with them; with or without
an apically prolonged rachilla. The rachilla disarticulating
between the florets or (not uncommonly) persistent. Glumes
equal or unequal; similar to dissimilar; sometimes carinate;
sometimes awned. Lower glume (0— )1(— 3) nerved; upper
glume l-3(-12) nerved. Incomplete florets sometimes absent;
usually present, then usually distal only (both distal and proximal
in Ctenium, Entoplocamia). Female-fertile florets 1-45. Lemmas

386

rarely firmer than the glumes (not becoming indurated); entire or
variously incised; hairy or hairless, the hairs rarely in tufts but
not in transverse rows; carinate or not; without a germination
flap; 1 — 5(— 1 1 ) nerved; awnless, mucronate or awned. Lemma
awns 1, 3 or 5; non-geniculate. Palea usually relatively long;
entire or notched; usually 2-nerved and 2-keeled, the keels
sometimes winged. Lodicules usually present, fleshy, glabrous.
Anthers often very short. Ovary glabrous; stigmas 2, white, red
or brown. Pericarp sometimes free or loose. Hilum short.
Embryo usually large.

Abaxial leaf blade epidermis. Microhairs nearly always
present; sometimes panicoid-type (e.g. Eragrostis ), usually
chloridoid-type. Papillae often present. Mid-intercostal long-
cells rectangular. Costal short-cells usually in long rows, the
silica bodies saddle-shaped or panicoid-type. Transverse section
of leaf blade , physiology . C4 (except Eragrostis walteri) ; PCK or
NAD-ME; XyMS+. Midrib conspicuous to inconspicuous, with
one bundle or a simple arc. Usually with all the vascular bundles
accompanies by sclerenchyma.

Chromosome base number, x = usually 10 (occasionally 9,
rarely 7 or 12).

Panicoideae

Culms 10-400 cm high; mostly herbaceous but occasionally
woody and persistent; more often branching above than
unbranched. Culm intemodes more often solid than hollow.
Leaves generally not basally aggregated, usually non-auriculate;
without auricular setae. Blades mostly linear to ovate-lanceolate,
flat or rolled; occasionally pseudopetiolate; occasionally with
conspicuous transverse veins; rarely disarticulating. Adaxial
ligule an infringed membrane, a fringed membrane or a fringe of
hairs (all states common). Abaxial ligule occasionally present.
Inflorescence determinate; commonly paniculate, but almost
as often of spikelike main branches (sometimes digitate),
occasionally a spike, a raceme or falsely spicate with clusters of
spikelets; spatheate or not; the axes persistent or disarticulating.
Spikelets commonly in distinct long-and-short combinations.
Female-fertile spikelets most commonly compressed
dorsiventrally, less often compressed laterally or terete; nearly
always falling with the glumes, only occasionally disarticulating
above them; the rachilla not prolonged above the uppermost
female-fertile floret. Glumes usually 2, occasionally 1; usually
relatively large; equal to very unequal; frequently very
dissimilar; (the longer) usually long relative to the adjacent
lemma. Lower glume nerveless-1 1 nerved; upper glume
(0-) 1— 9(— 1 3) nerved. Incomplete florets nearly always present,
proximal; 1. Female-fertile florets nearly always 1, very
occasionally 2. Lemma with or without a germination flap;
usually non-carinate; nerveless-1 1 nerved. Palea nerveless or 2
nerved, keel-less or two-keeled, entire or notched. Lodicules
usually present, 2; fleshy; usually glabrous. Stamens (l-)3.
Ovary glabrous. Styles free or (often) joined at the base. Stigmas
2; usually red-pigmented, rarely white or brown. Fruit a
caryopsis; rarely longitudinally grooved; usually dorsiventrally
compressed. Hilum occasionally long-linear, usually short.
Endosperm hard, without lipid; starch grains sometimes
compound, but usually simple. Embryo usually large; without
an epiblast; scutellar tail and mesocotyl internode present;
embryonic leaf margins overlapping.

Abaxial leaf blade epidermis. Microhairs present; panicoid-
type. Mid-intercostal long-cells usually rectangular, occasionally
fusiform (occasionally without typical long-cells); the walls
usually but not always markedly sinuous. Papillae sometimes
present; in various configurations; occasionally present on the
stomatal subsidiaries. Costal silica bodies nearly always
‘panicoid-type’ (exception: Oxyrhachis ), occasionally sharp-
pointed. Stomatal guard-cells nearly always flush with to
overlapping the interstomatals; subsidiaries usually
triangular or dome-shaped, very rarely parallel-sided. Costal
short-cells nearly always in long rows, occasionally in other
configurations. Transverse section of the leaf blade, physiology .
C3 or C4. XyMS+ or XyMS-. C4 types PCK, NAD-ME or NADP-
ME. Blade commonly adaxially flat, the ribs when present of
equal size. Midrib inconspicuous or (usually) conspicuous;
sometimes with one bundle, usually with a simple arc; often with
adaxial colourless tissue. Mesophyll rarely traversed by
colourless columns; without arm-cells; very rarely with
fusoids. Bulliforms present or absent; infrequently associated

with colourless cells to constitute deeply-penetrating fans.
Commonly with the smallest bundles unaccompanied by
sclerenchyma. Very rarely having sclerenchyma not directly
associated with the bundles.

Chromosome base numbers x ~ mostly 5, 9 or 10. 2-18 ploid.
Mean diploid 2c DNA value 1.6-5. 2 pg, group mean 3.03.

Helophytic or mesophytic, less often hydrophytic or
xerophytic.

Rusts: Dasturella . Phakospora. Physopella. Puccinia. Smuts:
Entyloma. Melanotaenium, Tilletia, Sorosporium,
Sphacelotheca, Tolyposporella. Tolyposporium, Ustilago.

Panicodae

Leaf blades linear to ovate-lanceolate; occasionally with
conspicuous transverse veins. Adaxial ligule sometimes an
unfringed membrane, but more often a fringed membrane or a
fringe of hairs. Abaxial ligule occasionally present. Inflorescence
usually espatheate; occasionally with its branches naked-tipped
or terminating in bristles; the axes occasionally disarticulating
(then nearly always falling entire, frequently in the form of
condensed spikelet clusters), but usually persistent. Spikelets
often secund and biseriate when borne on spicate inflorescence
branches; sometimes associated with bristles (reduced
inflorescence branches or branch tips); occasionally in long-and
short combinations, but then the members usually alike in form
and sexuality. Female-fertile spikelets usually compressed
dorsiventrally, but not infrequently compressed laterally;
occasionally disarticulating above the glumes, but usually
disarticulating below them, occasionally shed in clusters. Glumes
very unequal or less frequently more or less equal; similar or
dissimilar in about equal frequency. Female-fertile lemma
usually at least as firm as the upper glume, frequently firmer
and becoming hardened in the fruit; nearly always with a
germination flap; infrequently awned and usually entire (save
in Arundinelleae); (0— )3— 1 1 nerved. Palea present, nearly always
well developed and relatively long; usually entire (except
Arundinelleae, Rhynchelytrum)', usually 2-nerved, often
2-keeled. Hilum occasionally long-linear.

Transverse section of the leaf blade. C3 or C4 (occasionally
genuinely intermediate). XyMS+ or XyMS-. C4 types PCK,
NAD-ME and NADP-ME. Mesophyll of C3 forms occasionally
Isachne-typc, occasionally traversed by colourless columns.

Basic chromosome number* = mostly 9, occasionally 10, 12
etc. Mean diploid 2c DNA value 1.6-2. 7 pg, group mean 2.3.

Paniceae: Acroceras, Alloteropsis, Anthephora, Axonopus,
Brachiaria, Cenchrus, Digitaria, Echinochloa, Entolasia,
Eriochloa. Leucophrys, Megaloprotachne, Melinis,
Odontelytrum, Oplismenus, Oryzidium, Panicum. Paratheria ,
Paspalidium. Paspalum, Pennisetum. Pseudechinolaena,
Sacciolepis, Setaria. Stenotaphrum, Stereochlaena, Tarigidia,
Tricholaena, Urochloa.

Culms 10-300(-800) cm high; mostly herbaceous; commonly
branching above. Intemodes often solid. Leaves usually not
basally aggregated. Leaf blades linear to lanceolate (rarely
ovate), 1-30 mm wide; occasionally cordate (e.g. Acroceras ), or
even sagittate (Cymbosetaria)', sometimes pseudopetiolate; rarely
with readily visible transverse veins; occasionally disarticulating.
Adaxial ligule sometimes an unfringed membrane, more often a
fringed membrane or a fringe of hairs. Abaxial ligule
occasionally present. Inflorescence commonly of spikelike main
branches (sometimes digitate or subdigitate) or a panicle,
occasionally falsely spicate with spikelet clusters, rarely a
raceme; very rarely spatheate; the axes sometimes naked-tipped
or terminating in a bristle. The spikelet-bearing inflorescence
axes persistent or disarticulating (then falling entire as spikelet
clusters). Female-fertile spikelets usually dorsiventrally
compressed, but terete or compressed laterally in Acroceras ,
Pseudechinolaena , Rhynchelytrum , Sacciolepis , Tricholaena ;
when orientation discernable, more often abaxial than adaxial;
usually disarticulating below the glumes; the rachilla not
prolonged apically. Glumes occasionally 1, usually 2 and
unequal; usually very dissimilar; hardly ever carinate. 1
incomplete floret present; proximal; paleate or epaleate; male
or sterile; the lemma less firm than to as firm as the female-fertile
one. Female-fertile floret 1. Lemma nearly always entire
(sometimes incised in Melinis, Rhynchelytrum ); occasionally
mucronate or with an apical, non-geniculate awn; usually
hairless; usually with a germination flap; rarely carinate;
mostly 3-7 nerved; usually firmer than the glumes, often
becoming indurated. Palea relatively long; usually entire;

387

usually similar in texture to the lemma (often indurated); 2

nerved. Lodicules usually present; fleshy; glabrous. Ovary
glabrous; styles sometimes joined at their bases; stigmas usually
red (occasionally white or brown). Fruit, embryo. Fruit usually
compressed dorsiventrally. Hilum usually short (long-linear in
Acroceras). Embryo large.

Abaxial leaf blade epidermis. Microhairs present; nearly
always panicoid-type. Mid-intercostal long-cells rectangular.
Papillae occasionally present. Costal short-cells usually in long
rows; costal silica-bodies usually panicoid-type. Transverse
section of leaf blade, physiology. C4 (including all three
biochemical types), or C3; XyMS-t- or XyMS-. C4 anatomical
organization 'conventional', except in Alloteropsis (q.v.).
Mesophyll without 'circular' (‘distictive’) cells; without arm
cells; without fusoids. Midrib usually more or less conspicuous,
often with an arc of bundles. Commonly exhibiting small
vascular bundles unaccompanied by sclerenchyma.

Chromosome base number, x = usually 9 (occasionally 7, 10,
11 12, 15 or 17).

Arundinelleae : Arundinella, Danthoniopsis, Loudetia,

T richopteryx, T ristachya .

Culms (2— )25— 300(— 500) cm high; herbaceous. Leaves rarely
basally aggregated. Leaf blades linear to lanceolate; not cordate,
not sagittate; without readily visible transverse veins; rarely
pseudopetiolate. Adaxial ligule a fringed membrane or a fringe
of hairs. Inflorescence an open or contracted panicle (rarely
reduced to a raceme); non-digitate; the axes persistent;
espatheate; terminated by spikelets. Spikelets solitary, paired or
in triplets. Female-fertile spikelets compressed laterally to
terete; 1.5-45 mm long; disarticulating above the glumes and
between the florets; usually with a hairy callus and without an
apically prolonged rachilla. Glumes usually very unequal; non-
carinate; dissimilar to similar; 3 or 5 nerved. Incomplete florets
present; proximal only; 1; male or sterile. Proximal lemma less
firm than the female-fertile one, or similar in texture. Female-
fertile floret 1. Lemma not becoming indurated; usually
2-toothed or lobed, sometimes deeply cleft; usually awned, the
awns 1 (median) or 3 (median and lateral), the (median) awn from
the sinus, geniculate; non-carinate; with or without a germination
flap; hairy (the hairs sometimes in tufts, occasionally in
transverse rows) or hairless; ( 1— )5— 7(— 9) nerved. Palea usually
notched; not indurated; 2 nerved; 2 keeled. Lodicules 2; fleshy;
glabrous. Ovary usually glabrous (hairy in Tristachya) ; stigmas
free to their bases. Fruit, embryo. Caryopsis often
longitudinally grooved. Hilum usually long-linear. Embryo
large.

Abaxial leaf blade epidermis. Microhairs present; panicoid-
type. Mid-intercosta; long-cells ususally rectangular. Papillae
absent. Costal short-cells usually in long rows, the costal silica-
bodies usually panicoid-type (sometimes round or crescentic in
Arundinella ). Transverse section of the leaf blade, physiology.
C4; XyMS- (except sometimes in Loudetiopsis). Mesophyll often
with ‘colourless’ (‘circular’, ‘distinctive’) cells; without arm-
cells; without fusoids. Midrib conspicuous to inconspicuous; wiih
one bundle, or an arc. Often with small vascular bundles
unaccompanied by sclerenchyma.

Chromosome base number, x = 10 or 12 (less often 6-7, 9 or
14).

Andropogonodae

Leaf blades linear to lanceolate; very rarely disarticulating.
Adaxial ligule rather more often an unfringed membrane than a
fringed membrane or a fringe of hairs. Abaxial ligules absent.
Inflorescence diverse, but not exhibiting naked branch-tips or
reduced-branch bristles; often comprising spicate 'racemes',
which may be variously (sometimes greatly) reduced; frequently
spatheate and comprising ‘partial inflorescences’ (i.e. with the
limits of the 'inflorescence' only arbitrarily definable); the axes
usually disarticulating. Spikelets usually paired (or in
triplets); usually in long-and-short combinations; frequently
heterogamous, the members of a combination predictably
different in sexuality (the short-pedicelled or sessile members
usually female or hermaphrodite, the longer-pedicelled
members male or sterile). Female-fertile spikelets usually
falling in combination with the adjoining member and their rachis
segment; usually dorsiventrally compressed. Glumes 2, usually
more or less equal; usually very dissimilar. The single
proximal lemma nearly always larger and more substantial
than the female-fertile one, which is frequently reduced and
hyaline, sometimes comprising a mere stipe. Female-fertile
lemma often (minutely) bifid, often with a geniculate awn from

the apex or the sinus; sometimes virtually reduced to the awn;
without a germination flap; not forming a hardened protection
for the fruit; rarely more than 3-nerved, often nerveless or
1-nerved. Palea commonly absent or vestigial, relatively short
or well developed; 2-nerved or (more often) nerveless; entire
or notched.

Transverse section of leaf blade, physiology. Seemingly
exclusively C4, XyMS- and NADP-ME type. Midrib usually
conspicuous; usually with an arc of bundles, usually with adaxial
colourless tissue. Blade sometimes ribbed adaxially, but more
often adaxially flat. Bulliforms and associated colourless cells
sometimes forming arches over the smaller vascular bundles (and
the adaxial epidermis sometimes extensively 'bulliform').
Usually with the smallest vascular bundles unaccompanied by
sclerenchyma.

Chromosome base number, x = mostly 5 or 10, occasionally
9, 12 etc.

Mostly helophytic or mesophytic.

Andropogoneae :

Culms 10-1200 cm high; mainly herbaceous. Leaf blades
occasionally pseudopetiolate; without conspicuous transverse
veins. Adaxial ligule a fringed or unfringed membrane, less
commonly a fringe of hairs. Plants usually bisexual, with
bisexual spikelets, and possessing hermaphrodite florets;
(Hypogynium monoecious with all the fertile spikelets unisexual),
occasionally with no hermaphrodite florets in Heteropogon also.
The spikelets usually in long-pedicel/short-pedicel (or
pedicellate/sessile) pairs or triplets, the members of each pair or
triplet commonly differing in sexuality (heterogamous); the
sessile or short-pedicelled members then usually female or
hermaphrodite and the pedicelled or longer-pedicelled members
usually male-only or sterile (but the situation reversed in
Trachypogon)’, or the spikelets all alike (homogamous:
Cleistachne , Eulalia, Imperata, Oxyrhachis, Saccharum etc.).
Inflorescence of spikelike main branches (‘racemes’) or
paniculate (the panicles then often readily interpretable as made
up of ‘partial inflorescences’ with reduced andropogonoid
‘racemes’); commonly spatheate. The spikelet-bearing axes
usually disarticulating at the joints, the fruiting spikelet then
falling with the adjoining rachis internode and non-fruiting
spikelet(s); but occasionally the axes persistent, and the spikelets
disarticulating individually beneath the glumes (Cleistachne,
Imperata, Sorghum spp., Trachypogon). Female-fertile spikelets
usually compressed dorsiventrally, sometimes laterally (e.g.
Arthraxon, Chrysopogon, Vetiveria); without an apically
prolonged rachilla; with a proximal incomplete floret (this
occasionally missing in Eulalia), and with one female-fertile
(hermaphrodite or female) floret above it. Glumes often very
dissimilar. The proximal floret paleate or more often epaleate,
usually sterile but sometimes male; its lemma usually larger and
more substantial than the (commonly reduced) female-fertile
lemma. Female-fertile lemma usually more or less reduced
(often to a stipe); entire or incised (commonly minutely so);
muticous, mucronate or awned (the single awn from the sinus or
apical; geniculate); without a germination flap; often nerveless
or 1 nerved, rarely with more than 3 nerves. Palea sometimes
relatively long, but more often more or less reduced, vestigial or
absent; 2 nerved or (more often) nerveless. Lodicules usually
present; fleshy; occasionally ciliate. Ovary glabrous; stigmas
usually red. Fruit, embryo. Hilum short. Embryo large.

Leaf blade epidermis. Microhairs nearly always present;
panicoid-type. Papillae often present. Costal short-cells usually
in long rows; costal silica-bodies usually panicoid-type.
Transverse section of the leaf blade, physiology. C4; type NADP-
ME; XyMS-. Mesophyll without arm-cells; without fusoids;
without ‘circular cells’.

Chromosome base number, x = mainly 5 or 10, less often 9
(rarely 7, 11, 12, 15, 17 or 20).

(i) Andropogoninae (‘awned Andropogoneae’): Andropogon,
Arthraxon, Bothriochloa, Chrysopogon, Cleistachne,
Cymbopogon, Dichanthium, Diheteropogon, Elymandra,
Eriochrysis, Eulalia, Heteropogon, Hyparrhenia, Hyperthelia,
Imperata, Ischaemum, Microstegium, Miscanthus,
Monocymbium, Schizachyrium, Sehima, Sorghastrum, Sorghum,
Thelepogon, Themeda, Trachypogon, Vetiveria.

Spikelet-bearing axes usually ‘racemes’ (only rarely
‘spikelike’), these sometimes much reduced; usually with slender
rachides; elaiosome usually absent; ‘articles’ usually hairy. Leaf
blade abaxial epidermis often papillate. Adaxial surface of the
leaf blade usually adaxially flat. Female-fertile lemma awned
(except Hypogynium and a few species in other genera);
commonly incised.

388

(ii) Rottboelliinae: Coelorhachis, Elionurus, Hackelochloa,
Hemarthria, Oxyrhachis, Phacelurus, Rhytachne, Rottboellia,
Urelytrum, Vossia.

Spikelet bearing axes ‘spikes’ with substantial rachides;
elaiosome usually present; ‘articles’ usually glabrous. Pedicel of
the pedicellate spikelet sometimes more or less fused to the
rachis. Female-fertile lemma awnless; entire. Papillae very rare
in the abaxial epidermis of the leaf blade. Leaf blade usually
adaxially ribbed or ‘nodular’ in section.

Maydeae : Coix.

Culms herbaceous, robust. Leaf blades 30-70 mm wide.

Plants monoecious, with all the fertile spikelets unisexual;
without hermaphrodite florets. The male- and female-fertile
spikelets in different inflorescences, on different branches of

the same inflorescence, or in different parts of the same
inflorescence branch. Female spikelet with a paleate or epaleate,
sterile proximal incomplete floret; with a single fertile lemma;
without an apically prolonged rachilla. Female-fertile lemma
awnless or mucronate; nerveless, 3 nerved or 4—5 nerved.
Lodicules absent. Fruit, embryo. Hilum short. Embryo large.
Epiblast absent; scutellar tail and mesocotyl internode present;
embryonic leaf margins overlapping.

Abaxial leaf blade epidermis. Microhairs present; panicoid
type or ‘balanoform’. Papillae absent. Costal silica-bodies
panicoid-type. Transverse section of the leaf blade, physiology.
C4; type NADP-ME; XyMS-. Mesophyll without arm-cells or
fusoids.

Chromosome base number, x = 5 or 9-10; 2 n = 10-108.

389

APPENDIX 2: DELTA

World grass genera - character list

The characters and their states recorded for each taxon are the
heart of the DELTA system. In DELTA, several character types
are used: unordered multistate characters, ordered multistate
characters, real numbers, integer numbers and text characters. A
character list is never complete but is constantly being
augmented, changed and refined as research progresses. The full
data for each genus in Watson's database of world grass genera
is recorded using the following character list, which demonstrates
the wide range of data from a number of disciplines that can be
used for taxonomic comparisons. The figure numbers in this
character list refer to the illustrations in Watson & Dallwitz
(1988), and the literature references are given in full as part of
the automated data set.

#1. <Synonyms: i.e. 'genera' included in the current description
— for most nomenclatural literature references, see
Clayton and Renvoize 1986>/

#2. <= Sensu lato genus: i.e. genus in which this taxon might
reasonably be (or sometimes is) included>/

Habit, vegetative morphology.

#3. cLongevity of plants>/

1. annual <or biennial, without remains of old sheaths or

culms>/

2. perennial <with remains of old sheaths and/or culms>

<Figs 1, 2, 18>/

#4. <Reeds>/

1. reeds <helophytic, tall, to (2-)3 m or more in height;

culms woody and persistent, always leafy>/

2. not reeds <imp!icit>/

#5. <Habit>/

1. long-rhizomatous/

2. long-stoloniferous/

3. caespitose <Figs 1, 7>/

4. decumbent <including ‘rooting at the nodes’> <Fig.

2>/

#6. The flowering culms <whether having foliage leaves>
<intended mainly for bamboos>/

1 . leafless/

2. leafy/

#7. <Mature> culms cmaximum height: data unreliable for
large genera>/
cm high/

#8. Culms <whether woody or herbaceous>/

1. woody and persistent/

2. herbaceous <not woody, not persistent>/

#9. Culms reaching <maximum diameter: note cm units,
intended for bamboos>/
cm in diameter/

#10. Culms <shape: intended for bamboos>/

1 . cylindrical/

2. flattened on one side/

#11. Culms <whether scandent>/

1. scandent/

2. not scandent <self-supporting, scrambling or

floating> <implicit>/

#12. Culms <whether branched above>/

1. branched <vegetatively> above <Fig. 2>/

2. unbranched <vegetatively> above <Figs 1, 7>/

#13. <Number of> primary branches per mid-culm node

<intended mainly for bamboos>/

#14. Culms <whether tuberous at base>/

1. tuberous <at base> <Fig. 3>/

2. not tuberous <at base — implicit;*/

#15. <Culm> nodes <whether hairy or glabrous>/

1 . hairy <Figs 4, 33>/

2. glabrous <Fig. 4 >/

#16. <Culm> nodes <number of ridges: bamboos>/
ridged/

#17. Culm sheaths persistence (intended mainly for
bamboos)>/

1. <or at least their bases> persistent/

2. deciduous in their entirety/

#18. <Mid> culm internodes <whether solid or hollow: avoid
the 'peduncle'>/

1 . solid <or spongy>/

2. <conspicuously> hollow/

#19. <Bambusoid habit, unicaespitose or pluricaespitose
(intended for bamboos)>/

1. unicaespitose/

2. pluricaespitose/

#20. Rhizomes <form (intended mainly for bamboos)>/

1. pachymorph <sympodial>/

2. leptomorph <monopodial>/

#2 1 . Plants <whether conspicuously armed>/

1. conspicuously armed <specify how>/

2. unarmed/

#22. Young <vegetative> shoots <whether extra- or
intravaginal: poorly recorded>/

1. extravaginal cbursting through the bases of

subtending sheaths> <Fig. 6>/

2. intravaginal <emerging from between subtending

sheath and stem> <Fig. 5>/

#23. The <fresh> shoots <whether aromatio/

1. aromatic <when crushed>/

2. not aromatic <when crushed — implicit:*/

#24. Leaves <whether mainly basal, or mainly on the culms>/

1 . mostly basal <Figs 7, I4>/

2. not <distinctly> basally aggregated <i.e., the culms

leafy> <Figs 1, 2, 9, 16, 33>/

#25. Leaves <whether differentiated into sheath and blade>/

1. clearly differentiated into sheath and blade

<implicit>/

2. not clearly differentiated into sheath and blade/

#26. Leaves <phyllotaxy>/

1. spirally disposed <Figs 8, 9>/

2. distichous <the near-universal condition — implicit;*/
#27. Leaves <whether auricles present or absent: see Clifford

and Watson 1977, for definition:*/

1 . auriculate <Fig. 10>/

2. without auricles <Figs 12, 19 etc.>/

#28. Leaves presence of auricular setae (data poor, except for
bamboos)>/

1. with auricular <‘oral’> setae <Fig. 1 1 >/

2. without auricular <‘oral’> setae <Fig. 12 etc.>/

#29. <Leaf> sheath margins <whether joined>/

1. joined <to at least one-quarter of their length: 'sheaths

tubular’;*/

2. free <implicit>/

#30. <Comments on sheaths>/

#31. Leaf blades <extreme reduction>/

1. <all> greatly reduced <with main functions

transferred elsewhere>/

2. not all greatly reduced <implicit>/

#32. Leaf blades <shape: data incomplete>/

1 . linear/

2. linear-lanceolate/

3. lanceolate/

4. ovate-lanceolate/

5. ovate/

6. elliptic <oblong>/

7. obovate/

#33. Leaf blades <texture>/

1. leathery/

2. flimsy/

3. neither leathery nor flimsy <to become implicit/

#34. Leaf blades <whether broad or narrow (specify the true

range)>/

1. broad <maximum (flattened) width greater than 1

cm>/

2. narrow cmaximum (flattened) width less than 1 cm>/
#35. Leaf blades cmid-width: data very incomplete>/

mm wide <in the middle>/

390

#36. Leaf blades <whether cordate or sagittate>/

1. <at least some of them> cordate <Fig. 13>/

2. <at least some of them> sagittate/

3. not cordate, not sagittate <implicit>/

#37. Leaf blades <whether setaceous>/

1 . setaceous <i.e., fine and bristle-like: not to be

confused with pungent, subulate etc.> <Fig. 14>/

2. not setaceous <implicit>/

#38. Leaf blades <folded/rolled>/

1 . flat/

2. folded <Fig. 42>/

3. rolled/

4. acicular/

#39. Leaf blades <whether needle-like>/

1. hard, woody, needle-like <and plants prickly, e.g.

Triodia> <Fig. 15>/

2. not needle-like <plants not prickly> <implicit>/

#40. Leaf blades <whether pseudopetiolate>/

1. pseudopetiolate <Figs 11, 42 >/

2. not pseudopetiolate <implicit>/

#41. Leaf venation <layout>/

1. pinnate <Fig. 16 >/

2. palmate/

3. neither pinnate nor palmate <implicit>/

#42. Leaf blades <whether with conspicuous transverse veins>/

1 . with readily visible transverse veins <at least

abaxially> <Fig. 17>/

2. without readily visible transverse veins/

#43. Leaf blades cwhether disarticulating>/

1. cor at least many of them, ultimately> disarticulating

from the sheaths <Fig. 9>/

2. not disarticulating/

#44. Leaf blades cwhether vernation rolled or folded>/

1. rolled in bud/

2. once-folded in bud/

3. folded like a fan in bud/

#45. <Adaxial> ligule <presence>/

1 . <consistently> present <implicit>/

2. absent, at least from upper leaves/

#46. <Adaxial> ligule cform — avoid seedlings>/

1 . an unfringed membrane cmay be variously hairy or

ciliolate> <Fig. 19>/

2. a fringed membrane cFigs 20, 21, 23>/

3. a fringe of hairs <Fig. 22 >/

4. a rim of minute papillae/

#47. <Adaxial> ligule cshape of apex>/

1 . truncate/

2. not truncate cacute, obtuse or rounded> <Fig. 19 >/
#48. <Adaxial> ligule clength at middle: generally recorded

only for membranous, unfringed forms>/
mm long/

#49. <Outer> contra-ligule cpresence: data very incomplete;*/

1 . present cFigs 11, 24>/

2. absent <Fig. 12 etc.>/

Reproductive organization.

#50. Plants cwhether plants monoecious, with bisexual
spikelets, or dioecious>/

1. cbisexual, but> monoecious with all the fertile

spikelets unisexual/

2. bisexual, with cat least some> bisexual spikelets

cPlates 1:4, 1:5, 1:8, 2:10, 2:11, 2:13-2:17 etc.>/

3. dioecious cwith separate male and female-fertile

individuals> cFigs 25, 26>/

#51. Plants cwhether having hermaphrodite florets: not to be

confused with presence or absence of hermaphrodite
spikelets>/

1. with cat least some> hermaphrodite florets cPlate

2:14 etc.>/

2. without hermaphrodite florets/

#52. The spikelets cwhether heterospiculate: exclusive of
‘hidden’ spikelets>/

1. of cat least two> sexually distinct forms on the same

plant ce.g., female or hermaphrodite and sterile or
male-only. Vestigial spikelets represented only by
their pedicels have here been regarded as spikelets>
cPlates 1:3, 1:6; Figs 27-29, 33, 75, 76>/

2. all alike in sexuality con the same plant: ignore

hidden axillary spikelets, etc. Implicit:*/

#53. The male and female-fertile spikelets cdisposition on the
plant>/

1. in different inflorescences/

2. on different cmain> branches of the same

inflorescence/

3. segregated, in different parts of the same

inflorescence branch cPlate 1:6>/

4. mixed in the inflorescence/

#54. The spikelets cwhether heteromorphic (intended mainly
for heterospiculate andropogonoids)>/

1. overtly heteromorphic cPlate 1:6; Figs 27, 28, 75,

76 >/

2. cexternally> homomorphic/

#55. The spikelets cwhether the spikelet combinations are all
heterogamous: generally applied only to
andropogonoids>/

1. in both homogamous and heterogamous combinations,

2. all in heterogamous combinations/

#56. Plants cwhether outbreeding or inbreeding — data
extensively from Connor 1979>/

1. outbreeding callogamous>/

2. inbreeding cautogamous>/

#57. cCleistogamy — data mainly from Connor 1979: exposed
spikelets>/

1. exposed-cleistogamous cassociated with varying

degrees of spikelet and/or floret modification;*/

2. chasmogamous cunreliably implicit;*/

#58. Plants cpossession of hidden, cleistogamous spikelets in
leaf axils or on specialised rhizomes;*/

1. with hidden cleistogenes cmore or less hidden,

usually conspicuously modified cleistogamous
spikelets>/

2. without hidden cleistogenes cimplicit>/

#59. The hidden cleistogenes clocation>/

1. in the leaf sheaths/

2. subterranean crhizanthogenes>/

#60. cWhether apomixis observed: data mainly from Connor
1979>/

1 . apomictic/

2. reproducing sexually cunreliably implicit:*/

#61. cOccurrence of vivipary (poorly recorded);*/

1 . viviparous/

2. not viviparous cunreliably implicit:*/

Inflorescence.

#62. Inflorescence cwhether determinate (semelauctant) or
indeterminate (iterauctant or with a seemingly
indeterminate synflorescence): see McClure 1973,
Calderon and Soderstrom 1973 etc. for definitions:*/

1. determinate csemelauctant — implicit;*/

2. indeterminate citerauctant> cFig. 4 1 >/

#63. Inflorescence cwhether possessing pseudospikelets: see
McClure 1973 for definition^

1. with pseudospikelets chaving basal bracts with

axillary spikelets, in addition to or instead of the
usual barren glumes> cFig. 4 1 >/

2. without pseudospikelets cimplicit>/

#64. Inflorescence creduction>/

1 . reduced to a single spikelet/

2. of only two or three spikelets/

3. normally of more than three spikelets cimplicit>/

#65. Inflorescence cchasmogamous: overall form>/

1. a single spike cPlate 1:9; Fig. 30>/

2. of spike-like main branches cof spikes, narrow

racemes or narrow panicles;* cPlate 1 :4; Figs 3 1 , 38,
53, 54>/

3. a false spike, with clusters of spikelets on reduced

axes cPlate 1:5; Figs 42, 49, 50, 51, 77>/

4. a single raceme cat least some of the spikelets clearly

pedicellate> cFig. 33, 79>/

5. paniculate cand not readily referable to any of the

other states> cPlates 1:1, 1:3, 1:8, 2: 12; Figs 34, 35,
36, 37, 55>/

#66. Inflorescence ctumbleweeds>/

1. deciduous in its entirety as a ‘tumbleweed’ cFig. 26 >/

2. not deciduous cimplicit>/

#67. Inflorescence coverall form: mainly applied to panicles>/

1. open cPlates 1:8, 2:12; Figs 34, 35>/

2. contracted every compact, or narrow and spike-like>

cPlates 1:1, 1:3; Figs 36, 37>/

#68. Inflorescence ccompact, solitary: form — mainly applied
to panicles and solitary racemes;*/

1. capitate cmore or less spherical:* cPlate 1:3>/

2. more or less ovoid/

3. elongated-symmetrical, spike-like cFig. 37>/

4. more or less irregular cneither capitate nor ovoid, not

elongated-symmetrical:*/

#69. Inflorescence cwhether branches divaricate>/

1. with conspicuously divaricate branchlets cFig. 40>/

391

2. without conspicuously divaricate branchlets
<implicit>/

#70. Inflorescence <whether branchlets capillary: avoid

INTKEY use with non-paniculate inflorescences,
which have usually been scored as ‘inapplicable’>/

1. with capillary branchlets <Fig. 35>/

2. without capillary branchlets <implicit>/

#71. Inflorescence cwhether digitate or subdigitate;-/

1. digitate cincludes paired branches> <Figs 38, 54>/

2. subdigitate/

3. non-digitate <neither digitate nor ‘subdigitate’ —

implicit;-/

#72. With <number of primary inflorescence branches: applied
mainly to forms with spike-like main branches —
data very incomplete>/
primary inflorescence branches/

#73. Inflorescence <whether branches end in spikelets>/

1. with axes ending in spikelets <Figs 30, 32, 54>/

2. axes <often> not ending in spikelets <Figs 26, 31,

49>/

#74. Rachides <whether clearly (macroscopically) flattened,

hollowed or winged (states poorly defined, often left
uncoded)>/

1 . hollowed <Plates 1 :7, 1 :9; Figs 30, 56 >/

2. flattened <Fig. 39>/

3. winged/

4. neither flattened nor hollowed, not winged/

#75. Spikelets <whether embedded in the rachis>/

1. all <more or less> partially embedded in the rachis

<Plates 1:7, 1:9; Figs 30, 42, 56>/

2. not all embedded <implicit>/

#76. Inflorescence <whether spatheate (note: ‘spatheate’ not
currently distinguished from ‘spatheolate’)>/

1. spatheate <specify> <ignore mere early enclosure by

an unmodified flag leaf> <Figs 25, 26, 27, 43>/

2. espatheate/

#77. Inflorescence <whether comprising a complex of ‘partial
inflorescences' and intervening foliar organs ( =
leaves, spathes, spatheoles>/

1. a complex of ‘partial inflorescences’ and intervening

foliar organs <i.e., a ‘pseudo-inflorescence’> <Fig.
43>/

2. not comprising ‘partial inflorescences’ and foliar

organs/

#78. <Ultimate> spikelet-bearing axes <form> <intended
mainly for andropogonoids and bamboos>/

1. very much reduced <specify> <Plate 1:5; Figs 26, 42,

49, 50, 5 1 >/

2. spikes/

3. ‘racemes’ <Plate 1:6; Fig. 43 >/

4. spike-like <cf. Hemarthria> <Plate 1:7; Figs 27, 46,

47, 59, 60, 75, 76>/

5. paniculate/

6. capitate <= 1&5>/

#79. The spikelet-bearing axes <andropogonoid, number of
spikelet-bearing ‘articles’ (joints)>/

1. with only one spikelet-bearing ‘article’/

2. with 2-3 spikelet-bearing ‘articles’/

3. with 4—5 spikelet-bearing ‘articles’/

4. with 6-10 spikelet-bearing ‘articles’/

5. with more than 10 spikelet-bearing ‘articles’ <specify

the approximate number>/

#80. The racemes <whether spikelet bearing to the base>/

1. spikelet bearing to the base/

2. without spikelets towards the base/

#81. <Ultimate> spikelet-bearing axes <grouping> <intended
mainly for andropogonoids>/

1 . solitary/

2. paired/

3. clustered <in groups of three or mor e>/

#82. <Ultimate> spikelet-bearing axes <thickness of rachides>
cintended mainly for andropogonoids>/

1. with very slender rachides cPlate 1:6>/

2. with substantial rachides <Plate 1:7; Fig. 27>/

#83. Spikelet-bearing axes <whether disarticulating. Note that

spikelet-bearing axes may be greatly reduced>/

1. disarticulating <often manifested in clearly articulated

rachides. Excluding inflorescences falling whole
(tumbleweeds)> <Figs 26, 27, 44, 45, 46, 47, 59,
76>/

2. persistent <not disarticulating: implicit> <Figs 30, 38,

39, 53 >/

#84. Spikelet-bearing axes <manner of disarticulation>/

1. falling entire <Figs. 50, 5 1 >/

2. disarticulating at the joints <Figs 27, 44, 45, 46, 47,

59, 75, 76>/

#85. The pedicels and internodes of the rachis <Bothriochloa ,
Dichanthium and relatives>/

1. with a longitudinal, translucent furrow/

2. without a longitudinal, translucent furrow <implicit>/
#86. ‘Articles’ <( 'joints’ ) of the spikelet-bearing rachis, shape

(intended mainly for andropogonoids )>/

1. linear/

2. non-linear <Figs 44, 46, 47, 59, 75>/

#87. ‘Articles’ <of the spikelet-bearing rachis: whether bearing
an elaiosome>/

1. with a basal callus-knob <elaiosome> <Fig. 47>/

2. without a basal callus-knob/

#88. ‘Articles’ <of the spikelet-bearing rachis, whether

appendaged (intended mainly for andropogonoids)>/

1. appendaged <Figs 45, 48>/

2. not appendaged <Plate 1:7; Fig. 46>/

#89. ‘Articles’ <of the spikelet-bearing rachis, orientation of
disarticulation (intended mainly for
andropogonoids);-/

1. disarticulating transversely <Plate 1:7; Figs 44, 46,

47, 59, 75>/

2. disarticulating obliquely <Figs 27, 28>/

#90. ‘Articles’ <of the spikelet-bearing rachis, whether hairy
(intended mainly for andropogonoids>/

1. densely long-hairy/

2. somewhat hairy/

3. glabrous <Plate 1:7; Figs 46, 47, 59, 75>/

#91. Spikelets <and/or clusters, whether subtended by or

associated with ‘involucres’ or bristles representing
vestigial branches (note that ‘bristles’ must not be
confused with hairs)>/

1. <all> unaccompanied by bractiform involucres, not

associated with setiform vestigial branches
<implicit>/

2. (at least some of them) subtended by solitary ‘bristles’

<vestigial branches>/

3. <or clusters> with ‘involucres’ of ‘bristles’ cvestigial

branches;- <ignore true hairs> <Figs 49, 50, 5 1 >/

4. associated with bractiform involucres <Fig. 72>/

#92. The creduced branch> ‘bristles’ <form, coalescence>/

1. spiny, markedly coalescent basally <Fig. 50>/

2. relatively slender, not spiny <Figs 49, 5 1 >/

#93. The creduced branch;- ‘bristles’ cwhether deciduous>/

1. persisting on the axis <Fig. 49>/

2. deciduous with the spikelets cFigs 50, 5 1 >/

#94. The involucres cwhether deciduous or persistent;-/

1 . persistent on the rachis/

2. shed with the fertile spikelets/

#95. Spikelets cgrouping: recorded mainly in spikes and
racemes>/

1. cmainly> solitary cPlate 1:4; Figs 30, 38, 56>/

2. cconsistently> in pairs cPlate 1:6; Fig. 44>/

3. cconsistently> in triplets cFigs 46, 52, 59>/

#96. Spikelets cwhether secund: currently a catch-all character,
covering one-sidedness of inflorescence (e.g.,
Dactylis, dorsiventral rachides, etc .>/

1 . secund cPlates 1:2, 1 :4, 2: 15; Figs 3 1 , 32, 38, 39, 42,

53, 54, 55>/

2. not secund/

#97. Spikelets cinsertion>/

1. biseriate con one side of the rachis> cPlate 1:4; Figs

31, 38, 39, 53, 54>/

2. distichous cFig. 30 >/

3. not two-ranked cnot biseriate, not distichously

arranged> cto become implicit>/

#98. Spikelets cinsertion — revised version;-/

1. sessile cPlate 1:9; Figs 30, 56 >/

2. subsessile cFig. 54>/

3. pedicellate cPlates 1:1, 1:8, 1:5, 2:10-12 etc.; Figs

34, 35, 53, 63, etc.>/

#99. Pedicel apices cshape — recorded as yet only in Paniceae.
Data mainly from R.D. Webster 1985 >/

1. oblique cFig. 58 >/

2. truncate cFig. 58 >/

3. discoid cPlate 2:12; Fig. 58 >/

4. cupuliform cPlates 1:8, 2:10; Fig. 57>/

#100. Spikelets cdisposition, e.g. Diplachne/Leptochloa: not
widely recorded>/

1. imbricate/

2. distant cnot overlapping>/

#101. Spikelets cwhether in regular ‘long-and-short’
combinations, as exemplified in typical
andropogonoids>/

1. consistently in ‘long-and-short’ combinations ci.e.,
pedicellate/sessile or long-pedicel/short-pedicel pairs

392

or triplets: currently includes andropogonoid forms
with the pedicellate ‘spikelets’ reduced to their
pedicels> <Plates 1:6, 1:7; Figs 28, 44, 47, 59, 72,
76>/

2. not <consistently> in distinct iong-and-short’
combinations <implicit>/

#102. Spikelets <detail of ‘long-and-short’ combinations
(intended mainly for andropogonoids)>/

1. in pedicellate/sessile combinations <Figs 28, 44, 47,

59 >/

2. unequally pedicellate in each combination/

#103. Pedicels of the ‘pedicellate’ spikelets <whether fused

with the rachis: intended for andropogonoids>/

1. discernible, but <extensively> fused with the rachis

<Plate 1:7; Figs 47, 60>/

2. free of the rachis <Fig. 44, 75>/

#104. The ‘shorter’ <andropogonoid> spikelets <sessile or
shorter-pedicelled, sexuality>/

1. hermaphrodite <Fig. 59>/

2. female-only/

3. male-only/

4. sterile/

#105. The ‘longer’ <andropogonoid> spikelets <pedicelled or
longer-pedicelled, sexuality>/

1 . hermaphrodite/

2. female-only/

3. male-only/

4. sterile <comment if reduced to pedicels> <Figs 59,

60 >/

Female-sterile spikelets.

#106. <Description of female-sterile spikelets>/

Female-fertile spikelets.

#107. <Female-fertile> spikelets <whether morphologically
conventional/

1. morphologically ‘conventional’ <with readily

identifiable glumes, lemmas and paleas> <implicit>/

2. <very> unconventional <and hard to interpret;-/

#108. <Female-fertile> spikelets <approximate length,

excluding any awns: data unreliable for large
genera>/
mm long/

#109. <Female-fertile> spikelets <orientation of sessile to
subsessile forms>/

1. abaxial <G1 when present on the side away from the

rachis; in panicoid forms having a proximal
incomplete floret, the upper (female-fertile) lemma
backs onto the rachis> <Plate 1:7; Figs 59, 70, 79>/

2. adaxial <G1 when present against the rachis; in

panicoid forms having a proximal incomplete floret,
the upper (female-fertile) lemma is on the side away
from the rachis> <Plate 1:4>/

#110. <Female-fertile> spikelets <plane of compression^

1. compressed laterally <ly ing on the side when placed

on a flat surface> <Plates 1:2, 1:5, 1:9, 2:15, 2:17;
Figs 30, 54, 56, 61, 67, 68, 73, 1 19>/

2. not noticeably compressed <terete>/

3. compressed <dorsally, ventrally or> dorsiventrally

<lying on front or back when placed on a flat
surface> <Plates 1:4, 1:7, 2:10, 2:1 1; Figs 70, 74, 75,
82, 97, 98>/

#111. <Female-fertile> spikelets <shape of ‘dorsiventrally
flattened’ forms>/

1. planoconvex/

2. biconvex/

#112. <Female-fertile> spikelets <location of disarticulation
positions>/

1. <readily> disarticulating above the glumes <when

mature>/

2. falling with the glumes <when mature> <pending data

changes, including forms where the spikelets are
shed by inflorescence disarticulation>/

3. not disarticulating <common in cultivated cereals>/

#1 13. <Female-fertile> spikelets <whether rachilla

disarticulates between the florets of spikelets with
two or more fertile florets>/

1 . not disarticulating between the florets/

2. disarticulating between the florets/

#114. <Female-fertile> spikelets <rachilla internode spacings:
unsatisfactorily defined, and inadequately scored for
treating state 1 as implicit>/

1. with conventional intemode spacings/

2. with a distinctly elongated rachilla internode between

the glumes <Fig. 62>/

3. with a distinctly elongated rachilla intemode above

the glumes <Figs 63, 64, 8 1 >/

4. with distinctly elongated rachilla internodes between

the florets/

#1 15. <Presence or absence of Ichnanthus -type stipe:
Paniceae>/

1. the upper floret conspicuously stipitate <Fig. 64>/

2. the upper floret not stipitate/

#1 16. The stipe beneath the upper floret cthickness: Ichnanthus
relatives>/

1 . filiform/

2. not filiform <Fig. 64>/

#117. The stipe beneath the upper floret <shape: Ichnanthus
relatives;-/

1. straight and swollen <Fig. 64>/

2. curved, not swollen/

#118. The stipe beneath the upper floret <whether
heterogeneous Zuloaga 1987>/

1. heterogeneous cmembranous towards the base of the

palea. indurated on the lemma side>/

2. homogeneous/

#119. Rachilla <of female-fertile spikelets, whether terminated
by a female-fertile floret, or ‘prolonged’;-/

1. prolonged beyond the uppermost female-fertile floret

<i.e. not terminated by a female-fertile floret: note
that ‘racemose’ spikelets with three or more
female-fertile florets have all been awarded this
stato <Figs 41, 56, 61, 65>/

2. terminated by a female-fertile floret <not

‘prolonged’;-/

#120. Rachilla <of female-fertile spikelets, whether hairy>/

1. hairy <between the female-fertile florets, or above the

single one>/

2. hairless/

#121. The rachilla extension <beyond the uppermost

female-fertile floret of female-fertile spikelets,
rudiments;-/

1. with incomplete florets/

2. naked/

#122. Callus <presence/length: data very incomplete;-/

1. absent/

2. short/

3. long <Fig. 100>/

#123. Callus <whether blunt or pointed>/

1. pointed <Figs 28, 100>/

2. blunt/

#124. Hairy callus <presence: an unsatisfactory catch-all

character, but widely recorded and useful in keys>/

1. present <Figs 28, 63, 72, 100>/

2. absent/

#125. Callus hairs <presence, size: CalamagrostislAgrostis>l

1. present, more than 0.5 mm long/

2. absent, or if present less than 0.5 mm long/

#126. Glumes <of female-fertile spikelets, present or absent>/

1. present <implicit>/

2. absent/

#127. Glumes <of female-fertile spikelets, number: ‘glumes’
are barren, with neither axillary spikelets nor
florets;-/

1. one per spikelet/

2. two/

3. several/

#128. Glumes <whether glumes of the female-fertile spikelets
are all minute>/

1. minute <relative to the rest of the spikelet> <Plate

1:2; Fig. 63>/

2. relatively large <implicit>/

#129. Glumes <of female-fertile spikelets, whether markedly
unequal in the intact spikelet; regardless of any.
differences in form>/

1. very unequal <in length in the intact spikelet> <Plates

1:8, 2:10, 2:12; Figs 61, 68, 71>/

2. more or less equal <in length in the intact spikelet>

<Plates 1:1, 1:7,2:11; Figs 62, 66, 73, 79, 85, 89,

1 14, 1 19>/

#130. Glumes <length relative to the spikelet — applied only to
spikelets with 2 or more florets> currently for
key-making only>/

1 . markedly shorter than the spikelets <Figs 61, 68 >/

2. about equalling the spikelets <Plate 1:8; Figs 62, 93 >/

3. much exceeding the spikelets <Figs 66, 67, 73, 85,

1 14>/

#131. Glumes <of female-fertile spikelets, lengths relative to
proximal (adjacent) lemmas. Refers to the longer

393

glume when glumes unequal>/

1. decidedly shorter than the adjacent lemmas <in intact

spikelets> <Figs 61, 68, 89 >/

2. long relative to the adjacent lemmas cmore or less

equalling or exceeding them> <Plates 1:1, 1:7, 1:8,
2:15, 2:16; Figs 62, 66, 67, 71, 73, 79, 85, 114,

1 19>/

#132. Glumes <of female-fertile spikelets, whether free or
joined>/

1. joined <at least basally>/

2. free <implicit>/

#133. Glumes <of female-fertile spikelets, whether ventricose>/

1. conspicuously ventricose <basally> <Fig. 69>/

2. not ventricose <implicit>/

#134. Glumes <of sessile to subsessile female-fertile spikelets,
position relative to rachis>/

1. dorsiventral to the rachis <the entire spikelet

orientated dorsiventrally to flatwise> <Plates 1:7,

1:9; Figs 56, 70, 79>/

2. lateral to the rachis <the spikelets borne flatwise>/

3. displaced <e.g., lateral to each other on side away

from rachis>/

#135. Glumes <of female-fertile spikelets, whether hairy>/

1. hairy <Plates 1:1, 1:3, 1:8, 2:12, 2:13, 2:15, 2:16; Figs

52, 6 1 >/

2. hairless <Plate 1:5; Fig. 73>/

#136. Glumes <hairless, whether glabrous or scabrous>/

1. glabrous <Plate 1:8; Fig. 73>/

2. scabrous <Plate 1 :5>/

#137. Glumes <of female-fertile spikelets, hair disposition;*/

1 . with distinct hair tufts/

2. with distinct rows of hairs/

3. without conspicuous tufts or rows of hairs/

#138. Glumes <of female-fertile spikelets, shape of apex>/

1. pointed <Plates 1:3, 1:5, 2:13, 2:17; Figs 61, 67, 79,

85>/

2. not pointed <blunt or incised>/

#139. Glumes <of female-fertile spikelets, shape>/

1. subulate/

2. not subulate <to become implicit>/

#140. Glumes <of female-fertile spikelets, whether awned>/

1. awned <Plates 1:1, 2; 15; Fig. 32 >/

2. awnless <Fig. 73>/

#141. Glumes <of female-fertile spikelets, whether carinate
(i.e., one-keeled to middle or below)>/

1. carinate <one-keeled> <Plates 1:5, 2:17; Figs 54, 67,

73, 89, 1 14>/

2. non-carinate <includes forms with more than one

keel, as well as those with non-keeled glumes>
<Plates 1 :5, 1 :7>/

#142. Glumes <of female-fertile spikelets, whether

conspicuously winged on the median keel>/

1. with the keel conspicuously winged <Fig. 73>/

2. without a median keel-wing <implicit>/

#143. Glumes <of female-fertile spikelets, whether markedly
dissimilar in form or texture; ignore mere size
difference^

1 . very dissimilar <specify> <Plates 1 :7, 2:10, 2: 12; Figs

59, 61>/

2. <more or less> similar <Plates 1:1,2:11; Figs 66, 67,

73, 81, 85, 89 >/

#144. Lower glume <in situ length relative to upper glume of
female-fertile spikelet: not recorded if glumes more
or less equal>/

times the length of the upper glume/

#145. Lower glume clength relative to lowest lemma: not
widely recorded>/

1. shorter than the lowest lemma/

2. about equalling the lowest lemma/

3. much exceeding the lowest lemma/

#146. Lower glume <length relative to the lowest lemma
(originally introduced to deal with Colpodium/
Catabrosa)>l

1. much shorter than half length of lowest lemma/

2. longer than half length of lowest lemma/

#147. Lower glume <of female-fertile spikelets, whether

distinctly two-keeled to the middle or below
(intended mainly for andropogonoids)>/

1. two-keeled distinctly two-keeled to the middle or

below> <Plate 1:7; Figs 74, 75>/

2. not two-keeled <not distinctly two-keeled, at least

below the upper quarter> <Fig. 72>/

#148. Lower glume <of female-fertile spikelets, shape of back
(recorded mainly for andropogonoids)>/

1. convex on the back <Plate 1:7; Fig. 72>/

2. flattened on the back <Fig. 74>/

3. concave cbetween the keels> on the back/

4. sulcate on the back/

#149. Lower glume <of female-fertile spikelet, whether pitted
with 1-3 pits, cf. Bothriochloa ; not synonymous
with lacunose, qv. (intended for andropogonoids)>/

1. conspicuously pitted <Fig. 74>/

2. not pitted/

#150. Lower glume <of female-fertile spikelet, texture
(intended mainly for andropogonoids)>/

1. smooth <Plate 1:7; Fig. 60>/

2. lacunose with <several-to-many> deep depressions

<Figs 75, 76>/

3. rugose/

4. tuberculate <Fig. 59>/

5. muricate/

6. spiny <Fig. 77>/

#151. Lower glume <of female-fertile spikelet, mid-zone nerve
number>/
nerved/

#152. Upper glume <whether saccate: e.g. Sacciolepis>/

1. distinctly saccate/

2. not saccate <implicit>/

#153. Upper glume <(or the single glume) of female-fertile
spikelets, mid-zone nerve number>/
nerved/

#154. Upper glume <whether spiny>/

1 . spiny/

2. not spiny <implicit>/

#155. <Female-fertile> spikelets <whether containing sterile or
male-only florets in addition to female-fertile
florets>/

1. <normally> with female-fertile florets only/

2. <or at least some of them, normally> with incomplete

<sterile or male-only> florets <note that the situation
at the apex of spikelets with more than three florets
is often unknown or unclear> <Plates 1:8, 2:10,

2:12, 2:13, 2:16; Figs 61, 64, 71, 78, 79>/

#156. The incomplete cmale or sterile> florets <position in
spikelet>/

1. proximal to the female-fertile florets <Plates 1:8,

2:12, 2:13, 2:16; Figs 64, 71, 79>/

2. distal to the female-fertile florets <Figs 61, 78>/

3. both distal and proximal to the female-fertile florets/
#157. The distal <incomplete> florets <specialisation>/

1. merely underdeveloped <neither clearly specialised

nor peculiarly modified in form> <Fig. 6 1 >/

2. clearly specialised and modified in form <Fig. 78>/
#158. The distal <incomplete> florets <whether awned: data

very incomplete>/

1 . awned <Fig. 78>/

2. awnless/

#159. <Female-fertile> spikelets <presence or absence of
proximal incomplete florets. Strictly speaking, a
redundant character, but universally recorded and
very useful for key-making>/

1. with proximal incomplete florets <includes empty

lemmas> <Plates 1:8, 2:10, 2:12, 2:13, 2:16; Figs 64,
71, 79, 80, 84>/

2. without proximal incomplete florets <and no proximal

empty lemmas>/

#160. Proximal incomplete florets <of the female-fertile

spikelets, when present, number (intended mainly for
panicoids)>/

#161. Proximal incomplete florets <of the female-fertile
spikelets, whether paleate>/

1. paleate <Plate 2:13; Figs 71, 79, 80>/

2. epaleate <Fig. 64>/

#162. Palea of the proximal incomplete florets <development>/

1. fully developed <Fig. 84>/

2. reduced <or vestigial> <Plate 2:13; Fig. 80>/

#163. Palea of the proximal incomplete florets <whether

becoming hardened and enlarged laterally:

Paniceae>/

1. becoming conspicuously hardened and enlarged

laterally/

2. not becoming conspicuously hardened and enlarged

laterally/

394

#164. Proximal incomplete florets <of the female-fertile
spikelets: sexuality>/

1. male <Plate 2:13; Figs 71, 79, 80>/

2. sterile <Plate 2:16 >/

#165. <Proximal lemmas: shape comments>/

#166. The proximal <imperfect> lemmas <of the female-fertile
spikelets: whether awned>/

1 . awned/

2. awnless/

#167. The proximal <imperfect> lemmas <of the female-fertile
spikelets, mid-zone nerve number (intended mainly
for panicoids)>/
nerved/

#168. The proximal <imperfect> lemmas <of the female-fertile
spikelets, length relative to the female-fertile ones in
the intact spikelet (intended mainly for panicoids)>/

1. exceeded by the female-fertile lemmas <Fig. 80>/

2. more or less equalling the female-fertile lemmas <Fig.

82 >/

3. decidedly exceeding the female-fertile lemmas

<Plates 2:12, 2:16; Figs 64, 71, 84>/

#169. The proximal <imperfect> lemmas <of the female-fertile
spikelets, firmness relative to the female-fertile ones
(intended mainly for panicoids)>/

1. less firm than the female-fertile lemmas <Plate 2:16;

Figs 64,71, 80>/

2. similar in texture to the female-fertile lemmas/

3. decidedly firmer than the female-fertile lemmas/

#170. The proximal <imperfect> lemmas <of the female-fertile

spikelets, whether becoming indurated (intended
mainly for panicoids)>/

1. becoming indurated/

2. not becoming indurated <Plate 2:16; Fig. 84>/

#171. <Number of> female-fertile florets <per female-fertile

spikelet>/

#172. <Female-fertile> lemmas <insertion>/

1. conspicuously non-distichous/

2. not conspicuously non-distichous <implicit>/

#173. <Female-fertile> lemmas <shape comments:*/

#174. <Female-fertile> lemmas <whether convolute>/

1. convolute <and hiding the palea> <Fig. 85>/

2. not convolute <implicit: but data not yet reliable>/
#175. <Female-fertile> lemmas <whether saccate>/

1. saccate <Figs 81, 89>/

2. not saccate <to become implicit: but not yet reliably

so>/

#176. <Female-fertile> lemmas <firmness, relative to the
glumes>/

1. less firm than the <firmer of the> glumes/

2. similar in texture to the <firmer of the> glumes/

3. decidedly firmer than the <firmer of the> glumes

<Plate 2:16; Figs 64, 71, 81, 89, 114>/

#177. <Female-fertile> lemmas <texture: data provided for
Australian Paniceae by R.D. Webster>/

1. smooth <Figs 64, 8 1 >/

2. longitudinally, minutely> striate <rugulose> <Fig.

82 >/

3. <transversely> rugose <Figs 83, 114>/

#178. <Female-fertile> lemmas <whether becoming indurated>/

1. becoming indurated <cf. fingernails, when mature and

dry> <Plates 2:11, 2:16; Figs 81, 83, 85, 1 14>/

2. not becoming indurated <hyaline, membranous,

leathery, cartilaginous etc.> <Fig. 7 1 >/

#179. <Female-fertile> lemmas <shape of apex>/

1. entire <Figs 91, 97>/

2. incised <Plate 2:18; Figs 66, 86, 87, 9 1 >/

#180. <Female-fertile> lemmas <entire, whether pointed or
blunt>/

1. pointed <Fig. 97>/

2. blunt <Figs 88, 101 >/

#181. <Female-fertile> lemmas <number of lobes>/
lobed/

#182. <Female-fertile> lemmas <whether deeply cleft>/

1. deeply cleft <to a third or more> <Plate 2:18; Figs 66,

86, 87>/

2. not deeply cleft <Fig. 91 >/

#183. <Female-fertile> lemmas <whether crested, cf.
Cyrtococcum>/

1. crested at the tip <Fig. 89>/

2. not crested <implicit>/

#184. <Female-fertile> lemmas <whether mucronate or
awned>/

1. awnless <neither mucronate nor awned> <Figs 68, 71,
73, 89 etc.>/

2. mucronate <with a short, hard point or vestigial or

incipient awn> <Plate 2:16; Fig. 104>/

3. awned <Plates 1:5, 1:6, 2: 18; Figs 56, 61, 66, 67, 85,

87, 90, 91, 1 14>/

#185. Awns <of female-fertile lemmas, form>/

1. triple or trifid, commonly with a basal column

<Aristida typo <Figs 92, 96>/

2. not of the triple/trifid, basal column type <implicit>/
#186. Awns <of female-fertile lemmas, if present, number;*/
#187. Awns <of female-fertile lemmas, position:*/

1. median <Figs 56, 61, 67, 9 1 >/

2. median and lateral <Plate 2:18; Figs 66, 86, 87>/

3. lateral only/

#188. The median awn <whether different from the laterals in
form>/

1. different in form from the laterals <Figs 66, 86 >/

2. similar in form to the laterals <Plate 2:18; Fig. 87>/
#189. Awns <of female-fertile lemmas, position of (main,

median):*/

1. from the sinus <Figs 86, 9 1 >/

2. dorsal <Figs 56, 67, 90 >/

3. apical <Plate 1:5; Figs 61, 85, 87, 91, 92. 96, 1 14>/
#190. Awns <of dorsally awned female-fertile lemmas,

position;*/

1. from near the top <from the upper quarter, or near the

apex, or just behind an apical notch;* <Fig. 56 >/

2. from well down the back <from near the middle, or

below> <Figs 67, 90 >/

#191. Awns <of female-fertile lemmas, whether straight or
geniculate when dry >/

1 . non-geniculate <straight or curved:* <Plates 1:5,2:18;

Figs 56, 61,87>/

2. geniculate <usually twisted at the base> <Figs 66, 67,

86. 90>/

#192. Awns <main, median of the female-fertile lemmas,
hairiness:*/

1. hairless <glabrous or scabrous;* <Figs 61, 67, 92 >/

2. hairy <but not long-plumose> <Fig. 9 1 >/

3. long-plumose <Plate 2:18; Figs 87, 96 >/

#193. Awns <main, median of the female-fertile lemmas,

relative length>/

1 . much shorter than the body of the lemma <Plate 1 :5>/

2. about as long as the body of the lemma/

3. much longer than the body of the lemma <Plates 1:6,

2:18; Figs 61, 66, 67,86, 87, 9 1 >/

#194. Awns <of female-fertile lemmas, number of veins
entering base>/

1 . entered by one vein <Fig. 94>/

2. entered by several cthree or more> veins <Fig. 95>/
#195. Awns <of female-fertile lemmas, whether deciduous —

e.g. Stipa/Oryzopsis>/

1. deciduous/

2. persistent <to become implicit>/

#196. The lateral awns <relative length;*/

1 . shorter than the median <in the intact spikelet> <Figs

66, 86>/

2. about equalling the median/

3. exceeding the median/

#197. <Female-fertile> lemmas <whether hairy: excludes callus
and awns>/

1. <conspicuously> hairy <Plate 1:2; Figs 66, 86, 90>/

2. hairless <glabrous, scabrous, sparsely puberulent,

etc.> <Plate 2:16; Figs 61, 64>/

#198. The hairs <of the female-fertile lemmas>/

1. in tufts <Figs 66, 86>/

2. not in tufts <implicit>/

#199. The hairs <of the female-fertile lemmas>/

1. in transverse rows <Figs 66, 86>/

2. not in transverse rows <implicit>/

#200. <Female-fertile> lemmas <hairless, whether glabrous or
scabrous>/

1. glabrous/

2. scabrous/

#201. <Female-fertile> lemmas <whether carinate (i.e.,

one-keeled at least to the middle on the back>/

1. carinate <with a single median keel> <Plate 1:2; Figs

54, 68>/

2. non-carinate <rounded, flat, with two or more keels>

<Figs 82, 83, 86, 90, 9 1 >/

#202. <Female-fertile> lemmas <whether margins Digitaria or
Paspalum type (intended for Paniceae )>/

1. having the margins <at least over the upper
two-thirds> lying flat and exposed on the palea
<Digitaria- type> <Plate 2:13; Fig. 97>/

395

2. having the margins <at least over the lower
two-thirds> tucked in onto the palea
<Paspalum-type> <Plate 2:16; Fig. 98>/

#203. <Female-fertile> lemmas <presence of germination
flap>/

1 . with a clear germination flap <when mature> <Figs

99, 100>/

2. without a germination flap /

#204. <Female-fertile> lemmas cnumber of nerves traversing
mid-region>/
nerved/

#205. <Female-fertile> lemmas <confluence of nerves: data
very incomplete>/

1. with the nerves confluent towards the tip/

2. with the nerves non-confluent apically <Fig. 1 0 1 >/
#206. Palea <presence in female-fertile florets>/

1. present <within the female-fertile lemma>/

2. absent/

#207. Palea <female-fertile, relative size>/

1. relatively long <three-quarters or more of

female-fertile lemma length;- <Plates 2:13, 2:16;
Figs 102, 104>/

2. conspicuous but relatively short <less than

three-quarters of female-fertile lemma length> <Fig.
88>/

3. very reduced <or vestigial>/

#208. Palea <female-fertile, whether convolute>/

1 . convolute/

2. not convolute <implicit: but data not yet reliable>/
#209. Palea <female-fertile, whether gaping: especially

Aveneae>/

1 . gaping/

2. tightly clasped by the lemma <not gaping>/

#210. Palea <female-fertile, whether incised>/

1 . entire <Fig. 106>/

2. apically notched <Fig. 103>/

3. deeply bifid/

#211. Palea <female-fertile, whether with awns or setae>/

1. awnless, without apical setae <Figs 102, 103>/

2. with apical setae <Fig. 104>/

3. awned/

#212. Palea ctexture: data very incomplete>/

1. thinner than the lemma/

2. similar in texture to the lemma <Plates 2:11,2:16;

Figs 81, 97, 98>/

#213. Palea <female-fertile, whether indurated>/

1 . indurated <Plates 2:11,2:16; Figs 8 1 , 87>/

2. not indurated <Figs 103, 105 etc.>/

#214. Palea <female-fertile, nerve number>/

1 . 1-nerved <truly 1-veined, or with two contiguous

veins>/

2. 2-nerved <with two well-separated nerves> <Figs

102, 103, 106>/

3. with several nerves <specify>/

4. nerveless/

#215. Palea <female-fertile, whether dorsally 2-keeled,
one-keeled (carinate), or keel-less>/

1. one-keeled/

2. 2-keeled <Figs 65, 88, 102, 103, 105, 106>/

3. keel-less/

#216. Palea keels <whether winged: data very incomplete;*/

1. winged <Figs 105, 106>/

2. wingless <Fig. 102>/

#217. Palea keels <female-fertile, hairiness>/

1 . glabrous/

2. scabrous <Fig. 105>/

3. hairy <Figs 102, 103>/

#218. Lodicules <presence in female-fertile florets>/

1 . present/

2. absent/

#219. Lodicules <number>/

#220. <Presence of third lodicule>/

1 . third lodicule present <Fig. 108>/

2. no third lodicule/

#221. Lodicules <of female-fertile florets, whether anterior pair
joined or free>/

1. joined <at least basally> <Fig. 106>/

2. free <Figs 103, 107-1 1 1>/

#222. Lodicules <of female-fertile florets, texture>/

1. <distally> fleshy dcuneate’; panicoid type> <Figs

103, 106, 107, 109>/

2. <distally> membranous <i.e. pooid typo <Plate 2:14;

Figs 108, 1 10, 1 I I >/

#223. Lodicules <of female-fertile florets, whether hairy>/

1 . ciliate <or hairy> <Figs 103, 108, 1 10>/

2. glabrous <Figs 106, 107, 109, 1 1 1 >/

#224. Lodicules <of female-fertile florets, whether toothed>/

1. toothed/

2. not toothed/

#225. Lodicules <of female-fertile florets, vascularization.

Note: this fairly unsatisfactory character is not
equivalent to ‘presence or absence' of xylem>/

1. heavily vascularized <cf. bamboos> <Figs 108, 1 12>/

2. not or scarcely vascularized <i.e. the norm> <Figs

103, 106, 107, 1 09- 1 1 1 >/

#226. Stamens <number per female-fertile floret (not applicable
to male spikelets or male florets)>/

#227. Stamens <whether filaments joined>/

1. with free filaments <implicit>/

2. monadelphous/

3. diadelphous/

4. triadelphous/

#228. Anthers <of female-fertile florets, length: data very
incomplete, unreliable for large genera>/
mm long/

#229. Anthers <whether penicillate>/

1. penicillate <Fig. 1 14>/

2. not penicillate <Fig. 1 13>/

#230. Anthers <whether connective apically prolonged;-/

1. with the connective apically prolonged/

2. without an apically prolonged connective/

#231. Ovary <of female-fertile florets, whether apex glabrous
or hairy>/

1 . glabrous <Plate 2: 14; Figs 103, 106, 116, 11 8>/

2. hairy <Figs 110, 112, 115, 117>/

#232. Ovary <whether with a conspicuous apical appendage>/

1 . with a conspicuous apical appendage <Fig. 1 15>/

2. without a conspicuous apical appendage <implicit>

<Figs 103, 1 16, 1

#233. The <ovary> appendage <form: intended for bamboos>/

1. long, stiff and tapering/

2. broadly conical, fleshy/

#234. Styles <whether fused>/

1 . fused <at least basally; each stigma assumed to

represent one stylo <Figs 107, 112, 117, 1 18>/

2. free to their bases <Figs 103, 107, 1 10, 1 16>/

#235. Stigmas <number>/

#236. Stigmas <colour, in chasmogamous spikelets>/

1. white <Plates 1:2, 1:5, 1:7, 2:14; Figs 1 10, 111, 1 15>/

2. red <anthocyanin> pigmented <i.e. red, pink, purple

or black> <Plates 1:3, 1:4, 1:6, 1:8, 2: 10 etc.; Figs
79, 107>/

3. <golden> brown <Plate 2: 1 2>/

Fruit, embryo and seedling.

#237. Disseminule constitution: data not yet entered>/

1 . a naked seed/

2. a free caryopsis/

3. a caryopsis enclosed in but free of the lemma and

palea/

4. a caryopsis enclosed within and partially fused with

the lemma and palea/

5. consisting of the abscised spikelet/

6. consisting of the abscised spikelet and its pedicel/

7. comprising the rachis segment and associated

structures/

8. consisting of the disarticulated spikelet-bearing

inflorescence unit/

9. constituted by the complete, deciduous inflorescence/
#238. Fruit <adherence>/

1. adhering to lemma and/or palea <Fig. 123>/

2. free from both lemma and palea <but may be

enclosed:*/

#239. Fruit <length when mature>/

1 . small dess than 4 mm>/

2. medium sized <4-10 mm>/

3. large <more than 10 mm long>/

#240. Fruit <shape>/

1. linear/

2. fusiform/

3. banana-shaped/

4. ellipsoid <Fig. 1 2 1 >/

5. subglobose/

6. pyriform/

#241. Fruit <whether grooved in transverse section;-/

1. longitudinally grooved <sulcate> <Fig. 121>/

396

2. not grooved <includes terete, triangular in section,
etc.; specify> <Figs 1 22— 1 24>/

#242. Fruit <plane of compression;*/

1. compressed laterally/

2. compressed <dorsally, ventrally or> dorsiventrally

<Figs 121—123 >/

3. not noticeably compressed <Figs 1 1 9— 1 20>/

4. trigonous/

#243. Fruit <or grain surface pattern;-/

1. sculptured <Fig. 127>/

2. <relatively> smooth <the near-universal condition>

<Figs 1 2 1— 1 26>/

#244. Fruit <hair distribution;-/

1. with hairs confined to a terminal tuft <Fig. 121 >/

2. hairy on the body/

#245. Hilum <form>/

1. short <punctiform or shortly elliptical, less than half

length of fruit> <Figs 122, 124>/

2. long-linear cmore than half as long as fruit> <Figs

121, 123>/

#246. Pericarp <texture>/

1 . thin <Figs 1 1 9— 1 26>/

2. thick and hard/

3. fleshy <fruit a berry>/

#247. Pericarp <whether fused or loose (or free)>/

1. free <Figs 119, 120>/

2. loosely adherent <fairly easily removable when

soaked>/

3. fused <Figs 1 2 1 — 1 26>/

#248. Embryo <relative siz e>/

1. large <at least one-third as long as fruit> <Fig. 125>/

2. small <less than one-third as long as fruit> <Fig.

126 >/

#249. Embryo <whether waisted in surface view>/

1. waisted <Fig. 125>/

2. not waisted <Fig. 126>/

#250. Seed <whether endospermio/

1. endospermic <implicit>/

2. not endospermic/

#251. Endosperm <hard or liquid: data extensively from Terrell
1971, Rosengurtt et al. 1972>/

1. liquid <soft or milky> in the mature fruit/

2. hard/

#252. Endosperm <presence of lipid: data mainly from
Rosengurtt et al. 1972>/

1 . with lipid/

2. without lipid/

#253. Endosperm <form of starch grains: data mainly from
Tateoka 1954, 1955, 1962>/

1. containing only simple starch grains <each with only

one hilum> <Fig. 129>/

2. containing <at least some; compound starch grains

<with at least some grains having two or more hila>
<Fig. 128>/

#254. Embryo <presence of epiblast. Embro section data

extensively from Reeder 1967, 1962 and Decker
1 964>/

1. with an epiblast <Fig. 132>/

2. without an epiblast <Fig. 133>/

#255. Embryo <presence of scutellar tail>/

1. with a scutellar tail <i.e. with a cleft between

scutellum and coleorhiza> <Figs 132, 133>/

2. without a scutellar tail /

#256. Embryo <relative length of mesocotyl intemode>/

1. with an elongated mesocotyl intemode <Figs 132,

133>/

2. with a negligible <short> mesocotyl intemode/

#257. Embryo <number of scutellum bundles>/

1. with one scutellum bundle <Fig. 131 >/

2. with more than one scutellum bundle/

#258. Embryonic leaf margins <whether overlapping or

meeting>/

1. meeting <Fig. 131 >/

2. overlapping <Fig. 130>/

#259. Seedling crelative length of mesocotyl: compiled data

probably unreliable, because germination conditions
should be standardized:*/

1. with a short mesocotyl <Figs 135, 136>/

2. with a long mesocotyl <Figs 134, 137, 138>/

#260. Seedling tightness of coleoptile: data extensively from

Muller 1978>/

1. with a loose coleoptile <at least near tip> <Fig. 135>/

2. with a tight coleoptile/

#261. First seedling leaf possession of lamina>/

1. with a well-developed lamina/

2. without a lamina/

#262. The <first seedling leaf> lamina <relative width: data on
seedling leaf characters mainly from Kuwabara
1960, 1961 and H.T. Clifford (pers . comm.>/

1. broad <length/breadth, ratio less than 20> <Figs 134,

137, 138>/

2. narrow <length/breadth ratio 20 or more> <Fig. 136>/
#263. The <first seedling leaf> lamina <carriage>/

1. erect <Fig. 136>/

2. curved <Figs 134, 137>/

3. supine <Fig. 138> <Fig. 138>/

#264. The <first seedling leaf> lamina cvein number, in
middle>/
veined/

Abaxial leaf blade epidermis.

#265. Microhairs presence in abaxial leaf blade epidermis:*/

1. present <Plates 3:19, 3:22; Figs 139-142, 145, 149,

150, 152, 156, 160, 161, 164. 172, 174, 180, 182,

186 etc.>/

2. absent/

#266. Microhairs <of abaxial leaf blade epidermis, shape>/

1 . more or less spherical/

2. elongated <to become implicit:*/

#267. Microhairs <of abaxial leaf blade epidermis, number of
cells visible:*/

1. ostensibly one-celled cusually indicative of a sunken

basal cell>/

2. clearly two-celled <to become implicit;*/

3. uniseriate/

#268. Microhairs <of abaxial leaf blade epidermis, form>/

1. panicoid-type <distal cell more or less parallel-sided

or tapered to the apex; usually relatively elongated,
thin-walled, often collapsed or missing:* <Plate 3:22;
Figs 139, 140, 149. 152, 156, 160-162, 164, 174,

177, 186>/

2. chloridoid-type <distal cell inflated or more or less

hemispherical, relatively short, usually thick-walled
relative to the panicoid type, persistent> <Plate 3:19;
Figs 141, 170, 180>/

3. Enneapogon- type <long, with very long basal cell and

relatively short, inflated apical cell> <Fig. 142>/
#269. Microhairs <whether with 'partitioning membranes;*/

1. with ‘partitioning membranes’/

2. without ‘partitioning membranes’/

#270. The ‘partitioning membranes’ <iocation>/

1. in the basal cell/

2. in the apical cell/

#271. Microhairs <of abaxial leaf blade, total external length:
for species sample, see attached list plus Metcalfe
1 960>/

microns long/

#272. Microhairs <of abaxial leaf blade, width at the septum:
for species sample, see attached list>/
microns wide at the septum/

#273. Microhair apical cells <of abaxial leaf blade, length: for
species sample, see the attached list plus Metcalfe
1960>/

microns long/

#274. Microhair apical cell/total length ratio <for species

sample, see attached list plus Metcalfe 1960. Useful
approximations: 0-0.3 (a.c. markedly shorter than
b.c.); 0.3-0. 7 (a.c. and b.c. about equal); 0. 7-1.0
(a.c. markedly longer than b.c.)>/

#275. Microhair total length/width at septum <for species
sample, see attached list. Useful ranges: 0.5-1. 5
(more or less spherical); 1.5-3 (decidedly plump);
3-8 (narrow); 8^10 (very narrow);*/

#276. <Whether abaxial leaf blade epidermis shows> costal/
intercostal zonation/

1. conspicuous <Plates 3:19, 3:21, 3:22; Figs 139-141,

143, 146, 147, 151-153, 158, 169-174 etc.>/

2. lacking <Figs 144, 185>/

#277. Intercostal zones <of abaxial leaf blade epidermis,
whether of typical long-cells>/

1. <fairly exclusively> of typical long-cells <Figs 139,

143, 144, 148, 152, 160, 169, 170, 172, 184, 186
etc.>/

2. having many atypical long-cells <Figs 153, 174>/

3. without typical long-cells <Fig. 146>/

397

#278. Long-cells <of abaxial leaf blade epidermis, whether
similar in shape costally and intercostally>/

1. similar in shape costally and intercostally <Figs 178,

180>/

2. markedly different in shape costally and intercostally

<Plate 3:21; Figs 172-174, 176, 186>/

#279. Long-cells <of abaxial leaf blade epidermis, whether
similar in thickness costally and intercostally>/

1. of similar wall thickness costally and intercostally

<Figs 178, 180, 186>/

2. differing markedly in wall thickness costally and

intercostally <Fig. 147>/

#280. Mid-intercostal long-cells <of abaxial leaf blade
epidermis, shape>/

1. more or less rectangular <Plates 3:19, 3:21; Figs 139,

143, 145, 147, 149-152, 159, 160, 163, 164, 169,
170, 172, 173, 176, 177, 179, 180, 184— 186>/

2. more or less fusiform <or narrowed at ends> <Plate

3:20; Figs 144, 145, 148, 163, 176, 184>/

#281. Mid-intercostal long-cells <of abaxial leaf blade

epidermis, whether walls straight or sinuous in
(outer) optical section>/

1. having markedly sinuous <tessellated> walls <Plates

3:19, 3:21; Figs 139, 143, 145, 147, 149, 150, 153,

159, 160, 164, 165, 168-175, 177-180, 182-186 >/

2. having straight or only gently undulating walls <Plate

3:20; Figs 144, 145, 148, 156, 163, 166, 1 8 1 >/

#282. Papillae <presence in the abaxial leaf blade epidermis>/

1. present <Plates 3:19, 3:22; Figs 139-141, 151-153,

160, 174, 179, 1 8 1 >/

2. absent/

#283. <Leaf blade abaxial epidermal> papillae <general
location: data very incomplete>/

1. costal <Figs 153, 160>/

2. intercostal <Plate 3:22; Figs 151-153, 160>/

#284. <Leaf blade abaxial epidermal> papillae <whether on the
subsidiaries: data very incomplete>/

1. present on the subsidiaries <Fig. 139>/

2. absent from the subsidiaries <Figs 140, 152, 153>/
#285. Intercostal papillae <of the abaxial leaf blade epidermis,

whether over-arching the stomata (at least at one
end)>/

1. <frequently> over-arching the stomata <Plate 3:22;

Figs 140, 151, 152, 1 8 1 >/

2. not over-arching the stomata <Figs 174, 179>/

#286. Intercostal papillae <of the abaxial leaf blade epidermis,

form, arrangement^

1. consisting of one oblique swelling per cell <Plate

3:22; Figs 140, 152, 1 8 1 >/

2. consisting of one symmetrical <conical or

finger-like> projection per cell <Plate 3:19; Figs
151, 174>/

3. several per cell <specify appearance> <Figs 139, 153,

179>/

#287. Crown cells <presence in the abaxial leaf blade
epidermis>/

1. present <Fig. 154>/

2. absent/

#288. Costal regions <of the abaxial leaf blade epidermis,
presence of horizontally elongated-sinuous or
elongated-crenate silica bodies>/

1. with ‘pooid-type’ <horizontally elongated-sinuous or

elongated-crenate> silica bodies <Plate 3:20; Figs
148, 155>/

2. without significant numbers of> ‘pooid-type’

silica-bodies/

#289. Costal regions <of the abaxial leaf blade epidermis,

presence of cross-to-dumb-bell shaped or nodular
silica bodies>/

1. with ‘panicoid type’ <cross-shaped to dumb-bell

shaped or nodular> silica bodies <specify> <Plates
3:21, 3:22; Figs 139-141, 143, 145, 149, 152, 156,
169, 171, 173, 175-177, 180-182, 184— 186>/

2. without significant numbers of> ‘panicoid-type’

silica bodies/

#290. Costal regions <of the abaxial leaf blade epidermis,
presence of tall-and-narrow silica bodies>/

1. with tall-and-narrow silica bodies <Fig. 157>/

2. without significant numbers of> tall-and-narrow

silica bodies/

#291. Costal regions <of the abaxial leaf blade epidermis,

presence of saddle-shaped (chloridoid-type) silica
bodies>/

1. with saddle-shaped silica bodies <Plate 3:19; Figs

150, 151, 158, 170>/

2. without significant numbers of> saddle-shaped silica

bodies/

#292. Costal regions <of the abaxial leaf blade epidermis,
presence of crescentic silica bodies>/

1. with crescentic silica bodies <Figs 159, 170>/

2. without significant numbers of> crescentic silica

bodies/

#293. Costal regions <of the abaxial leaf blade epidermis,

presence of oryzoid-type silica bodies: i.e. vertically
orientated dumb-bells or nodules>/

1. with oryzoid silica bodies <Figs 139, 160, 164>/

2. without significant numbers of> oryzoid silica

bodies/

#294. Costal regions <of the abaxial leaf blade epidermis,

presence of silica bodies with sharp points; includes
’acutely-angled’ sensu Metcalfe>/

1. with sharp-pointed silica bodies <Figs 146, 161, 164,

173, 181 >/

2. without significant numbers of> sharp-pointed silica

bodies/

#295. Costal regions <of the abaxial leaf blade epidermis,

presence of round or oval (or potato-shaped) silica
bodies>/

1. with round to oval silica bodies <Figs 162, 178, 183>/

2. without significant numbers of> round to oval silica

bodies/

#296. Costal regions <of the abaxial leaf blade epidermis,
presence of horizontally elongated-smooth silica
bodies>/

1 . with elongated-smooth silica bodies <Fig. 163>/

2. without significant numbers of> elongated-smooth

silica bodies/

#297. Stomata <abaxial, presence in the abaxial leaf blade
epidermis>/

1. absent or very rare <Fig. 183>/

2. common <abaxially>/

#298. Stomata <of the abaxial leaf blade, end to end guard cell
length: for species sample, see attached list>/
microns long/

#299. Stomata <of the abaxial leaf blade, guard-cells

overlapped or overlapping (Watson & Johnston
1978: Aust. J . Bot. 26)>/

1. having guard-cells overlapped by the interstomatals

<Plate 3:20; Figs 166, 167>/

2. having guard-cells overlapping to flush with the

interstomatals <Plate 3:21; Figs 165, 167>/

#300. Stomata <abaxial leaf blade, presence/abundance of
triangular subsidiaries>/

1. without triangular subsidiaries/

2. <commonly> with triangular subsidiaries <Plates

3:21, 3:22; Figs 139, 143, 147, 150, 152, 153, 160,
165, 168, 169, 171, 173, 174, 177, 178, 179>/

#301. Stomata <abaxial leaf blade, presence/abundance of
parallel-sided subsidiaries^

1. without parallel-sided subsidiaries/

2. <commonly> with parallel-sided subsidiaries <Plate

3:20; Figs 144, 146, 156, 166 >/

#302. Stomata <abaxial leaf blade, whether exhibiting a mixture
of parallel-sided and triangular subsidiaries on the
same leaf>/

1. exhibiting on the same leaf a mixture of stomatal

complexes with triangular and parallel-sided
subsidiaries/

2. not exhibiting parallel-sided and triangular

subsidiaries on the same leaf /

#303. Intercostal short-cells <abaxial leaf blade, presence/

abundance — prickles and hair bases not regarded as
short-cells>/

1. common <Figs 147, 149, 150, 152, 164, 169,

171-173, 175, 176, 179, 180, 183, 184, 186>/

2. absent or very rare <Figs 144, 148>/

#304. Intercostal short-cells <abaxial leaf blade epidermal,
arrangement^

1. in cork/silica-cell pairs <Figs 152, 171-173, 175, 176,

180, 183, 1 84>/

2. not paired cnote that some short-cells recorded as

‘solitary’ probably represent superposed cork/
silica-cell pairs> <Figs 152, 169, 179, 184>/

#305. Intercostal short-cells <abaxial leaf blade epidermal,
whether silicified>/

1 . silicified <Figs 152, 171-173, 175, 176, 179, 180,

1 83>/

398

2. not silicified <Figs 152, 169, 184>/

#306. Costal short-cells <abaxial leaf blade epidermal,

arrangement of short-cells; prickles, hair bases not
counted as short-cells>/

1. conspicuously in long rows <of five or more cells>

<Plates 3:19, 3:21, 3:22; Figs 140, 141, 143, 145,

146, 150-153, 156, 158, 160. 161, 164, 169,

171-175, 177, 180, 182, 184— 186>/

2. predominantly paired <Figs 159, 170, 178>/

3. neither distinctly grouped into long rows nor

predominantly paired <solitary; in short rows,
mixtures of solitaries, pairs, short rows, etc.> <Figs

147, 148, 157, 162, 163, 176, 1 8 1 >/

Transverse section of leaf blade, physiology, culm anatomy.

#307. <Maximum cells-distant count; indicating photosynthetic
pathway; see Hattersley & Watson 1975:
Phytomorphology 25>/

1 . <showing a maximum cells-distant count of one,

reliably predicting> C4 <Plates 3:23, 3:24; Figs 187,
188, 192-196, 198, 207-213, 222, 224-226>/

2. <showing a maximum cells-distant count of two or

more, reliably predicting> C3 <Plates 3:25, 3:26;

Figs 189-191, 197, 216, 218, 219, 223, 227>/

#308. The <C4> anatomical organization <of the leaf blade,
whether conventional/

1. conventional <to become implicit> <Plates 3:23, 3:24;

Figs 187-188, 192-196, 210, 212, 222>/

2. unconventional <Figs 198, 208-209. 211. 2I3>/

#309. Organization of <leaf blade> PCR tissue <when

unconventional/

1. Triodia type <with the PCR cells forming a layer

draping (at least in places) from one bundle to the
next, rather than constituting discrete bundle
sheaths> <Figs 198, 213>/

2. Alloteropsis type <with two bundle sheaths, the inner

being PCR> <Figs 208, 209>/

3. Aristida type <the PCR cells constituting a double

bundle sheath>/

4. Arundinella type <with single PCR files or groups in

the mesophyll, in addition to the conventional PCR
sheath> <Fig. 2 1 1 >/

#310. <C4> biochemical type <as determined by enzyme assay:
data from Hatch and Kagawa 1974, Gutierrez et al.
1974(a) and 1974(b), Hatch, Kagawa and Craig
1975, and Prendergast, Hattersley and Stone 1987.
Species samples in parentheses>/

1 . PCK/

2. NAD-ME/

3. NADP-ME/

#311. <Leaf blade XyMS: reliably indicative of C4 type

(Hattersley & Watson 1976: Aust. ./. Bot. 24. N.B.,
ascertainable from major vascular bundles only)>/

1 . XyMS+ <C3, or C4 'aspartate formers’ type PCK or

NAD-ME (exceptions: Eriachneae)> <Plates
3:24-26; Figs 187, 189-193, 195-198, 203-205,

212, 214, 220, 223, 224>/

2. XyMS- <C4 ‘malate formers’, type NADP-ME>

<Plate 3:23; Figs 188, 199, 208, 209, 222 >/

#312. <Leaf blade> PCR sheath outlines <in C4 forms> <data
extensively from Ellis 1977, and Prendergast and
Hattersley 1987>/

1. uneven <PCK or ‘PCK-like’, Figs 187, 193-195,207,

210>/

2. even <NAD-ME or ‘NAD-ME-like’, lFigs 188, 192,

196, 212>/

#313. <Leaf blade> PCR sheath extensions <presence> <data
mainly from H.D.V. Prendergast 1987>/

1. present <in at least some veins> <Figs 187, 193,

207 >/

2. absent <Figs 1 94 — 1 96>/

#314. Maximum number of deaf blade PCR sheath> extension
cells <data mainly from H.D.V. Prendergast 1987>/
#315. <Leaf blade> PCR cells <of C4 forms, presence of a
suberised lamella> <cf. Hattersley and Browning
1 98 1 >/

1. with a suberised lamella <Figs 199-203 >/

2. without a suberised lamella <Figs 204-206>/

#316. <Leaf blade> PCR cell chloroplasts <of C4 forms, shape>
<data from H.D.V. Prendergast 1987, Prendergast,
Hattersley and Stone 1987>/

1. ovoid <Figs 193, 195>/

2. elongated <Figs 194, 196, 205, 206>/

#317. <Leaf blade> PCR cell chloroplasts <of C4 forms,

whether granal. See Gutierrez et al. (1974). Carolin
et al. (1973), Hattersley and Browning (1981 )>/

1. with well developed grana <Figs 204-206>/

2. with reduced grana <Figs 1 99— 203>/

#318. <Leaf blade> PCR cell chloroplasts <position. Data
extensively from Ellis 1977, Brown 1960,
Prendergast and Hattersley 1987>/

1 . centrifugal/peripheral <sometimes NAD-ME. more

often indicative of NADP-ME or PCK> <Plate 3:23;
Figs 193, 195. 199, 201-203 >/

2. centripetal <NAD-ME: predominant in arid and

semiarid species> <Figs 192, 194, 196, 204-206>/
#319. <Leaf blade> PBS cells <of C3 forms>/

1. with a suberised lamella/

2. without a suberised lamella/

#320. Leaf blade chlorophyll a.b ratio <data from Prendergast
1987>/

#321. <Leaf blade> mesophyll <whether chlorenchyma radiate:
an ill-defined feature, not reliably indicative of
photosynthetic pathway>/

1. with radiate chlorenchyma <Plates 3:23, 3:24, 3:26;

Figs 187, 192, 195-197, 207-210 etc.>/

2. with non-radiate chlorenchyfna <Plate 3:25: Figs

189-191, 213 etc.>/

#322. <Leaf blade> mesophyll <presence of palisade>/

1. with <a clear> adaxial palisade <Figs 214, 216>/

2. without <any obvious> adaxial palisade/

#323. <Leaf blade> mesophyll <presence of Isachne- type

mesophyll>/

1. Isachne-lype <Plate 3:26; Fig. 197>/

2. not Isachne-lype/

#324. <Leaf blade> mesophyll <presence of ‘circular cells" (i.e.
isolated PCR cells or cell groups; ‘distinctive
cells’)>/

1. exhibiting ‘circular cells' <Fig. 21 1>/

2. without ‘circular cells’/

#325. <Leaf blade> mesophyll <whether traversed by (at least
some) columns of colourless cells>/

1. traversed by columns of colourless cells <P!ate 3:24;

Figs 192, 2 10>/

2. not traversed by colourless columns <Figs 187, 188,

193, 212, 224. 225 etc.>/

#326. <Leaf blade> mesophyll <presence of arm cells (=
‘ratchet’ cells)>/

1 . with arm cells <Figs 2)3, 215. 2 1 6>/

2. without arm cells/

#327. <Leaf blade> mesophyll <presence of fusoid cells>/

1 . with fusoids <Figs 214, 2 1 6— 220>/

2. without fusoids/

#328. The fusoids <whether part of the PBS>/

1 . an integral part of the PBS <Fig. 2 1 9 >/

2. external to <though contiguous with> the PBS <Figs

214-216, 218>/

#329. Leaf blade <ribbing>/

1. with distinct, prominent adaxial ribs <only> <Figs

187, 189. 190, 227, 228>/

2. ‘nodular’ in section <Plate 3:26; Fig. 191 >/

3. adaxially <more or less> flat <ignore mid-rib.

Includes forms with abaxial ribs only> <Plates 3:23,
3:24; Figs 212, 224, 226>/

#330. Leaf blade <adaxial ribs, relative sizes>/

1 . with the ribs more or less constant in size <Fig. 187>/

2. with the ribs very irregular in sizes <i.e. of two or

more size orders; ignore the mid-rib> <Figs 189,

190, 227>/

#331. Midrib <of the leaf blade, prominence>/

1. conspicuous <prominent in the outline, with

distinctive sclerenchyma, etc.> <Plate 3:25; Figs
221. 222>/

2. not readily distinguishable from other main veins

<other than by position;-/

#332. Midrib <of the mid leaf blade, vascularization>/

1. with one bundle only/

2. with a conventional arc of bundles <i.e. at least three

bundles> <Fig. 222>/

3. vascularization complex <i.e. more than one bundle,

not arranged in a conventional arc> <Fig. 22 1 >/
#333. Midrib <and/or middle part of leaf blade, whether
extensively of colourless cells adaxially>/

1. with <conspicuous> colourless tissue adaxially <Fig.

222 >/

2. without <conspicuous> colourless tissue adaxially/

399

#334. The lamina <in transverse section, symmetry around the
midrib>/

1 . distinctly asymmetrical on either side of the midrib

<usually involving marked asymmetry in the ribbing
and/or the form of the margin; e.g. as in many
bamboos>/

2. symmetrical on either side of the midrib/

#335. <Presence in the adaxial leaf blade of discrete adaxial
groups of bulliforms; exclude ‘hinges' flanking
midribs>/

1. bulliforms present in discrete, regular adaxial groups

<Plates 3:23, 3:24; Figs 187, 188, 191-197, 207,

208, 210, 212, 216, 221, 223-225 >/

2. no discrete, regular groups of adaxial bulliforms

<absent, exclusively in irregular groups or
constituting most of the epidermis> <Plate 3:25;

Figs 189, 219, 226>/

#336. <Presence in the adaxial leaf blade of simple fan-shaped
bulliform groups>/

1. bulliforms occurring in simple fan-shaped groups <i.e.

without associated colourless cells> <Figs 187, 190,
191, 196, 197, 207, 209, 216, 220, 223 >/

2. without simple fans of bulliforms <ignore midrib

‘hinges’>/

#337. <Presence in the adaxial leaf blade of deeply-penetrating
fans of combined bulliforms and colourless cells>/

1. having <at least some> bulliforms combined with

colourless cells to form deeply-penetrating
fan-shaped groups <Plate 3:24; Figs 187, 192, 210,
212>/

2. without deeply-penetrating fans of

bulliforms-plus-colourless cells/

#338. <Presence in the adaxial leaf blade of narrow groups of
bulliforms-plus-colourless cells>/

1. having <at least some> bulliforms associated with

colourless cells to form narrow groups penetrating
into the mesophyll <Fig. 224>/

2. without narrow-penetrating groups of

bulliforms-plus-colourless cells/

#339. <Whether bulliforms and associated colourless cells form
arches in the leaf blade>/

1. bulliforms and associated colourless cells

<sometimes> forming arches over small vascular
bundles <Figs 188, 225 >/

2. without bulliform-plus-colourless cell arches/

#340. <Presence in the leaf blade of small vascular bundles

unaccompanied by sclerenchyma>/

1. many of the smallest vascular bundles unaccompanied

by sclerenchyma <Plate 3:23; Figs 222, 226 >/

2. all <or nearly all> the vascular bundles accompanied

by sclerenchyma/

#341. <Presence in the leaf blade of vascular bundles

combining both adaxial and abaxial girders>/

1. exhibiting vascular bundles <at least some, if only the

midrib> combining both adaxial and abaxial girders
of sclerenchyma <Plate 3:24, 3:25; Figs 187-195,
197. 212, 216, 220, 223-225, 227>/

2. without vascular bundles combining adaxial with

abaxial girders of sclerenchyma <cf. Figs 194, 208,
209>/

#342. The combined girders <adaxial and abaxial sclerenchyma
girders, whether forming ‘anchors’, I’s or T’s in one
or more bundles of the leaf blade (include the
midrib)>/

1. forming ‘figures’ <‘anchors', I’s or T’s> <in at least

some bundles> <Plate 3:24; Figs 187-193, 195, 212,
216, 223, 227>/

2. nowhere forming ‘figures’ <l.e. no ‘anchors’, I’s or

T’s>/

#343. Sclerenchyma <whether all leaf blade sclerenchyma is
bundle-associated;*/

1. all associated with vascular bundles <apart from any

marginal fibres>/

2. not all <obviously> bundle-associated <Plate 3:25;

Fig. 227>/

#344. Culm internode bundles <arrangement; poorly recorded,
data mainly from Metcalfe 1960 >/

1. in one or two rings <ignore ‘outer rings’ of very few

bundles>/

2. in three or more rings/

3. scattered/

#345. Stem tissues of the culm bases <whether accumulating

abundant starch: data from Smith 1968, Smouter and
Simpson 1989 and original observations^

1. with abundant starch/

2. with little or no starch cimplying fructans and/or

sucrose>/

#346. Fructosans predominantly <short- or long-chain>/

1. short-chain/

2. long-chain/

Special diagnostic features.

#347. <Anomochloa>/

1 . inflorescence of 2-3 glumeless, bracteate spikelets,

the lodicules represented by a fringed annulus/

2. plant not as in Anomochloa <implicit>/

#348. <Arundo/Phragmites>/

1. female-fertile lemmas conspicuously hairy; ligule

hairs to 0.3 mm long, shorter than the membrane/

2. female-fertile lemmas hairless; ligule hairs longer

than 0.5 mm, longer than the membrane/

#349. <Atractantha>/

1. the inflorescences of very peculiar pseudospikelets,

characterized by development of rachides with long
terminal segments, each of which serves as the
pedicel of an abscissile spikelet/

2. the inflorescences not as in Atractantha <implicit>/
#350. <Briza>/

1. lemmas as broad as long, gibbous and umbonate,

cordate at base <Briza> <Fig. 88>/

2. lemmas not as in Briza <implicit>/

#35 1 . <Brylkinia>/

1. lemma awn winged, the wing extending down the

upper back of the lemma/

2. lemma not wing-awned <implicit>/

#352. <Buchloe>/

1. the male inflorescences elevated, with one to four

spicate, unilateral branches; female spikelets in
burr-like clusters, usually two burrs per
inflorescence, each burr on a short, stout rachis,
partially enclosed in a broad, bractlike leaf sheath,
falling entire with the indurate rachis united with the
upper glumes/

2. not as in Buchloe <implicit>/

#353. <Centrochloa>l

1. upper glume extended downwards into a conspicuous

spur/

2. upper glume not as in Centrochloa <implicit>/

#354. <C haetobromus>l

1. pedicels articulated and bearded with long hairs at and

above the joint/

2. pedicels not as in Chaetobromus/

#355. <Coix>/

1. inflorescences in hard, globular 6-12 mm utricles/

2. inflorescences not as in Coix <implicit>/

#356. <Cortaderia/Lamprothyrsus>/

1. the lemma awns lateral and median, the median

strongly flattened/

2. the median lemma awn not strongly flattened, laterals

present or absent/

#357. <Corynephorus>/

1 . lemmas awned, the awn bearing a ring of minute hairs

at the middle, and apically clavate <Fig. 93>/

2. lemmas without the characteristic Corynephorus awn

<implicit>/

#358. <Cyperochloa>/

1. the inflorescence of a few digitately-borne, bracteate

spikelets, subtended by a spatheate leaf atop a single
elongated culm internode, the plant very sedge-like
in appearance/

2. plants not as in Cyperochloa <implicit>/

#359. <Diandrostachya>/

1. the lower glume exceeding the female-fertile lemma/

2. the lower glume shorter than the female-fertile

lfemma/

#360. <Diarrhena>t

1. grain with a conspicuous whitish or yellowish, glossy

beak/

2. fruit not as in Diarrhena <implicit>/

#361. <Dichanthelium>/

1. plants from a short rosette of winter leaves, the

primary panicle producing secondary inflorescences
with cleistogamous spikelets/

2. plants not as in Dichanthelium/

#362. <Enneapogon/Schmidtia> ICottealKaokochloal

1. female-fertile lemmas 9-lobed, each lobe terminating
in an awn/

400

2. female-fertile lemmas 6-lobed and 5-awned, with an

awn arising between each pair of lobes/

3. female-fertile lemmas irregularly lobed, the lobes

produced into 7-1 1 awns/

4. female-fertile lemmas with an incurved-emarginate

apex, and a narrow awned lobe at each margin
(sometimes with 1-2 shorter, additional lobes)/

#363. <Eriochloa>/

1. spikelets supported on a peculiar, hardened,

cupuliform ‘callus’ <Plate 2:16 >/

2. no Eriochloa-type ‘callus’ <implicit>/

#364. <Hackelochloa/Hemarthria/Rottboellia>/

1. lower glume of female-fertile spikelet globose, pitted/

2. lower glume of female-fertile spikelet flattish, not

pitted; ‘pedicellate’ spikelets similar to the
female-fertile spikelets/

3. lower glume of female-fertile spikelet flattish, not

pitted; ‘pedicellate’ spikelets reduced, herbaceous/
#365. <Hubbardia>l

1. plants of wet places, the leaves remarkably thin and

delicate/

2. plants not as in Hubbardia <implicit>/

#366. <Hydrothauma>/

1. the adaxial surface of the leaf blade raised into

sinuous lamellae/

2. the adaxial surface of the leaf blade not as in

Hydrothauma/

#367. <Hygroryza>l

1. plants aquatic, with inflated leaf sheaths serving as

floats/

2. plants not as in Hygroryza <implicit>/

#368. <KoeleriafTrisetum>l

1. panicle dense, cylindrical, ovoid, not interrupted:

awns if present straight, subterminal, inconspicuous
in the inflorescence/

2. panicle loose, or if dense then interrupted, neither

cylindrical nor ovoid: awns usually present, usually
twisted, usually distinctly dorsal, conspicuous if
inflorescence compact/

#369. < Leptaspis and Scrotochloa>/

1. having female spikelets, with shell- or urn-shaped

lemmas which are closed save for an apical pore/

2. not having female spikelets as in Leptaspis or

Scrotochloa <implicit>/

#370. <Lombardochloa>l

1. female-fertile lemma very broad, with a conspicuous,

succulent, translucent region near the base of each
wing/

2. female-fertile lemma not as in Lombardochloa

<implicit >/

#37 1 . <Lopholepis>/

1. spikelets minute, shaped like cartoon birds’ heads/

2. spikelets not as in Lopholepis <implicit>/

#372. <Lygeum> /

1. plant coarsely tufted, with wiry leaf blades, the

inflorescence of one very peculiar spikelet enclosed
in a sheath/

2. plant and inflorescence not as in Lygeum <implicit>/
#373. <Manisuris>/

1. spikelets in ‘false pairs’, the pedicellate member of

the andropogonoid pair abscinding from its pedicel
but remaining attached to the base of the ‘article’
above, alongside the sessile member of that ‘article’/

2. spikelets not arranged as in Manisuris <implicit>/
#374. <Melica et al .>/

1 . spikelets with the distal incomplete florets and/or the

rachilla apex forming a terminal clavate appendage/

2. spikelets without a terminal clavate appendage

<implicit>/

#375. <Merxmuellera/Karroochloa/Chaetobromus/Schismus>/

1. female-fertile lemmas with a bent awn, the awn

twisted below/

2. female-fertile lemmas awnless, mucronate or with a

short straight awn/

#376. <Merxmuellera/Karroochloa>/

1. spikelets 8-25 mm long, inflorescence longer than 60

mm long/

2. spikelets 4 — 6(— 7) mm long, inflorescence 10-60 mm

long /

#377. <NasseIla>/

1. spikelet with a single gibbous floret, the lemma awn

placed off-centre/

2. spikelet not as in Nassella <implicit>/

#378. <Odontelytrum>/

1. the inflorescence a coarse, cylindrical ‘raceme’,

apparently representing a raceme of reduced
‘glomerules’, each glomerule shortly pedunculate,
comprising a single spikelet subtended crosswise by
a lobed scale forming an involucre-plus-bristle/

2. the inflorescence not as in Odontelytrum <implicit>/
#379. <PaspalumlEchinochloa/Paspalidium>l

1. glumes and/or sterile lemmas awned or

acuminate-mucronate/

2. spikelets awnless, muticous/

3. spikelets awnless, the female-fertile lemmas pointed

or apiculate but not mucronate/

#380. <Phaenosperma> /

1. seed dark brown, with ruminate endosperm/

2. seed not as in Phaenosperma <implicit>/

#381. <Phyllorhachis>/

1. spikelets borne on one side of a broad, leaflike rachis/

2. spikelets not borne on a broad, leaflike rachis

<implicit>/

#382. <Prosphytochloa>/

1. scandent via leaf blades with retrorsely scabrid

margins/

2. not scandent as in Prosphytochloa <implicit>/

#383. <Rhynchoryza> /

1 . female-fertile lemma with its tip extended beyond the

palea as a conical, herbaceous beak (flotation
device) composed of aerenchyma with transverse
septa, tapering into an awn/

2. female-fertile lemma not as in Rhynchoryza/

#384. <Sorghastrum/Sorghum>/

1. spikelets ostensibly solitary, each accompanied by a

barren pedicel/

2. spikelets paired, all the pedicels spikelet-bearing/
#385. <Spartochloa, Xerochloa>/

1. rush-like, with reduced leaf blades/

2. not rush-like <implicit>/

#386. <Spinifex>/

1. female inflorescence a large, deciduous globular head

of sessile, bristle-tipped racemes <Fig. 26>/

2. inflorescence not as in Spinifex <implicit>/

#387. <Steyermarkochloa>/

1. culms dimorphic, the fertile culms leafless, the

vegetative culms each with a single developed leaf,
this being eligulate and with a terete, culm-like
‘sheath’/

2. plants not as in Steyermarkochloa <implicit>/

#388. <Thuarea>/

1. flowering culms ultimately bending over, so as to

enclose the ripening fruit/

2. flowering culms not as in Thuarea <implicit>/

#389. <Thyridolepis>/

1. lower glume <of female-fertile spikelet> with a

rectangular window, surmounted by bristles <Fig.
70>/

2. without a Thyridolepis- type window <implicit>/

#390. <Urelytrum>/

1. the lower glume of the pedicellate spikelet with a

5-10 mm (or longer) awn/

2. the lower glume of the pedicellate spikelet awnless/
#391. <Viguierella>/

1. the inflorescence a spicate ‘raceme’, with each

spikelet subtended at its base by a tiny hyaline bract:
Madagascar/

2. not Viguierella <implicit>/

#392. <Zea>/

1. fruiting inflorescence a massive, spatheate cob, the

fruits in many rows/

2. fruiting inflorescence not as in Zea <implicit>/

#393. <Zygochloa>/

1. stems cane-like, spikelets in bracteate, globular 1-3.5

cm heads/

2. plants not as in Zygochloa <implicit>/

Cytology.

#394. Chromosome base number, x =/

#395. <Diploid chromosome numbers> 2 n =/

#396. <Recorded ploidy levels: data very incomplete;*/
ploid/

#397. Mean diploid 2c DNA value <range and number of
species studied in parenthesis: data mainly from
Bennett and Smith (1976) and Bennett, Smith and
Heslop-Harrison (1982)>/

Pg/

401

Taxonomy .

#398. <Subfamily: Watson et al. 1985>/

1. Pooideae/

2. Bambusoideae/

3. Arundirioideae/

4. Chloridoideae/

5. Panicoideae/

#399. <Supertribes of Watson et al. 1985, with name endings
changed>/

1. Triticodae/

2. Poodae/

3. Oryzodae/

4. Bambusodae/

5. Panicodae/

6. Andropogonodae/

#400. cTribe of Pooideae>/

1. Triticeae/

2. Brachypodieae/

3. Bromeae/

4. Aveneae including Agrostideae, Phalarideae>/

5. Meliceae/

6. Seslerieae/

7. Poeae including Hainardieae, Monermeae>/

#401. <Tribe of Bambusoideae>/

1 . Oryzeae/

2. Olyreae/

3. Centotheceae/

4. Anomochloeae/

5. Brachyelytreae/

6. Diarrheneae/

7. Ehrharteae/

8. Phaenospermateae/

9. Phyllorhachideae/

10. Phareae/

1 1 . Streptochaeteae/

12. Streptogyneae/

13. Guaduelleae/

14. Puelieae/

15. Bambuseae/

#402. cTribe of Arundinoideae>/

1. Stipeae/

2. Nardeae/

3. Lygeae/

4. Arundineae/

5. Danthonieae <and satellites>/

6. Micraireae/

7. Aristideae/

8. Eriachneae/

9. Steyermarkochloeae/

10. Spartochloeae/

1 1 . Cyperochloeae/

#403. cTribe of Chloridoideae>/

1 . Triodieae/

2. Pappophoreae/

3. Orcuttieae/

4. Chlorideae sensu lato cthe main chloridoid

assemblage, including Cynodonteae, Eragrosteae,
Sporoboleae, Aeluropodeae, Jouveae, Unioleae,
Leptureae, Lappagineae, Spartineae, Trageae,
Perotideae, Pommereulleae>/

#404. cTribe of Panicoideae>/

1. Isachneae/

2. Paniceae/

3. Neurachneae/

4. Arundinelleae/

5. Andropogoneae/

6. Maydeae/

#405. cSubtribe of Andropogoneae>/

1. Andropogoninae c'awned Andropogoneae’>/

2. Rottboelliinae c'awnless Andropogoneae’>/

#406. ‘Nearest neighbours’ cin ascending order of ‘distance’,

according to DIST calculations conducted in 1985.
Note that these are safely interpretable as ‘closest
taxonomic relatives’ only when reciprocal lists are in
agreement. The misleading appearance of the same
large genera (Poa, etc.) in many lists probably
reflects their internal variability;*/

Ecology, geography, regional floristic distribution.

#407. cNumber of species>/
species/

#408. cGeographic distribution;*/

#409. eWorld distribution: this ‘character’ is intended only for
convenience in key-making — for more precise
distributions, see ‘geographical distribution’;*/

1. Western Eurasia, U.S.S.R. cincludes Iran, Iraq,

Turkey;*/

2. Mediterranean/

3. Eastern Asia cJapan, China to India>/

4. Africa cand Saudi Arabia>/

5. Pacific cMalaysia, Indonesia, Australasia, Pacific

Islands>/

6. North America cCanada, Alaska, U.S.A., Mexico/

7. South and Central America, West Indies/

8. Arctic/

#410. <Whether commonly adventive on an intercontinental
scale>/

1. commonly adventive/

2. not commonly adventive <implicit>/

#411. <Habitat water requirement;*/

1. hydrophytic/

2. helophytic <i.e., in marshy places>/

3. mesophytic/

4. xerophytic/

#412. cHabitat light requirement;*/

1. shade species/

2. species of open habitats/

#413. <Salt tolerance, etc.>/

1 . halophytic/

2. glycophytic <= not halophytio/

#414. cHabitat notes: soil types, etc.>/

#415. cGeographical occurrence in Australasia — mainly after
Simon 1978>/

1 . Tasmania/

2. New South Wales/

3. Australian Capital Territory/

4. Victoria/

5. Western Australia/

6. Queensland/

7. Northern Territory/

8. South Australia/

9. New Guinea/

10. New Zealand/

1 1. not known in Australasia <implicit>/

#416. cGeographical distribution in southern Africa;*/

1. Namibia/

2. Botswana/

3. Transvaal/

4. Orange Free State/

5. Swaziland/

6. Natal/

7. Lesotho/

8. Cape Province/

9. not in southern Africa cimplicit>/

#417. cStatus in southern Africa>/

1. indigenous species/

2. naturalized species cin southern Africa>/

3. cultivated/

#418. cGeographical distribution in North America;*/

#419. cNumber of species in the Flora North America region:
data from Kartesz and Kartesz 1980>/
species in North America/

#420. cStatus in North America: Flora North America region>/

1. indigenous species/

2. naturalized species cin North America;*/

3. cultivated/

#421. cFloristic Kingdoms: after Takhtajan 1969. Data deduced
from information for Takhtajan’s floristic regions
(see below), provided by B. K. Simon 1987>/

1. Holarctic/

2. Paleotropical/

3. Neotropical/

4. Cape/

5. Australian/

6. Antarctic/

#422. cHolarctic Subkingdoms: after Takhtajan 1969>/

1. Boreal/

2. Tethyan cancient Mediterranean^

3. Madrean cSonoran>/

#423. cPaleotropical Subkingdoms: after Takhtajan 1969>/

1. African/

2. Madagascan/

3. Indomalesian/

4. Polynesian/

5. Neocaledonian/

402

#424. <Boreal Subkingdom regions: after Takhtajan 1969 >/

1. Arctic and Subarctic/

2. Euro-Siberian/

3. Eastern Asian/

4. Atlantic North American/

5. Rocky Mountains/

#425. cTethyan Subkingdom regions: after Takhtajan 1969 >/

1. Macaronesian/

2. Mediterranean/

3. Irano-Turanian/

#426. <African Subkingdom regions: after Takhtajan 1969 >/

1. Saharo-Sindian/

2. Sudano-Angolan/

3. West African Rainforest/

4. Namib-Karoo/

5. Ascension and St. Helena/

#427. <Indomalesian Subkingdom regions: after Takhtajan
1969 >/

1. Indian/

2. Indo-Chinese/

3. Malesian <Malayan>/

4. Papuan/

#428. Polynesian Subkingdom regions: after Takhtajan 1969 >/

1. Hawaiian/

2. Polynesian/

3. Fijian/

#429. <Neotropical regions: after Takhtajan 1969 >/

1 . Caribbean/

2. Venezuala and Surinam/

3. Amazon/

4. Central Brazilian /

5. Pampas/

6. Andean/

7. Fernandezian/

#430. <Australian regions: after Takhtajan 1969>/

1. North and East Australian/

2. South-West Australian/

3. Central Australian/

#431. <Antarctic regions: after Takhtajan 1969 >/

1. New Zealand/

2. Patagonian/

3. Antarctic and Subantarctic/

#432. <Euro-Siberian Subregions>/

1 . European/

2. Siberian/

#433. <Atlantic North American Subregions>/

1 . Canadian-Appalachian/

2. Southern Atlantic North American/

3. Central Grasslands/

#434. <Sudano-Angolan Subregions>/

1. Sahelo-Sudanian/

2. Somalo-Ethiopian/

3. South Tropical African/

4. Kalaharian/

#435. <North and East Australian Subregions>/

1. Tropical North and East Australian/

2. Temperate and South-Eastern Australian/

#436. <Antarctic and Subantarctic Subregions>/

1 . South Temperate Oceanic Islands/

2. Antarctic/

Hybrids.

#437. dntergeneric hybrids>/

Rusts and smuts.

#438. Rusts — <genera: data from Cummins 1971, his

classification amended by D.B.O. Savile (pers.
comm.). Updating beyond Cummins confined as yet
to grass nomenclature. Unnamed species of
Agropyron , Elymus, Panicum etc. ignored>/

1. Dasturellal

2. Phakopsora/

3. PhysopellaJ

4. Stereostratuml

5. Puccinia <including Uromyces>l

6. no rusts recorded <by Cummins 1 97 1 >/

#439. The Puccinia species from <morphological Group - —
after Cummins 1971, amended by D.B.O. Savile
(pers. comm.)>/

1. Group 1/

2. Group 2/

3. Group 5/

4. Group 6/

5. Group 7/

6. Group 8/

#440. The Puccinia species from <the Group 1 species, Savile’s
subgroups>/

1. subgroup 1(a)/

2. subgroup 1(b)/

3. subgroup 1(c)/

4. subgroup 1(d)/

#441. The Puccinia species from <the Group 2 species, Savile’s
subgroups;-/

1. subgroup 2(a)/

2. subgroup 2(b)/

#442. The Puccinia species from <the Group 5 species, Savile’s
subgroups;-/

1 . subgroup 5(a)/

2. subgroup 5(b)/

3. subgroup 5(c)/

4. subgroup 5(d)/

5. subgroup 5(e)/

6. subgroup 5(f)/

7. subgroup 5(g)/

8. subgroup 5(h)/

9. subgroup 5(i)/

10. subgroup 5(j)/

11. subgroup 5(k)/

#443. The Puccinia species from <the Group 6 species, Savile’s
subgroups;-/

1. subgroup 6(a)/

2. subgroup 6(b)/

3. subgroup 6(c)/

4. subgroup 6(d)/

5. subgroup 6(e)/

#444. Wide-ranging <rust> species: <wide-ranging here =
recorded on 3 or more host genera by Cummins
( 1 97 1 )>/

1 . Dasturella divina/

2. Phakopsora incompleta/

3. Physopella clemensiael

4. Stereostratum corticoidesl

5. Puccinia chaetochloael

6. Puccinia stenotaphri/

7. Puccinia microspora/

8. Puccinia polysoral

9. Puccinia miscanthael

10. Puccinia nakanishikii/

1 1. Puccinia longicornis/

12. Puccinia kusanoi/

13. Puccinia eritraeensis/

14. Puccinia graminella/

15. Puccinia dolosa/

16. Puccinia oriental is/

17. Puccinia graminis/

18. Puccinia levis/

19. Puccinia substriata/

20. ‘ Uromyces ' setariae-italicae/

21. ‘ Uromyces ' schoenanthi/

22. Puccinia emaculata/

23. Puccinia cacabata/

24. Puccinia coronata/

25. Puccinia striiformis/

26. Puccinia montanensis/

27. Puccinia pygmaea/

28. Puccinia brachypodii-phoenicoidis/

29. Puccinia brachypodii/

30. Puccinia praegracilis/

3 1 . Puccinia poarum/

32. Puccinia hordei/

33. Puccinia recondita/

34. ‘ Uromyces ' turcomanicuml

35. 'Uromyces' fragilipesl

36. 'Uromyces' dactylidis/

37. 'Uromyces' hordeinus/

38. Puccinia monoica/

39. Puccinia versicolor/

40. Puccinia boutelouae/

4 1 . Puccinia chloridis/

42. Puccinia schedonnardi/

43. 'Uromyces' clignyi/

44. 'Uromyces' eragrostidis/

45. Puccinia miyoshiana/

46. Puccinia cesatii/

47. Puccinia esclavensis/

48. Puccinia aristidae/

403

49. no wide-ranging rust species <i.e. the positive
records limited to rusts with restricted host ranges,
as given by Cummins 1971: implicit>/

#445. Smuts <families: data not yet updated from Watson
(1972), and Panicum, Danthonia , Agropyron ,

Elymus etc. omitted pending nomenclatural checking
of records>/

1. from Tilletiaceae/

2. from Ustilaginaceae/

3. not recorded <implicit: but see qualification;-/

#446. <Smut genera> Tilletiaceae — /

1 . Entyloma/

2. Melanotaenium/

3. Neovossia/

4. Tilletia/

5. Urocystis/

#447. <Smut genera> Ustilaginaceae — /

1 . Sorosporium/

2. Sphacelotheca/

3. Tolyposporellal

4. Tolyposporium /

5. Ustilago/

Economic importance .

#448. Significant weed species: clist extended from Hiifliger
and Scholtz 1980>/

#449. Cultivated fodder:/

#450. Important native pasture species:/

#451. Grain crop species:/

#452. Lawns and/or playing fields:/

#453. Commercial essential oils:/

#454. <Miscellaneous economic/ethnic data> clittle yet
entered>/

References, etc.

#455. Morphological/taxonomic references: <articles of special
interest listed here have only rarely provided most of
the morphological descriptive data. The latter reflect
compilations from the separately listed ‘main
sources’, plus original observations by Watson and
associates (notably S.G. Aiken, H.T. Clifford, C.R.
Frylink, G.E. Gibbs Russell and T.D. Macfarlane>/
#456. Leaf anatomical references: <‘original observations’ by
Watson, or for Pooideae from Macfarlane 1979
supplemented by Watson. Note the need to account
for taxonomic realignments (especially in Triticeae)
when using Metcalfe 1960>/

Special comments.

#457. <Special comments>/

Additional characters under consideration.

#458. Plants <diameter>/

cm in diameter/

#459. Culms cnumber of aerial nodes>/

noded/

#460. Culms <habit>/

1. self-supporting <implicit>/

2. decumbent/

3. scrambling/

4. scandent/

5. pendent/

6. floating/

#461. Culm leaves <presence>/

1 . present/

2. absent/

#462. Upper culm leaf blades/

1 . fully developed/

2. reduced/

3. vestigial <i.e. leaves reduced to sheaths — not to be

confused with blade ahscission>/

#463. Culm nodes <exposure>/

1. exposed/

2. hidden by leaf sheaths/

#464. The <distal> incomplete florets <number>/

#465. Awn <of female-fertile lemmas, when non-geniculate,
shape>/

1. straight/

2. recurving/

3. flexuous/

#466. Awn bases <of female-fertile lemmas, whether twisted>/

1. twisted/

2. not twisted/

#467. Awn bases <of female-fertile lemmas, whether
flattened;*/

1. flattened/

2. not flattened/

#468. Palea back <indumentum>/

1. glabrous/

2. scabrous/

3. hairy/

#469. Styles <fusion>/

1. completely fused/

2. joined below/

3. free/

#470. Style bases <degree of separation^

1. adjacent/

2. widely separated/

#471. Fruit <indumentum>/

1 . glabrous/

2. scabrous/

3 hairy/

#472. <Fruit colour>/

#473. Fruiting lemma <of Paniceae, colour of mature L2: data
from Webster 1986>/

1 . white/

2. yellow/

3. brown/

4. black/

#474. <Cornucopiae> /

1. spikelets in numerous small, compact, short-branched

panicles, each panicle at the tip of a stout, recurved
peduncle and enclosed by a leathery, toothed
involucre, the peduncles themselves subtended by
the inflated sheaths of the (modified) upper leaves/

2. spikelets not borne as in Cornucopiae/

#475. Haploid nuclear DNA content <2c value divided by
ploidy: ranges and means>/

Pg/

#476. The ‘extra’ sclerenchyma <location of leaf blade

sclerenchyma not associated with vascular bundles
— exclusive of any in the midrib>/

1. in abaxial groups/

2. in a continuous abaxial layer/

3. within the mesophyll/

4. in adaxial groups/

#477. The ‘extra’ sclerenchyma <position of groups within the
lamina — exclusive of midrib>/

1 . abaxial-hypodermal, the groups isolated <opposite

bulliforms and/or furrows>/

2. abaxial-hypodermal, the groups continuous with

colourless columns/

3. adaxial-hypodermal, contiguous with the bulliforms/
#478. Awns <of female-fertile lemmas, whether hooked

(‘uncinate’)>/

1. hooked/

2. not hooked <implicit>/

Southern African grass species
- character list

405

In contrast to the generic character list above, the following
character list for southern African species has been deliberately
designed to be as short and simple as possible, in order to fulfil
its role as a prototype for the next level of approximation of
species data in the Taxon component of PRECIS (Gibbs Russell
& Arnold 1989). This character list incorporates the ‘common
knowledge’ characters included as ‘type one’ data in the ILDIS
legume database (ILDIS Coordinating Centre 1986), with the
addition of diagnostic characters and voucher specimens as
recommended by the 1987 meeting of the Herbarium Curators
Working Group.

Obviously, a full-scale character list at species level detailed
enough to carry sufficient data to allow classification, key
generation and detailed descriptions would require several
hundred characters, and data-capture would be the work of years.
The copious data held in text characters for distinguishing
between species and for habitat information will provide a firm
basis for development of a complete species-level character list.
Expansion to full-scale automated descriptions in DELTA will
be an important next step in the study of grasses in southern
Africa.

#1. References. <genera only>/

#2. <Synonyms>/

#3. <Vernacular names>/

#4. <Life form, following Raunkiaier 1936>/

1. tree or large shrub <phanerophyte>/

2. shrub or dwarf shrub <chamaephyte>/

3. perennial cherb - hemicryptophyte>/

4. bulb or corm <cryptophyte>/

5. annual <therophyte>/

6. biennial/

#5. <Habit>/

1. epiphyte/

2. climber/

3. scrambler/

4. parasite/

5. hydrophyte/

6. <long-> rhizomatous/

7. stoloniferous/

8. tufted/

9. succulent/

10. <other>/

#6. <Height of plant in mm>/
mm tall/

#13. cHabitat, e.g., moisture, insolation, substrate, etc.>/

#14. Conservation status: - IUCN categories 1986>/

1. extinct <Ex>/

2. endangered <E>/

3. vulnerable <V>/

4. rare <R>/

5. conservation status indeterminate <I>/

6. not endangered <0>/

7. conservation status not <or insufficiently> known

<K>/

#15. <Abundance, modified from Radford et al. 1974, but rare
categories included above in Conservation Status>/

1 . infrequent/

2. locally common <state area or habitat>/

3. common <abundant>/

4. locally dominant <state area or habitat>/

5. <widely> dominant/

#16. <Whether indigenous or naturalized;*/

1. indigenous <implicit>/

2. naturalized/

3. invader/

#17. <Area of origin for naturalized taxa >/

#18. distribution - FSA territory>/

1. Namibia/

2. Botswana/

3. Transvaal/

4. Orange Free State/

5. Swaziland/

6. Natal/

7. Lesotho/

8. Cape/

9. Other territories <specify>/

#19. distribution -> biome: <after Rutherford & Westfall
1986>/

1. Fynbos/

2. Savanna/

3. Grassland/

4. Nama-Karoo/

5. Succulent Karoo/

6. Desert/

7. Forest/

8. Afromontane/

#20. distribution - outside southern Africa>/

#7. Leaf blades <length>/
mm long/

#8. Leaf blades <width>/
mm wide/

#9. Spikelets <length>/
mm long/

#10. Spikelets <width>/
mm wide/

#11. distinguishing species characters>/

#12. Flowering <months>/

1. July <7>/

2. August <8>/

3. September <9>/

4. October <10>/

5. November <11 >/

6. December <12>/

7. January <1 >/

8. February <2>/

9. March <3>/

10. April <4>/

11. May <5>/

12. June <6>/

#21. dmportance to man, after SEPASAL; give details as
comment>/

1 . food and drink/

2. domestic use <e.g., ornaments, utensils and tools>/

3. timber deluding fuel>/

4. pasture <specify whether planted>/

5. barrier/

6. erosion control/

7. ornamental <specify whether established or

potential/

8. indicator <specify indicator of what>/

9. fibers/

10. poisonous/

1 1. medicinal/

12. traditional medicine/

13. chemicals/

14. weed <or other problem plant, or invasive>/

#22. dotes and comments>/

406

#23. <Reference to> description:/

1. Stapf 1898-1900/

2. Hitchcock & Chase 1950/

3. Chippindall 1955/

4. Clayton et al. 1970-1982/

5. FI. PI. Afr./

6. <other>/

#24. <Reference to> illustration:/

1 . Chippindall 1955/

2. Clayton et al. 1970-1982/

3. Flower. PI. Afr./

4. Hitchcock & Chase 1950/

5. <other>/

#25. <Reference to map>/

#26. Voucher: <state collector and specimen number>/

#27. PRECIS code/

#28. Species treatment by/

#29. <Divisions of the Cape>/

1. northern/

2. central/

3. eastern/

4. southern/

5. southwestern/

6. northwestern <Namaqualand>/

#30. <Divisions of Namaqualand>/

1. Richtersveld/

2. Namaqualand Rocky Hills/

3. Sandveld/

4. Knersvlakte/

407

Parameters for generic keys

The keys to genera were produced by the program KEY
(Dallwitz 1974, Dallwitz & Paine 1986) from Watson’s
database of world grass genera. The following are the
program parameters for each part of the generic key.

Key to Keys

Characters - 447 in data, 3 included, 3 in key.

Items - 4 in data, 4 included, 4 in key.

RBASE = 1.40, ABASE = 2.00, REUSE = 1.01, VARYWT
= .70

Preset characters (character, column: group) - 445,1:1
64,3:1

Characters included - 64 156 445
Character reliabilities - 64,7 156,8 445,9

Key 1.

Characters - 447 in data, 50 included, 4 in key.

Items - 206 in data, 6 included, 6 in key.

RBASE = 1.40, ABASE = 2.00, REUSE = 1.01, VARYWT
= .70

Characters included - 3-6 8 — 49 444-447
Character reliabilities - 3,7 4,6 5-6,7 8-10,7 1 1,8 12-13,7

14,6 15,5 16-17,7 18,6 19-21,7 22,5 23,7 24,6 25-26,8
27-29,7 30,1 31-36,7 37-38,6 39-43,7 44,1 45-46,7
47-48,4 49,5 444-445,7 446,8 447,7
Items included - 14 17 123 129 142 183

Key 2.

Characters - 447 in data, 261 included, 1 13 in key.

Items - 206 in data, 87 included, 129 in key.

RBASE = 1.40, ABASE = 2.00, REUSE = 1 .01, VARYWT
= .70

Preset characters (character, colummgroup) - 111,4:3
172,5:3 165,7:10 102,8:5 109,8:7 151,8:11 100,9:9
Characters included - 3-6 8-55 61-104 106-244 342 346
348 353 357 363-365 374 376 393-394 397 431-443
Character reliabilities - 3,7 4,6 5-6,7 8-13,7 14,6 15,5
16-17,7 18,6 19-21,7 22,5 23,7 24,5 25-26,8 27-29,7 30,1
31-36,7 37-38,6 39-43,7 44,1 45-16,7 47^18,4 49,5 50,8

51.6 52-55,7 61-64,7 65,6 66-68,7 69,6 70-72,7 73-75,5
76-81,7 82,4 83-88,7 89,6 90-92,7 93,4 94,7 95,5 96,7

97,5 98-104,7 106,1 107-116,7 117,5 118,6 119-142,7

143.8 144-151,7 152,5 153-155,7 156,8 157,7 158,6
159-162,7 163,6 164-165,7 166,5 167,8 168-173,7 174,6
175-177,7 178,5 179-181,7 182,8 183-189,7 190,5
191-193,7 194,6 195-213,7 214-220,4 221,2 222,5 223,4

224.6 225-226,4 227-229,5 230,6 231-243,5 244,3 342,6

346.8 348,6 353,6 357,6 363-365,6 374,7 376,6 393,5
394,4 397,1 431-435,6 436-439,5 440-443,7

Items included - 2 5 7-9 1 1-13 16 18 20-21 29 33 35 37-38
43—44 50 54-56 60-61 63-64 69-70 72-74 79 83-84 87-9 1
101 105 107 110 113-115 118 121 123-124 126-128 130
132-135 140-142 149-150 155-157 159 162 166-169
173-174 180-181 184-185 187 190 192 197 199 201-203

Key 3.

Characters - 447 in data, 261 included, 73 in key.

Items - 206 in data, 58 included, 87 in key.

RBASE = 1.40, ABASE = 2.00, REUSE = 1.01, VARYWT
= .70

Preset characters (character, column .group) - 64,1:1 70,2:2
46,2:3 131,4:4 46,5:15 109,6:13

Characters included - 3-6 8-55 61-104 106-244 342 346
348 353 357 363-365 374 376 393-394 397 431-443
Character reliabilities - 3,7 4,6 5-6,7 8-13,7 14,6 15,5
16-17,7 18,6 19-21,7 22,5 23,7 24,6 25-26,8 27-29,7 30,1
31-36,7 37-38,6 39-42,7 43,5 44,1 45-46,7 47-48,4 49,5

50.8 51,6 52-55,7 61-64,7 65,6 66-68,7 69,6 70-72,7
73-74,5 75-8 1 ,7 82,4 83-88,7 89,6 90-94,7 95,5 96-104,7
106,1 107-116,7 117,5 118,6 119-142,7 143,8 144-151,7

152.5 153-155,7 156,8 157-165,7 166,5 167,8 168-173,7

174.6 175-181,7 182,8 183-189,7 190,5 191-193,7 194,6
195-213,7 214-220,4 221,2 222,5 223,4 224,6 225-226,4
227-229,5 230,6 231-243,5 244,3 342,8 346,8 348,8 353,8

357.8 363-365,8 374,8 376,8 393,5 394,4 397,1 431-435,8
436—443,7

Items included - 1 19 22-24 26 28 31-32 36 42 46 49 52
57-59 62 65-66 68 71 74-75 77 80-81 86 98-100 102-104
109 111-112 117 119 125 131 139 145-146 148 154
164-165 170 172 182 186 188-189 191 193 204 206

Key 4.

Characters - 447 in data, 261 included, 92 in key.

Items - 206 in data, 67 included, 100 in key.

RBASE = 1.40, ABASE = 2.00, REUSE = 1 .01, VARYWT
= .70

Preset characters (character, column.group) - 46,1:1
111,2:1 111,2:2 123,4:1 180,4:4 434,6:4 52,11:4 107,15:3
Characters included - 3-6 8-55 61-104 106-244 342 346
348 353 357 363-365 374 376 393-394 397 43 1-143
Character reliabilities - 3-4,6 5-6,7 8-1 1,7 12,6 13,7 14,6

15,5 16-17,7 18,6 19-21,7 22,5 23,7 24,6 25-26,8 27,5
28-29,7 30,1 31-36,7 37-38,6 39-12,7 43,5 44,1 45-46,7
47—18,4 49,5 50,8 51,6 52-55,7 61-64,7 65,6 66-68,7 69,6
70-72,7 73-74,5 75-81,7 82,4 83-88,7 89,6 90-94,7 95,5
96-104,7 106,1 107-116,7 117,5 118,6 119-142,7 143,8
144-151,7 152,5 153-155,7 156,8 157-165,7 166,5 167,8
168-169,7 170,5 171-173,7 174,6 175-177,7 178,5
179-180,7 181-182,8 183-187,7 188,8 189,7 190,5
191-193,7 194,6 195-205,7 206,6 207-208,7 209,5 210,7
211,4 212,7 213,6 214-220,4 221,2 222,5 223-226,4
227-229,5 230,6 231-239,5 240,4 241-242,5 243,4 244,3

342,8 346,6 348,8 353,6 357,6 363,8 364-365,6 374,6

376,6 393,5 394,4 397,1 431-432,6 433-434,7 435,6
436^139,5 440-443,7

Items included - 3—4 6 10-11 13-15 24-28 30-31 34 36
39 — 4 1 47-48 51-53 59 67 71 76-78 82 85 92-97 108-109
111 120 122 136-138 143-144 147 151-153 160-161 163
171-172 175-179 189 195 200 204

409

LITERATURE REFERENCES

The following list contains references quoted in the
introductory sections and appendices, as well as major taxonomic
references applicable to several genera. An additional 125
taxonomic references pertain only to a single genus or species.
They are not included here but appear in abbreviated form in the
generic and species treatments.

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ADAMSON, R.S. & SALTER, T.M. 1950. Flora of the Cape
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AVDULOW, N.P. 1931. Karyo-systematische Untersuchung der
Familie Gramineen. Bulletin of Applied Botany, Genetics
and Plant Breeding, Supplement 44.

BENTHAM, G. 1883. Genera plantarum, Vol. 3. L. Reeve,
London.

BJORKMAN, O. 1976. Adaptive and genetic aspects of C4
photosynthesis. In R.H. Burris & C.C. Black, Metabolism
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BOR, N.L. 1985. Gramineae. In R.D. Meikle, Flora of Cyprus
Vol. 2. Bentham-Moxon Trust, Royal Botanic Gardens,
Kew.

BROWN, W.V., HARRIS, W.E. & GRAHAM. J.D. 1959. Grass
morphology and systematics I. The internode. Southwestern
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CAMPBELL, C.S. & KELLOGG, E.A. 1987. Sister group
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C.S. Campbell & M.E. Barkworth, Grass systematics and
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CHIPPINDALL, L.K.A. 1955. A guide to the identification of
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CHIPPINDALL, L.K.A. & CROOK, A.O. 1976. 240 grasses of
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CLARKE, L.G. & FISHER, J.B. 1987. Vegetative morphology
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CLAYTON, W.D. 1970. Gramineae (Part 1). In E. Milne-
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Crown Agents, London.

CLAYTON, W.D. 1972. Gramineae. In F.N. Hepper, Flora of
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CLAYTON, W.D. 1972. The awned genera of Andropogoneae.
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CLAYTON, W.D. 1972. The awnless genera of Andropogoneae.
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CLAYTON, W.D. 1981. Evolution and distribution of grasses.
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CLAYTON, W.D., PHILLIPS, S.M. & RENVOIZE, S.A. 1974.
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CLAYTON, W.D. & RENVOIZE, S.A. 1982. Gramineae (Part
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CLAYTON, W.D. & RENVOIZE, S.A. 1986. Genera graminum.
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CLIFFORD, H.T. 1987. Spikelet and floral morphology. In T.R.
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CLIFFORD, H.T. & WATSON, L. 1977. Identifying grasses:
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CRONQUIST, A. 1981. An integrated system of classification of
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DALLWITZ, M.J. 1974. A flexible computer system for
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DALLWITZ, M.J. 1980. A general system for coding taxonomic
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ELLIS, R.P. 1984. Eragrostis walteri, a first record of non-Kranz
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413

GLOSSARY

abaxial: the side away from the central axis
(opposite: adaxial).

achene: a small dry indehiscent fruit with a
single seed and a thin pericarp. True
achenes may not occur in Poaceae, but in
a few genera (e.g. Pentameris) a
caryopsis that resembles an achene is
formed when the endocarp and/or
mesocarp collapses at a late stage of
development (see caryopsis, utricle).

acicular: needle-shaped, i.e, narrow, stiff,
pointed, and round in cross-section (a
solid shape).

acropetal: development from the base
towards the tip, i.e., with the youngest
cells at the base and maturing toward the
tip (see basipetal).

arenicolous: growing in sand.

aristate: tipped with a bristle-like point.

articles: segments of a structure that
separate at maturity.

articulate: with a joint between parts that
separate cleanly at maturity.

articulation: a joint, e.g. between the
column and the lower part of the lemma
in Aristida and Stipagrostis.

ascending: curved upwards and approaching
erect. —

auricle: an ear-like outgrowth, e.g. of the
leaf sheath mouth or blade base (see Fig.
5,p. 7).

acuminate: tapering gradually to a point,
with the sides of the apex somewhat
concave.

acute: tapering to a point, with the sides of
the apex straight or somewhat convex.

adaxial: the side toward the central axis
(opposite: abaxial).

adnate: united with another organ (compare
oppressed).

adventive: a non-indigenous species that is
established in a new region but is not
expanding its range (compare
naturalized).

amplexicaul: with the base of a leaf blade
clasping the stem.

awn: a long thin stiff appendage at the tip (or
less commonly from the back or base) of
a glume, lemma or palea (see PI. 10, 12,
13, p. 358).

axil: the angle between a stem and its branch
(or leaf).

axis: a generalized term for the main stem of
the plant, of an inflorescence or of
inflorescence parts such as racemes or
spikelets (plural: axes).

basal: at or towards the base (opposite:
apical).

basipetal: development from the tip toward
the base, i.e., with the youngest cells at
the tip and maturing toward the base (see
acropetal).

annual: completing the life cycle in a year,
usually passing the unfavourable season as
a seed (see biennial, perennial).

beak: in Aristida, a slight narrowing of the
lemma below the awns, as distinct from
a longer straight column.

annular: ring-like, or arranged in a circle.

bearded: with a tuft of long stiff hairs.

anther: the part of a stamen that contains the
pollen (see filament, also Fig. 8, p. 11).

anthesis: the period during which the flower
is open and pollination takes place.

antrorse: pointing upward or forward
(opposite: retrorse).

apical: at or towards the apex (opposite:
basal).

biennial: living two years, setting seed and
dying in the second year; rare in grasses
(Pentaschistis) (see annual, perennial).

bifid: cleft into two lobes at the tip.

biseriate: lying side-by-side on one side of
the rachis (see distichous).

blade: the part of the leaf distal to the sheath
and ligule (see Fig. 5, p. 7).

apiculate: tipped abruptly with a small sharp
point.

apomixis: asexual production of seeds.

appressed: pressed against another organ,
but not united with it (compare adnate).

bract: a reduced leaf in the inflorescence,
including structures such as spathes,
spatheoles, glumes, lemmas and paleas.

C3: photosynthesis in which atmospheric
carbon is first fixed in three-carbon
chains. It is indicated anatomically by the

414

separation of chlorenchymatous cells
from the nearest PCR cells by two to
many other chlorenchymatous cells (see

Q).

C4: photosynthesis in which atmospheric
carbon is first fixed in four-carbon
chains. It is indicated anatomically by the
separation of chlorenchymatous cells
from the nearest PCR cells by no more
than one other chlorenchymatous cell
(see C3).

caespitose: growing in tufts, e.g.

Hyparrhenia hirta.

culm: the stem of a grass plant.

cuneate: wedge-shaped, widest near the
apex and tapering to a narrow base (a
solid shape).

cupuliform: cup-shaped (a solid shape).

deciduous: falling off naturally (abscising)
at a particular stage of growth (see persis-
tent).

decumbent: growing horizontally at the base
and then curving upwards (see Fig. 4, p.
6).

callus: a hard projection at the base of a
floret, spikelet or inflorescence segment
that aids in seed dispersal.

capillary: very fine and hair-like.

capitate: forming head-like clusters.

carinate: boat-shaped with one median keel
(a solid shape).

cartilaginous: tough but elastic.

caryopsis: the fruit (‘grain’) of most grasses,
in which the seed coat is adnate to the
pericarp (see achene, utricle).

dfeflexed: bent downward, but not to 180

degrees.

dentate: toothed, with teeth perpendicular to
the margin.

dichotomous: with equally forked paired
branches.

digitate: like the fingers of a hand, with the
members arising from the same point.

dioecious: a species with separate male and
female plants (see monoecious).

disarticulate: to break apart at the joints.

cataphyll: scale-like leaf on rhizomes,
stolons or at plant bases (see Fig. 3, p. 5).

caudate: with a long tail-like tip.

discoid, disc-shaped (a solid shape).

disseminule: the part of the plant released
with the seeds and aiding their dispersal.

chartaceous: papery in texture and usually
not green in colour.

distal: farther from the point of attachment
(opposite: proximal).

chasmogamous: with the florets opening for
pollination (see cleistogamous).

distichous: two-ranked, on opposite sides of
a stem (see biseriate).

ciliate: fringed with spreading, stiff hairs,
ciliolate: minutely ciliate.
clavate: club-shaped (a solid shape),
claveilate: barely clavate.

cleistogamous: with the florets not opening
for pollination,and therefore obligately
self-fertilizing (see chasmogamous).

divaricate: spreading widely.

dorsal: an ambiguous term for the back,
abaxial or outer surface of an organ
(opposite: ventral). The term abaxial is
preferable.

dorsiventral: the plane from the ‘dorsal’
(abaxial) to the ‘ventral’ (adaxial) sur-
faces; having distinct upper and lower
faces.

cleistogene: a floret that does not open for
pollination.

column: 1) in Aristida and Stipagrostis, a
straight structure between the apex of the
lemma and the branching point of the
awns (see beak)\ 2) in geniculate awns,
the part of the awn below the bend that
is often twisted (see Pis. 12 and 13, p.
358, respectively).

convolute: rolled from one side, with one
margin inside and one outside (see
involute).

cordate: heart-shaped, with rounded lobes (a
solid shape).

coriaceous: leathery in texture,
crateriform: cup-shaped (a solid shape).

dorsiventrally flattened: structures that are
compressed on the adaxial and abaxial
sides.

eglandular: without glands (see gland).

elaiosome: part of a disseminule specialized
to accumulate oils that attract ants which
disperse the seeds.

elliptic: rounded and broadest at the middle
and gradually narrowed to both ends,
with the width about 1/2 the length (a flat
or outline shape).

embryo: the rudimentary plant inside the
seed.

endemic: a species that is native to a
particular area and occurs naturally
nowhere else in the world (compare
indigenous, naturalized).

endosperm: food reserve tissue in the seed,
containing starch, oil and protein.

gibbous: swollen on one side, e.g. the lemma
of Sacciolepis or Nassella.

entire: with a continuous margin or apex, not
indented in any way.

erect: growing straight up (see Fig. 4, p. 6).

erose: a margin or apex that is irregularly
notched, as if gnawed. —

exserted: projecting beyond a containing
structure (opposite: included).

extravaginal: branching in which the young
shoot breaks through the leaf sheath (see
intravaginal).

falcate: sickle-shaped (a solid shape).

glabrous: without hairs, but not necessarily
smooth (opposite: pubescent).

gland: a secretory structure that can be either
raised or depressed.

glaucous: covered with a greyish or whitish
waxy bloom obscuring the natural colour.

globose: spherical (a solid shape, in contrast
to orbicular).

glomerate: densely clustered in heads.

glume: one of a pair of empty bracts at the
base of a spikelet.

false spike: a very narrow panicle with the
spikelets borne in tight clusters on much
reduced side-branches, as in some species
of Setaria and Pennisetum (see Fig. 6, p.
9).

fascicle: a fairly tight cluster.

female-fertile: florets (or spikelets) with an
ovary that can develop into a fruit. Fertile
stamens may or may not be present (see
sterile).

filament: the stalk of a stamen (see anther,
also Fig. 8, p. 11).

filiform: thread-like, cylindrical and very
slender. ~~

Fimbriate: fringed with long slender

processes.

glycophytic: receiving its moisture from
fresh water (compare halophytic).

grain: the caryopsis or naked fruit of a grass.

granular: with a bumpy surface.

granulose: with a slightly bumpy surface.

halophytic: growing in salty water, or salty
soil (compare glycophytic).

helophytic: growing in marshy places (see
me sophy tic , xerophytic).

hermaphrodite: bisexual; a plant, spikelet
or floret with both male and female parts.

heterogamous: of different sexes, e.g.. with
sterile and female-fertile spikelets (see
homogamous).

flabellate: fan-shaped; applied to flattened
basal leaf sheaths, e.g. in Eustachys
paspaloides (a solid shape).

flexuous: 1) not rigid; 2) zig-zag or wavy.

floret: an individual grass flower, usually
consisting of lodicules, stamens and a
pistil enveloped by the lemma and palea.

heteromorphic: of two different forms, e.g..
paired sessile and pedicellate spikelets
that differ in appearance (see
homomorphic , also PI. 92, p. 367).

heterospiculate: inflorescences with two
different kinds of spikelets, e.g. male-
fertile and female-fertile spikelets that
differ in appearance.

fragile: easily broken, especially along a line
of abscission.

free: a structure not united to any other
structures.

fruit: the ripe ovary with its adnate parts,
containing the seed. In most grasses it is
difficult to distinguish the fruit (caryop-
sis) from the seed.

fusiform: spindle-shaped; slender, but

broadest at the middle and tapering to
both ends (a solid shape).

geniculate: bent abruptly, like a knee.

geophytic: plants with the growing point
below the soil surface, often with bulb-
or corm-like structures.

hilum: the scar on the caryopsis marking the
site of attachment of the pericarp and seed
coat. It is on the side opposite the embryo.

hispid: hairy with bristly, straight, erect, stiff
hairs.

homogamous pairs: in Andropogoneae,
spikelets of a pair that are similar to each
other in sexuality (see PI. 21, p. 359), in
contrast to those which are paired but
differ in sexuality (see heterogamous)

homomorphic: of similar form, e.g. pairs of
spikelets in the Andropogoneae, in which
the spikelets of the pair are similar to
each other in form (see heteromorphic ,
also PI. 21, p. 359).

hyaline: thin and transparent.

germination flap: a line of thinner tissue in
hardened lemmas, that allows the root of
the germinating embryo to emerge.

hydrophytic: growing in water,
imbricate: overlapping.

416

imperfect: a flower lacking functional male
or female parts (opposite: perfect).

incised: cut or deeply toothed.

included: not protruding from a containing
structure (opposite: exserted).

indigenous: a species native to a particular
area, but not restricted to that area
(compare endemic, naturalized).

indurated: hardened at maturity.

inflorescence: the spikelet-bearing system
of branches (see Fig. 6, p. 9).

ligule: a membrane or line of hairs on the
adaxial leaf surface at the junction of the
sheath and the blade (see Fig. 5, p. 7).
Uncommonly an external ligule, or
contraligule, is present on the abaxial side
also, as in Alloteropsis semialata.

linear: long and narrow, with parallel sides,
usually more than ten times longer than
wide (a flat or outline shape).

lodicules: small rounded or scale-like

structures at the base of the stamens and
pistil in the grass flower which become
turgid at anthesis, thus opening the
lemma and palea (see Fig. 8, p. 11).

internode: the portion of a stem lying
between two nodes.

interrupted: with broken continuity, applied
to dense inflorescences with occasional
gaps.

intravaginal: branching in which the young
shoot emerges between the culm and the
sheath mouth (see extravaginal).

introgression: a series of hybrid generations
in which the hybrid individuals breed
back to one parental species, eventually
introducing into the parental species
characteristics of the other.

invader: an indigenous or non-indigenous
species that aggressively replaces natural
vegetation (compare naturalized).

maritime: growing by the sea.

membranous: thin and semitransparent but
not dry.

mesophytic: growing in places with fairly
abundant moisture (see helophytic,
xerophytic).

midrib: the main central vein of a leaf (see
Fig. 5, p. 7).

monoecious: plants bearing both male and
female flowers on the same individual
(see dioecious).

mucro: a minute sharp point or shortly
excurrent central nerve.

mucronate: with a mucro or mucros.

involucre: a series of bracts or bractlike
structures below a spikelet or spikelet
cluster, applied variously to the bristles of
Pennisetum and Setaria, the sterile
spikelets at the raceme bases in Themeda
or the lower glumes in Anthephora.

involute: rolled from both margins toward
the middle, thus with both margins inside
(see convolute).

muricate: rough with sharp, hard, irregular
protruberances.

muticous: blunt, without a point.

NAD-ME: a biochemical variant of C4
photosynthesis in which aspartate com-
pounds are formed (see NADP-ME and
PCK , as well as Fig. 1 and pp. 2-A for
particulars).

keel: a sharp fold or ridge on a compressed
sheath, blade, glume, lemma or palea (see
carinate).

Kranz anatomy: leaf blade anatomical
organization usually associated with C4
photosynthesis (see non-Kranz anatomy ,
as well as Fig. 1 and pp. 2-4 for
particulars).

lacerate: torn at the edges or irregularly
cleft.

lacunose: with depressions, or pitted with
shallow irregular holes.

lanceolate: lance-shaped; widest in the basal
third and gradually narrowed apically,
approximately three times longer than
wide (a flat or outline shape).

lateral: relating to the side. Spikelets whose
lemmas are 1 -keeled are usually ‘laterally
compressed’.

NADP-ME: a biochemical variant of C4
photosynthesis in which malate com-
pounds are formed (see NAD-ME and
PCK, as well as Fig. 1 and pp. 2-4 for
particulars).

naturalized: a non-indigenous species that
forms, self-sustaining populations under
local conditions and is capable of
expanding its range (compare adventive,
endemic, indigenous, invader).

nerves: the ‘veins’ of the blades, glumes,
lemmas and paleas.

node: the part of the stem where leaves
and/or branches arise (see Fig. 5, p. 7).

nodding: bent over and hanging down.

non-Kranz anatomy: leaf blade anatomy
indicative of C3 photosynthesis (see
Kranz anatomy, as well as Fig. 1 and pp.
2-4 for particulars).

lemma: the lower of the two bracts enclosing oblate: broadly elliptic,

the grass flower (see palea and Fig. 8, p.

11).

417

oblong: with parallel sides and longer than
wide (a flat or outline shape).

I

perennial: a plant that lives for more than
two years (see annual , biennial).

obovate: rounded, broadest above the
middle and narrower toward the base (a
flat or outline shape, see ovate).

obtuse: blunt and rounded at the apex (a flat
or outline shape).

orbicular: round (a flat or outline shape, in
contrast to globose).

perfect: a flower with both male and female
parts functional (opposite: imperfect).

pericarp: the outer layer of the grass fruit,
formed from the wall of the ovary.

persistent: remaining attached for a long
time, usually after other parts have been
shed (see deciduous).

ovary: the female part of the flower enclos-
ing the ovule that develops into the seed
(see Fig. 8, p. 11).

pilose: hairy with very long, soft, rather
straight hairs, not dense but somewhat
shaggy.

ovate: like the outline of a hen’s egg,
broadest below the middle and narrower
toward the apex (a flat or outline shape,
see obovate). ~ *

paired: spikelets occurring in groups of two,
usually with one spikelet short-
pedicellate or sessile and the other long-
pedicellate (see PI. 165, p. 375).

palea: the upper of the two bracts enclosing
the grass flower (see lemma and Fig. 8,
P- 11).

pallid: pale in colour.

panicle: an inflorescence in which the
primary axis bears branched secondary
axes and pedicellate spikelets (see Fig. 6,
p. 9).

papillate (or papillose): having small
protruberances (papillae).

partial inflorescences: portions of the large
compound inflorescence in Andropogon-
eae, separated from each other by leaves,
spathes and spatheoles (see Fig. 6, p. 9).

PCA tissue: in plants with C4

photosynthesis, specialized mesophyll
tissue where primary carbon assimilation
from atmospheric C02 takes place (see
PCR tissue).

PCK: a biochemical variant of C4

photosynthesis, in which aspartate com-
pounds are formed (see NAD-ME and
NADP-ME, as well as Fig. 1 and pp. 2-4
for particulars).

PCR tissue: in plants with C4

photosynthesis, specialized cells, often
but not always sheathing the vascular
bundles, where secondary carbon

reduction takes place (see PCA tissue).

pectinate: comb-like.

pedicel: the stalk of the spikelet (see Fig. 7,

p. 10).

pistillate: bearing pistils only and no
stamens; the term may be applied to a
flower, a floret, a spikelet, an inflores-
cence or an entire plant (see staminate).

plicate: folded lengthwise several times
(pleated).

procumbent: lying on the ground but not
rooting at the nodes (see Fig. 4, p. 6).

prophyll: a scale-like modified leaf with two
keels.

prostrate: lying flat on the ground.

proximal: nearer to the base or point of
attachment (opposite: distal).

pseudopetiolate: a grass leaf in which the
blade is narrowed to a slender stem-like
structure just distal to its junction with the
sheath.

puberulent: hairy with very short, erect,
straight hairs barely visible to the naked
eye.

pubescent: a generalized term for hairy,
lacking definition of the type of hairs
(opposite: glabrous).

punctiform: in the shape of a dot or point.

pungent: sharp-pointed.

raceme: an unbranched inflorescence in
which the primary axis directly bears
pedicellate spikelets (see Fig. 6, p. 9).

rachilla: the axis of a spikelet.

rachis: the axis of a spike or raceme.

, radical: pertaining to the roots.

reflexed: abruptly bent downward or

backward to about 180 degrees.

retrorse: pointing downward or backward
(opposite: antrorse).

pedicellate: with a pedicel (see sessile).

peduncle: 1) the specialized uppermost part
of a culm bearing an inflorescence; 2) the
stalk of a raceme or cluster of spikelets.

penicillate: with a tuft of fine hairs at the tip.

rhizome: an underground stem (see stolon
and Fig. 3, p. 5).

rhizomatous: with a rhizome.

rosette: a spreading and radiating basal clus-
ter of leaves.

418

rugose: wrinkled and ridged, e.g. the female-
fertile lemma of Panicum maximum.

saccate: bag- or sac-shaped, as in the lower
lemma of Sacciolepis (a solid shape).

sagittate: arrowhead-shaped; with down-
ward pointing acute lobes at the base (a
solid shape).

scaberulous: minutely scabrous.

scabrid, scabrous: rough to the touch, with
minute teeth or scattered short broad-
based harsh hairs.

scandent: scrambling, often on other plants.

scarious: thin, dry and shrivelled, not green.

secund: one-sided, or arranged on one side.

sensu lato: a Latin phrase meaning

‘interpreted in a broad sense’ (abbrevi-
ated: s.l.). For example, Melinis s.l.
means all the species assigned to the
genus, including those formerly classified
in Rhynchelytrum (see sensu stricto).

sensu stricto: a Latin phrase meaning
‘interpreted in a narrow sense’ (abbrevi-
ated: 5.5.). For example, Melinis s.s.
means the species assigned to the genus
before Rhynchelytrum was included in it
(see sensu lato).

sessile: without a pedicel (see pedicellate).

setaceous: bristle-like.

setae: bristles.

sheath: the basal part of a grass leaf that is
normally wrapped around the culm (see
Fig 5, p. 7).

sinus: the angle between two lobes.

spathe: a bract or bladeless leaf sheathing
the inflorescence or a major component
of it.

spatheole: a secondary spathe within a
compound inflorescence in the Andropo-
goneae.

spike: an inflorescence in which a single axis
bears sessile spikelets (see Fig. 6, p. 9).

spikelet: the basic unit of a grass

inflorescence, composed of glumes,
rachilla and florets (see Fig. 7, p. 10 and
Fig. 8, p. 11).

spreading: held outward, at about right
angles to the main axis.

stamen: the pollen-bearing part of a flower,
usually composed of filament and anther
(see Fig. 8, p. 11).

staminate: bearing stamens only and no
pistils; the term may be applied to a
flower, a floret, a spikelet, an inflores-
cence or an entire plant (see pistillate).

sterile: without functional male or female
parts. An ambiguous term, in the past
sometimes used to mean ‘not producing
seed or pollen’ (Chippindall 1955) and
sometimes meaning ‘without pistils. ..may
be staminate or neuter’ (Hitchcock &
Chase 1950), thus leaving male fertility
in doubt (see female-fertile).

stigma: the part of the pistil that receives the
pollen (see Fig. 8, p. 11).

stipe: a stalk to an organ that is part of the
organ itself and not a separate branch.

stipitate: with a stipe.

stolon: a stem that creeps above the ground,
roots and gives rise to new plants (see
rhizome and Fig. 3, p. 5).

stoloniferous: with a stolon.

striate: with fine parallel lines or ridges.

styles: branches of the pistil that bear the
stigmas (see Fig. 8, p. 11).

sub-: slightly or somewhat less than.

subulate: awl-shaped; tapering from base to
apex and usually sharp-pointed (a solid
shape).

sulcate: with a groove or furrow.

sward: lawn; continuous grass cover

produced by stoloniferous species.

taxon: an invented term that signifies any
taxonomic group irrespective of its
classification level.

terete: cylindric and slender.

tiller: a leafy side-branch from a main culm,
which may eventually flower, sometimes
beginning in one year and flowering in
the second year.

tomentose: hairy with somewhat matted,
curly, wooly hairs appressed to the
surface.

triad: three spikelets borne together (see PI.
38, p. 361 and PI. 209, p. 380).

trifid: 3-branched.

trigonous: 3-sided, with the sides convex (a
solid shape).

triquetrous: 3-sided, with the sides concave
(a solid shape).

truncate: ending abruptly as if cut off.

tuberculate: a surface with small

projections.

tussock: a dense tuft (Afrikaans: pol).

utricle: a bladdery fruit in which the seed
coat is separate from the pericarp; true
utricles may not occur in Poaceae, but

utricle-like caryopses occur mainly in
Sporobolus, Eleusine and a few other
chloridoids (see caryopsis).

vascularization: the pattern of vascular
tissue (veins, nerves, xylem and phloem)
in an organ.

ventral: an ambiguous term for the front,
adaxial or inner surface of an organ (see
dorsal). The term adaxial is preferable.

verrucose: a surface with warts or nodules.

villous: hairy with moderately erect, dense,
long, soft, often curly hairs.

viscid: sticky.

viviparous: young plantlets being produced
within the parental inflorescence.

whorled: with several branches arising from
a single node.

winged: with a thin projection or border.

woolly: hairy with dense, long, soft,

entangled, curled hairs not appressed to
the surface.

xerophytic: growing in arid places (see
helophytic, mesophytic).

XyMS+: a code that signifies the presence of
mestome sheath cells between the large
metaxylem elements and the PCR sheath
cells in primary vascular bundles (see
XyMS -, as well as Fig. 1 and pp. 2-4 for
particulars).

XyMS-: a code that signifies the absence of
mestome sheath cells between the large
metaxylem elements and the PCR sheath
cells in primary vascular bundles (see
XyMS+, as well as Fig. 1 and pp. 2-4 for
particulars).

421

INDEX TO SCIENTIFIC NAMES, SYNONYMS AND COMMON NAMES

Correct scientific names for southern African taxa are
printed in bold type, as is the page number on which the
treatment of each taxon appears. Synonyms are printed in
italic type. Vernacular names, names of taxa that do not
occur in southern Africa and names with an undecided
synonymy are printed in mild roman type.

Abyssinian finger grass 107

Achneria capensis (Steud.) Dur. & Schinz 261

Acrachne Chiov., 29, 31, 385

Acrachne racemosa (Roem. & Schult.) Ohwi 31 (fig. 10),
357 (pi. 1)

Acrachne verticillata (Roxb.) Chiov. 31
Acroceras Stapf 29, 32, 386, 387
Acroceras macrum Stapf 32 (fig. 1 1), 357 (pi. 2)
Acroceras pilgeranum Schweick. 241
African finger millet 129
Agropyron Gaertn. 132, 181, 275, 292
Agropyron distichum (Thunb.) Beauv. 336
Agropyron repens (L.) Beauv. vii, 132
Agrostis L. 10,12 29, 32, 77, 275, 382, 392
Agrostis avenacea Gmel. 34
Agrostis barbuligera Stapf
var. barbuligera 34

var. longipilosa Goossens & Papendorf 34
Agrostis bergiana Trin.
var. bergiana 34
var. laevisulca Stapf 34
Agrostis continuata Stapf 34
Agrostis eriantha Hack,
var. eriantha 33 (fig. 12), 34, 357 (pi. 3)
var. planifolia Goossens & Papendorf 34
Agrostis gigantea Roth 34
Agrostis griquensis Stapf 275
Agrostis huttoniae (Hack.) C.E. Hubb. 34
Agrostis lachnantha Nees
var. lachnantha 34

var. glabra Goossens & Papendorf 34, 35
Agrostis montevidensis Spreng. ex Nees 35
Agrostis natalensis Stapf 34
Agrostis polypogonoides Stapf 35
Agrostis schimperiana Steud. 34
Agrostis schlechteri Rendle 35
Agrostis semiverticillata (Forssk.) C. Christ. 276
Agrostis subulifolia Stapf 35
Agtdaepluimgras 154
Aira L. vii, 29, 35, 382
Aira carophyllea L. 36

Aira cupaniana Guss. 35 (fig. 13), 36, 265, 357 (pi. 4)
Alloteropsis Presl 2, 29, 36, 386, 387
Alloteropsis cimicina (L.) Stapf 37
Alloteropsis papillosa Clayton 37
Alloteropsis semialata auctt., non Gibbs Russell 37
Alloteropsis semialata (R. Br.) Hitchc.
subsp. eckloniana (Nees) Gibbs Russell 2, 37, 357 (pis.
5 & 6)

subsp. semialata 2, 7, 36 (fig. 14), 37, 416
var. ecklonii (Stapf) Stapf 37
Ammophila Host. 8, 29, 38, 382

Ammophila arenaria (L.) Link vii, 1, 38 (fig. 15), 77, 357
(pi. 7)

Andropogon L. 5, 10, 30, 38, 387
Andropogon abyssinicus Fresen. 40

Andropogon abyssinicus sensu Chippind., non Fresen. 40

Andropogon amethystinus Steud. 40
Andropogon amplectens Nees 1 15

Andropogon appendiculatus Nees 40

Andropogon brazzae Franch. 40

Andropogon chinensis (Nees) Merr. 39 (fig. 16), 40, 357
(pi. 8)

Andropogon distachyos L. 40, 41
Andropogon eucomus Nees 40, 41
Andropogon fastigiatus Swartz 41, 223
Andropogon festuciformis Rendle 41
Andropogon filifolius (Nees) Steud. 1 15
Andropogon gayanus Kunth
var. polycladus (Hack.) Clayton 41
var. squamulatus (Hochst.) Stapf 41
Andropogon huillensis Rendle 40, 41
Andropogon lacunosus J.G. Anders. 41
Andropogon laxatus Stapf 40, 41
Andropogon mannii Hook. f. 41
Andropogon pilosellus Stapf 40
Andropogon platybasis J.G. Anders. 41
Andropogon ravus J.G. Anders. 6, 42
Andropogon schinzii Hack. 40
Andropogon schirensis A. Rich. 42
var. angustifolius Stapf 42
Annual bluegrass 271
Annual bristle grass 47
Annual swamp grass 285
Annual wool grass 43

Anthephora Schreb. 10, 29, 42, 331, 386, 416
Anthephora angustifolia Goossens 43

Anthephora argentea Goossens 43, 131
Anthephora pubescens Nees 42 (fig. 17), 43
Anthephora ramosa Goossens 43, 357 (pi. 9)
Anthephora schinzii Hack. 43, 358 (pi. 10)
Anthoxanthum L. 29, 43, 381, 382
Anthoxanthum brevifolium Stapf 44
Anthoxanthum dregeanum (Nees) Stapf 44, 45
Anthoxanthum ecklonii (Nees ex Trin.) Stapf 44 (fig. 18),
358 (pi. 11)

Anthoxanthum odoratum L. 44, 45
Anthoxanthum tongo (Trin.) Stapf 45
Apochaete 347

Apochaete hispida (L. f.) J.B. Phipps 348
Aristida L. iii, 2, 3, 4, 9, 29, 45, 52, 286, 319, 320, 385,
410,414

Aristida adscensionis L. 47, 50, 358 (pi. 12)
subsp. guineensis (Trin. & Rupr.) Henr. 47
Aristida aequiglumis Hack. 48, 51
Aristida alopecuroides Hack. 49
Aristida amabilis Schweick. 320
Aristida andoniensis Henr. 53
Aristida angolensis C.E. Hubb. 287
Aristida argentea Schweick. 5 1
Aristida barbie ollis Trin. & Rupr. 48
Aristida bipartita (Nees) Trin. & Rupr. 48, 50, 53
Aristida brevifolia (Nees) Steud. 321
Aristida canescens Henr.
subsp. canescens 48, 52, 53
subsp. ramosa De Winter 48
Aristida capensis Thunb. 329
var. barbata Stapf 328
var. canescens Trin. & Rupr. 329
var. dieterleniana Schweick. 329
var. genuina Henr. 329
var. macropus (Nees) Trin. & Rupr. 329
Aristida ciliata Desf. 321
var. capensis Trin. & Rupr. 321
var. pectinata Henr. 321
var. tricholaena Hack. 321
var. villosa Hack. 321

422

Aristida ciliata sensu Desf., non Steud. & Hochst. ex Steud.

321

Aristida congesta Roem. & Schult.
subsp. barbicollis (Trin. & Rupr.) De Winter 47, 48
subsp. congesta 45 (fig. 19), 47, 49, 50
Aristida curvata (Nees) Dur. & Schinz 47
Aristida damarensis Mez 321
Aristida dasydesmis (Pilg.) Mez 49, 51
Aristida dewinteri Giess 49
Aristida diffusa Trin.
subsp. burkei (Stapf) Meld. 49, 50, 55
subsp. diffusa 49, 50, 55
var. burkei (Stapf) Schweick. 49
var. genuina Henr. 49

var . pseudo-hystrix (Trin. & Rupr.) Henr. 49
Aristida dinteri Hack. 322
Aristida dregeana (Nees) Trin. & Rupr. 322
Aristida effusa Henr. 48, 50, 53
Aristida engleri Mez
var. engleri 50, 55
var. ramosissima De Winter 50
Aristida fastigiata Hack. 322
Aristida fontismagni Schweick. 54
Aristida galpinii Stapf 50
Aristida garubensis Pilg. 322
Aristida geminifolia (Nees) Trin. & Rupr. 322
Aristida gonatostachys Pilg. 323
Aristida graciliflora Pilg. 53
Aristida gracilior Pilg.
var. gracilior 323
var. intermedia Schweick. 327
var. pearsonii Henr. 324
Aristida hermannii Mez 323
Aristida hirtigluma Steud. ex Trin. & Rupr. 323
Aristida hochstetteriana Beck ex Hack. 324
Aristida hordeacea Kunth 49, 50
Aristida hubbardiana Schweick. 47, 49, 50
Aristida jucunda Schweick. 287
Aristida junciformis Trin. & Rupr.
subsp. galpinii (Stapf) De Winter 48, 50, 52
subsp. junciformis 6, 48, 49, 51, 52, 55
subsp. welwitschii (Rendle) Meld. 51
Aristida lanipes Mez 324
Aristida longicauda Hack. & Henriques 49
Aristida lutescens (Nees) Trin. & Rupr. 324
Aristida marlothii Hack. 324
Aristida meridionalis Henr. 51 (fig. 20), 53, 54
Aristida mollissima Pilg.
subsp. argentea (Schweick.) Meld. 51, 54
subsp. mollissima 52, 54
Aristida monticola Henr. 52, 55
Aristida namaquensis (Nees) Trin. & Rupr. 325
Aristida obtusa Del. 326
Aristida parvula (Nees) De Winter 52
Aristida pilgeri Henr. 48, 52, 53
Aristida proximo Steud. 326
Aristida recta Franch. 52
Aristida rhiniochloa Hochst. 53
Aristida sabulicola Pilg. 326
Aristida scabrivalvis Hack,
subsp. borumensis (Henr.) Meld. 53
subsp. contracta (De Winter) Meld. 48, 50, 53
subsp. scabrivalvis 48, 50, 53
Aristida schaeferi Mez
var. biseriata Henr. 327
var. schaeferi 327
Aristida schlechteri Henr. 329
Aristida sciurus Stapf 48, 52, 53
Aristida secalina Henr. 324
Aristida sericans Hack, apud Schinz 329
Aristida spectabilis Hack. 51, 53
Aristida stipitata Hack,
subsp. graciliflora (Pilg.) Meld. 52, 53
subsp. robusta (Stent & Rattray) Meld. 54
subsp. spicata (De Winter) Meld. 54
subsp. stipitata 12, 52, 54
var. graciliflora (Pilg.) De Winter 53

Aristida stipoides Lam. 51, 54
Aristida subacaulis Nees ex Steud. 327
Aristida submucronata Schumach. 47
Aristida transvaalensis Henr. 49, 51, 52, 54
Aristida uniplumis Licht.
var. neesii Trin. & Rupr. 327
var. pearsonii Henr. 328
var. uniplumis 328

Aristida vestita Thunb. 49, 50, 51, 55
Aristida wildii Meld. 50
Arrhenatherum Beauv. vii, 29, 55, 58, 382
Arrhenatherum elatius (L.) Presl 55 (fig. 21 ), 358 (pi. 13)
var. biaristatum (Peterm.) Peterm. 55
var. bulbosum (Willd.) Spenner 55
Arrow grass 298
Arthraxon Beauv. 30, 56, 387
Arthraxon lanceolatus (Roxb.) Hochst.

var. lanceolatus 56 (fig. 22), 358 (pi. 14)

Arthraxon prionodes (Steud.) Dandy 56

Arundinaria tessellata (Nees) Munro 333

Arundinella Raddi 3, 29, 56, 387

Arundinella nepalensis Trin. 57 (fig. 23), 358 (pi. 15)

Arundo L. 29, 57, 384, 399

Arundo donax L. 5, 7, 58 (fig. 24), 358 (pi. 16)

Assegaaigras 179

Asternatherum Nevski 80

Asthenatherum forskahlei auctt., non (Vahl) Nevski 81
Asthenatherum glaucum (Nees) Nevski 81
Asthenatherum mossamedense (Rendle) Conert 8 1
Avena L. vii, 4, 10, 29, 55, 58, 382
Avena barbata Brot. 58, 358 (pi. 17)

Avena byzantina K. Koch. 59
Avena fatua L. 59 (fig. 25)

Avena sativa L. 1, 59
subsp. macrantha Hack. 59
subsp. praegravis Krause 59
subsp. sativa 59
Avena sterilis L. 59

subsp. ludoviciana (Dur.) Gillet & Magne 59
subsp. sterilis 59
Avena strigosa Schreb. 59
Axonopus Beauv. 29, 59, 386
Axonopus affinis Chase 60 (fig. 26), 358 (pi. 18)
Axonopus compressus sensu Chippind., non (Swartz)
Beauv. 60

Bahia grass 246

Bahia paspalum 246

Bambusa Schreber 14, 29, 60, 383

Bambusa balcooa Roxb. ex Roxb. 61, 359 (pi. 19)

Bana grass 250
Bankrotkweek 246
Barley 1

Barnyard millet 120

Bastersinjaalgras 65

Batavian quick grass 178

Bearded thatching grass 186

Beckeropsis Fig. & De Not. 5, 247, 250

Beckeropsis uniseta (Nees) K. Schum. 250

Beddinggras 249

Beesgras 57

Bent grass 34

Berggras 347

Bergpluimgras 161

Bergsetaria 296

Bergsteekgras 53

Besemeragrostis 157

Besemgras 155, 207, 221, 346, 348

Bewsia Goossens 29, 61, 385

Bewsia biflora (Hack.) Goossens 61 (fig. 27), 359 (pi. 20)

Big quaking grass 72

Bitter turpentine grass 95

Blackseed finger grass 1 13

Black wild sorghum 302

Blinkaarboesmangras 328

Blougras 40

Blouklosgras 62

423

Blousaadgras 61, 162, 241, 309, 342

Blousaadsoetgras 239

Bloutamboekiegras 187

Blown grass 34

Blue couch 1 10

Blue grama 105

Boat thatching grass 184, 187

Bokbaardgras 169

Bootjietamboekiegras 184

Borseltjiegras 43

Bosbuffelsgras 235

Bosluisgras 150

Bothriochloa Kuntze 1 1, 30, 62, 105, 387, 391, 393
Bothriochloa bladhii (Retz.) S.T. Blake 9, 62
Bothriochloa glabra (Roxb.) A. Camus 62
Bothriochloa insculpta (A. Rich.) A. Camus 11, 62 (fig.
28), 63, 359 (pi. 21)
var. vegetior (Hack.) C.E. Hubb. 62
Bothriochloa pertusa auctt., non (L.) A. Camus 63
Bothriochloa radicans (Lehm.) A. Camus 63
Bottlebrush grass 266

Brachiaria (Trin.) Griseb. 2, 5, 10, 12, 29, 63, 1 10, 386
Brachiaria advena Vickery 64, 66, 67
Brachiaria arrecta (Dur. & Schinz) Stent 65, 68
Brachiaria bovonei (Chiov.) Robyns 65, 68
Brachiaria brizantha (A. Rich.) Stapf 65, 359 (pi. 22)
Brachiaria chusqueoides (Hack.) Clayton 65, 66, 240
Brachiaria deflexa (Schumach.) C.E. Hubb. ex Robyns 65,
66, 68 (fig. 30), 359 (pi. 23)

Brachiaria dictyoneura (Fig. & De Not.) Stapf 65, 67
Brachiaria dura Stapf
var. dura 66, 68
var. pilosa J.G. Anders. 66
Brachiaria eruciformis (J.E. Sm.) Griseb. 65, 66, 67
Brachiaria filifolia Stapf 68
Brachiaria glomerata (Hack.) A. Camus 66, 67
Brachiaria grossa Stapf 65, 66
Brachiaria humidicola (Rendle) Schweick. 66
Brachiaria latifolia Stapf 65

Brachiaria malacodes (Mez & K. Schum.) Scholz 65, 67
Brachiaria marlothii (Hack.) Stent 6, 67
Brachiaria nigropedata (Fical. & Hiern) Stapf 67
Brachiaria poaeoides Stapf 67

Brachiaria psammophila (Welw. ex Rendle) Launert 66,
67

Brachiaria rugulosa Stapf 65

Brachiaria schoenfelderi C.E. Hubb. & Schweick. 67
Brachiaria serrata (Thunb.) Stapf 5, 7, 9, 63 (fig. 29), 67,
359 (pi. 24)

var. gossypina (A. Rich.) Stapf 67, 68
var. serrata 67

Brachiaria subulifolia (Mez) Clayton 65. 68
Brachiaria xantholeuca (Schinz) Stapf 66, 68
Brachyachne (Benth.) Stapf 29, 68, 385
Brachyachne patentiflora (Stent & Rattray) C.E. Hubb. 69
(fig. 31), 359 (pi. 25)

Brachychloa S.M. Phillips 29, 69, 385
Brachychloa fragilis S.M. Phillips 70
Brachychloa schiemanniana (Schweick.) S.M. Phillips 70
(fig. 32), 359 (pi. 26)

Brachypodium Beauv. 29, 70, 382

Brachypodium bolusii Stapf 71

Brachypodium distachyon (L.) Beauv. 71

Brachypodium flexum Nees 71 (fig. 33), 359 (pi. 27)

Bradley grass 96

Brak paspalum 247

Brakvleigras 309

Breeblaar andropogon 1 15

Breeblaar polgras 297

Bristle grass 295

Bristle three-awn 50

Briza L. 29, 72, 382, 399

Briza maxima L. 72, 360 (pi. 28)

Briza minor L. 72 (fig. 34)

Briza subaristatum Lam. 73, 360 (pi. 29)

Briza triloba Nees 73

Broad-leaved bluestem 115
Broad-leaved panicum 236
Bromus L. 7, 10, 1 1, 29, 73, 168, 382

Bromus alopecurus Poir. 74

Bromus catharticus Vahl 7, 9, 73 (fig. 35), 74, 360 (pi
30)

Bromus commutatus Schrad. 74
Bromus diandrus Roth 74, 75, 76
Bromus firmior (Nees) Stapf 75, 76
var .firmior 75
var. leiorhachis Stapf 75
Bromus hordeaceus L.
subsp. ferronii (Mabille) P.M. Sm. 75
subsp. molliformis (J. Lloyd) Maire & Weiller 75
Bromus inermis Leyss. 75
Bromus japonicus sensu Chippind., non Thunb.
var .japonicus 76

var. velutinus (Nocc.) Aschers. & Graebn. 76
Bromus leptoclados Nees 75
Bromus madritensis L. 75
Bromus molliformis Lloyd 75
Bromus natalensis Stapf 75, 76
var. lasiophilus Stapf 75
Bromus pectinatus Thunb. 76
Bromus rigidus Roth 74, 76
Bromus rubens L. 76
Bromus speciosus Nees 75, 76
Bromus speciosus sensu Compton, non Nees 75
Bromus tectorum L. 76
Bromus unioloides H.B.K. 74
Bromus willdenowii Kunth 74
Bronsaartamboekiegras 186
Bronze awned thatching grass 186
Broodsinjaalgras 65
Broom trident grass 348
Brown finger grass 108
Brown seed finger grass 109
Bruinhoenderspoor 168
Bruinsaadgras 147
Buchugras 95
Buffalo grass 314
Buffelsgras 80
Bulbous bluegrass 271
Bulgras 250
Bur bristle grass 300
Bushman grass 329

Calamagrostis Adans. 29, 33, 34, 76, 382, 392
Calamagrostis epigeios (L.) Roth
var. capensis Stapf 38, 77 (fig. 36), 360 (pi. 3 1 )

Cape bushman grass 328
Cape cord grass 303
Carpet grass 60

Catabrosia aquatica auctt., non (L.) Beauv. 90
Catalepis Stapf & Stent 29, 77, 385
Catalepis gracilis Stapf & Stent 78 (fig. 37), 360 (pi. 32)
Catapodium Link 29, 78, 382

Catapodium rigidum (L.) C.E. Hubb. 78 (fig. 38), 79, 360
(pl. 33)

Caterpillar grass 174
Catstail grass 116, 311
Cenchrus L. 5, 10, 29, 79, 247, 386
Cenchrus biflorus Roxb. 79
Cenchrus brownii Roem. & Schult. 12, 80
Cenchrus ciliaris L. 4, 5, 79 (fig. 39), 80, 249, 360 (pl. 34)
Cenchrus incertus M.A. Curtis 79, 80
Cenchrus pauciflorus Benth. 79, 80
Centipede grass 349
Centropodia Reichenb. 29, 80, 384
Centropodia glauca (Nees) T.A. Cope 2, 6, 80 (fig. 40),
81, 360 (pl. 35)

Centropodia mossamedensis (Rendle) T.A. Cope 81
Chaetobromus Nees 29, 81, 384, 399, 400
Chaetobromus dregeanus Nees 81 (fig. 41), 82, 360 (pl
36)

Chaetobromus involucratus (Schrad.) Nees 82

424

Chascolytrum Desv. 72

Chascolytrum subaristatum (Lam.) Desv. 73

Chilean love grass 163

Chloris O. Swartz 10, 29, 82, 96, 137, 168, 315, 385

Chloris diluta Renvoize 83, 84

Chloris flabellata (Hack.) Launert 83

Chloris gayana Kunth 84

Chloris mossambicensis K. Schum. 84

Chloris myriostachya Hochst. 84

Chloris prieUrii Kunth 137

Chloris pycnothrix Trin. 7, 83 (fig. 42), 84

Chloris roxburghiana Schult. 84

Chloris transiensis Pilg. 291

Chloris truncata R. Br. 83, 84

Chloris virgata Swartz 84, 361 (pi. 37)

Chrysopogon Trin. 30, 85, 387
Chrysopogon montanus Trin.

var. tremulus (Hack.) Stapf 85
Chrysopogon serrulatus Trin. 9, 85 (fig. 43), 361 (pi. 38)
Cladoraphis Franch. 29, 85, 385
Cladoraphis cyperoides (Thunb.) S.M. Phillips 86
Cladoraphis spinosa (L. f.) S.M. Phillips 7, 86 (fig. 44),

361 (pi. 39)

Cleistachne Benth. 30, 86, 387

Cleistachne sorghoides Benth. 87 (fig. 45), 361 (pi. 40)
Coast finger grass 1 12
Coastal wheat grass 336
Cocksfoot 99

Coelachyrum Hochst. & Nees 29, 87, 385
Coelachyrum yemenicum (Schweinf.) S.M. Phillips 87
(fig. 46), 88, 361 (pi. 41)

Coelorachis Brongn. 30, 88, 388

Coelorachis capensis Stapf 88 (fig. 47), 361 (pi. 42)

Coix L. 30, 89, 388, 399

Coix lacryma-jobi L. vii, 89 (fig. 48), 361 (pi. 43)
Colpodium Trin. 29, 89, 382, 393

Colpodium hedbergii (Meld.) Tzvel. 90 (fig. 49), 361 (pi.
44)

Common Canary grass 269
Common finger grass 1 10
Common russet grass 207
Common signal grass 65
Common thatching grass 185
Common turpentine grass 95
Common wild-sorghum 302
Cortaderia Stapf vii, 29, 90, 384, 399
Cortaderia jubata (Lem.) Stapf 10, 90
Cortaderia selloana (Schult.) Aschers. & Graebn. 1, 7, 91
(fig. 50), 361 (pi. 45)

Corynephorus Beauv. 14, 29, 91, 382, 399
Corynephorus divaricatus (Pourret) Breistr. 92
Corynephorus fasciculatus Boiss. & Reut. 91 (fig. 51), 92,

362 (pi. 46)

Cottonwool grass 190

Couch grass 97

Couch panicum 241

Couch paspalum 246

Crab finger grass 1 13

Craspedorhachis Benth. 14, 29, 92, 385

Craspedorhachis africana Benth. 92, 93, 362 (pi. 47)

Craspedorhachis rhodesiana Rendle 92 (fig. 52)

Craspedorhachis sarmentosa (Hack.) Pilg. 356

Craspedorhachis uniflora (Hochst. ex A. Rich.) Chippind.

199

Creeping carrot-seed grass 338
Creeping paspalum 246
Creeping signal grass 66
Crinipes Hochst. 330
Crinipes gynoglossa Goossens 330
Critesion Raf. 180

Critesion marinum (Huds.) Loeve 181
Critesion murinum (L.) Loeve 182
Critesion stenostachys (Godr.) Loeve 1 82
Crowfoot 100

Ctenium Panzer 29, 93, 385

Ctenium concinnum Nees 93 (fig. 53), 362 (pi. 48)

Curly leaf 1 59

Curly-leaved three-awned grass 48
Cymbopogon Spreng. 7, 9, 1 1, 94
Cymhopogon afronardus Stapf 95
Cymbopogon caesius (Hook. & Arn.) Stapf 95
Cymbopogon dieterlenii Stapf ex Phill. 95
Cymbopogon excavatus (Hochst.) Stapf ex Burtt Davy 7,
94 (fig. 54), 95

Cymbopogon giganteus Chiov. 95

Cymbopogon marginatus (Steud.) Stapf ex Burtt Davy 95,
362 (pi. 49)

Cymbopogon nardus (L.) Rendle 95
Cymbopogon plurinodis (Stapf) Stapf ex Burtt Davy 2, 6,
7, 95

Cymbopogon pospischilii (K. Schum.) C.E. Hubb. 95
Cymbopogon prolixus (Stapf) Phill. 95
Cymbopogon validus (Stapf) Stapf ex Burtt Davy 95
Cymbosetaria Schweick. 5, 293, 294, 386
Cymbosetaria sagittifolia (A. Rich.) Schweick. 298
Cynodon Rich. 9, 10, 29, 83, 95, 97, 385
Cynodon aethiopicus Clayton & Harlan 96, 97
Cynodon bradleyi Stent 96, 97

Cynodon dactvlon (L.) Pers. 1, 5, 6, 7, 9, 96 (fig. 55), 97,
‘ 362 (pi. 50)

Cynodon hirsutus Stent 97
Cynodon incompletus Nees 97
Cynodon nlemfuensis Vanderyst 96, 97
Cynodon plectostachyus (K. Schum.) Pilg. 97
Cynodon polevansii Stent 97
Cynodon transvaalensis Burtt Davy 97
Cynosurus L. 29, 98, 382
Cynosurus aureus L. 196

Cynosurus coloratus Lehm. ex Nees 98, 362 (pi. 51)
Cynosurus echinatus L. vii, 98 (fig. 56)

Cypholepis yemenica (Schweinf.) Chiov. 88

Daba grass 221

Dactylis L. 29, 99, 382, 391

Dactylis glomerata L. 99 (fig. 57), 362 (pi. 52)

Dactyloctenium Willd. 29, 99, 385

Dactyloctenium aegyptium (L.) Willd. 100

Dactyloctenium australe Steud. 100

Dactyloctenium geminatum Hack. 100

Dactyloctenium giganteum Fisher & Schweick. 100 (fig.

58), 101, 362 (pi. 53)

Dallis grass 246

Danthonia DC. 5, 1 18, 213, 278
Danthonia arundinacea (Berg.) Schweick. 215
Danthonia aureocephala J.G. Anders. 215
Danthonia brachyphylla Stapf 278
Danthonia cincta Nees 215
Danthonia curva Nees 193
Danthonia davyi C.E. Hubb. 215
Danthonia disticha Nees 216
Danthonia drakensbergensis Schweick. 216
Danthonia dura Stapf 216
Danthonia glauca Nees 81
Danthonia lanata (Schrad.) Schrad.
var. lanata 217
var. maior Nees 217

Danthonia lupulina (Thunb.) Beauv. ex Roem. & Schult.
217

Danthonia macowanii Stapf 2 1 7
Danthonia macrantha Schrad. 279
Danthonia macrocephala Stapf 217
Danthonia mossamedensis Rendle 8 1
Danthonia papposa Nees 217
Danthonia pumila Nees 118

Danthonia purpurea (Thunb.) Beauv. ex Roem. & Schult.
193

Danthonia rangei Pilg. 217
Danthonia stereophylla J.G. Anders. 218
Danthonia stricta Schrad. 218
Danthonia tenella Nees 193
Danthonia zeyheriana Steud.
var. trichostachya Stapf 215
var. zeyheriana 215

425

Danthoniopsis Stapf 29, 101, 387

Danthoniopsis chimanimaniensis (J.B. Phipps) Clayton 102
Danthoniopsis dinteri (Pilg.) C.E. Hubb. 101 (fig. 59),
102, 362 (pi. 54)

Danthoniopsis lignosa C.E. Hubb. 102
Danthoniopsis parva (J.B. Phipps) Clayton 102
Danthoniopsis pruinosa C.E. Hubb. 102
Danthoniopsis ramosa (Stapf) Clayton 102
Danthoniopsis scopulorum (J.B. Phipps) J.B. Phipps 102
Darnel 203
Dekgras 293
Dektamboekiegras 185
Deschampsia Beauv. vii, 29, 103, 265, 382
Deschampsia cespitosa (L.) Beauv. vii, 103 (fig. 60), 363
(pi. 55)

Deschampsia flexuosa (L.) Trin. 103
Deschampsia flexuosa (L.) Trin.

var. afromontana C.E. Hubb. 103
Desmazeria Dumort. 78
Desmazeria rigida (L.) C.E. Hubb. 79
Diandrochloa De Winter 29, 104, 385
Diandrochloa namaquensis (Nees) De Winter 11, 104
(fig. 61 ), 363 (pi. 56)

Diandrochloa pusilla (Hack.) De Winter 104
Dichanthium Willemet 9, 30, 62, 105, 387, 391
Dichanthium annulatum (Forssk.) Stapf
var. papillosum (A. Rich.) De Wet & Harlan 105 (fig. 62),
363 (pi. 57)

Dichanthium aristatum (Poir.) C.E. Hubb. 105
Dichanthium nodosum sensu Acocks, non Willemet 105
Dichanthium papillosum (Hochst.) Stapf 105
Diectomis Beauv. 5

Diectomis fastigiata (Swartz) Kunth 41
Digitaria Haller iii, vii, 7, 9, 10, 29, 60, 106, 208, 3 15, 33 1,
386

Digitaria abyssinica (A. Rich.) Stapf 107
Digitaria acuminatissima Stapf 107
subsp. inermis Goetghebeur 107
Digitaria adscendens Henr. 108
Digitaria alhomarginata Stent 1 10
Digitaria angolensis Rendle 108
Digitaria apiculata Stent 1 1 1
Digitaria argyrograpta (Nees) Stapf 108
Digitaria bechuanica (Stent) Henr. 1 10
Digitaria borbonica Desv. 112
Digitaria brazzae (Franch.) Stapf 108
Digitaria ciliaris (Retz.) Koeler 108
Digitaria comifera Pilg. 109
Digitaria debilis (Desf.) Willd. 6, 108 (fig. 64), 109
Digitaria decumbens Stent 1 10
Digitaria diagonalis (Nees) Stapf
var. diagonalis 109 (fig. 65), 110
Digitaria didactyla Willd. 110, 111
Digitaria dinteri Henr. 1 10
Digitaria diversinervis (Nees) Stapf 110
var. woodiana Henr. 1 10

Digitaria eriantha Steud. 1, 4, 6, 9, 106 (fig. 63), 110, 111,
112, 113, 363 (pi. 58)
subsp. pentzii (Stent) Kok 1 10
subsp. stolonifera (Stapf) Kok 1 10
subsp. transvaalensis Kok 1 10
var. stolonifera Stapf 1 10
Digitaria eylesii C.E. Hubb. 109, 110
Digitaria flaccida Stapf 110
Digitaria gayana (Kunth) Stapf 110
Digitaria gazensis Rendle 111, 112
Digitaria geniculata Stent 1 10
Digitaria glauca sensu Stent, non A. Camus 110
Digitaria gymnostachys Pilg. Ill
Digitaria littoralis sensu Stent, non Salisb. 1 12
var .prostrata Stent 112
Digitaria longiflora (Retz.) Pers. Ill
Digitaria macroglossa Henr. 1 1 2
var. prostrata (Stent) Henr. 1 12
Digitaria maitlandii Stapf & C.E. Hubb. Ill
Digitaria maniculata Stapf 111

Digitaria marginata Link 108
Digitaria melanochila Stapf 1 1 3

Digitaria milanjiana (Rendle) Stapf 111
Digitaria monodactyla (Nees) Stapf 111, 131, 363 (pi. 59)
var. explicata Stapf 1 1 1
Digitaria natalensis Stent 112
subsp. stentiana Henr. 1 12
Digitaria nuda Schumach. 112
Digitaria pentzii Stent 1 10
var. stolonifera (Stapf) Henr. 1 10
Digitaria perrottetii (Kunth) Stapf 112
Digitaria polevansii Stent 1 13
Digitaria polyphylla Henr. 112
Digitaria remotigluma (De Winter) Clayton 112
Digitaria rigida Stent 112
Digitaria rukwae Clayton 111, 112
Digitaria sanguinalis (L.) Scop. 6, 106, 113 (fig. 66)
Digitaria scalarum (Schweinf.) Chiov. 107
Digitaria seriata Stapf 113
Digitaria setifolia Stapf 113
Digitaria setivalva Stent 1 10
Digitaria smutsii Stent 1 10
Digitaria stentiana Henr. 110
Digitaria swazilandensis Stent 110
Digitaria ternata (A. Rich.) Stapf 113
Digitaria thouaresiana (Fluegge) A. Camus 113
Digitaria tricholaenoides Stapf 7, 114 (fig. 67)

Digitaria trichopodia Stent 109
Digitaria tricostulata (Hack.) Henr. 1 13
Digitaria uniglumis (A. Rich.) Stapf 109
Digitaria valida Stent 1 10
var. glauca Stent 1 10
Digitaria velutina (Forssk.) Beauv. 114
Digitaria vestita Fig. & De Not.

var. scalarum (Schweinf.) Henr. 107
Digitaria violascens Link. 1 14
Digitaria zeyheri (Nees) Henr. 1 14
Digitariella De Winter 106
Digitariella remotigluma De Winter 1 12
Diheteropogon Stapf 12, 30, 114, 387
Diheteropogon amplectens (Nees) Clayton 7, 42, 95, 115
(fig. 68), 363 (pi. 60)

Diheteropogon filifolius (Nees) Clayton 115
Dinebra Jacq. 29, 116, 385
Dinebra retroflexa (Vahl) Panz.

var. condensata S.M. Phillips 116 (fig. 69), 363 (pi. 61)
Diplachne Beauv. 29, 116, 385, 391
Diplachne biflora Hack, ex Schinz 61
Diplachne cinerea Hack. 226
Diplachne cuspidata Launert 117, 118
Diplachne eleusine Nees 117

Diplachne fusca (L.) Beauv. ex Roem. & Schult. 7, 117
(fig. 70), 118, 363 (pi. 62)

Diplachne gigantea Launert 118
Diplachne malabarica (L.) Merr. sensu Adamson 1 17
Disc dropseed 308
Dogstail 98

Dolichochaete rehmannii (Hack.) J.B. Phipps 348
Donsgras 190
Draadbloustam 115
Drabok 203

Dregeochloa Conert 29, 118, 217, 384
Dregeochloa calviniensis Conert 118
Dregeochloa pumila (Nees) Conert 118 (fig. 71), 363 (pi.
63)

Dronkgras 209, 246
Dropseed 306, 309
Duck grass 100
Duinekweek 320
Durban grass 100
Dwarf grass 229

Echinochloa Beauv. 7, 29, 119, 386, 400
Echinochloa colona (L.) Link 119, 120
Echinochloa crus-galli (L.) Beauv. 1 19 (fig. 72), 120, 364
(pi. 64)

426

Echinochloa crus-pavonis (Kunth) Schult. 120
Echinochloa haploclada (Stapf) Stapf 1 19, 120
Echinochloa holubii (Stapf) Stapf 120
Echinochloa jubata Stapf 120

Echinochloa pyramidalis (Lam.) Hitchc. & Chase 120
Echinochloa stagnina (Retz.) Beauv. 120, 121
Echinochloa subverticillata Pilg. 120
Echinochloa ugandensis Snowden & C.E. Hubb. 121
Eenjarige assegaaigras 179
Eenjarige borseltjiegras 43

Ehrharta Thunb. iii, 4, 5, 10, 11, 29, 121, 220, 383
Ehrharta aphylla Schrad. 126

Ehrharta barbinodis Nees ex Trin. 123
Ehrharta brevifolia Schrad.
var. brevifolia 123
var. cuspidata Nees 123
Ehrharta bulbosa J.E. Sm. 124

Ehrharta calycina J.E. Sm. 4, 12, 124 (fig. 74), 364 (pi.

65)

var. angustifolia 1 24
var. versicolor 124

Ehrharta capensis Thunb. 7, 121 (fig. 73), 124, 364 (pi.

66)

Ehrharta delicatula (Nees) Stapf 125
Ehrharta dodii Stapf 1 27
Ehrharta dura Nees ex Trin. 7, 125
Ehrharta eburnea Gibbs Russell 125
Ehrharta erecta Thunb.
var. abyssinica (Hochst.) Pilg. 125
var. erecta 1 1, 122, 125
var. natalensis Stapf 125
Ehrharta gigantea Thunb. 1 28
Ehrharta longiflora J.E. Sm. 125
Ehrharta longifolia Schrad. 125, 364 (pi. 67)

Ehrharta longigluma C.E. Hubb. 126
Ehrharta melicoides Thunb. 126
Ehrharta microlaena Nees ex Trin. 7, 126
Ehrharta ottonis Kunth ex Nees 126
Ehrharta pusilla Nees ex Trin. 126
Ehrharta ramosa (Thunb.) Thunb.,
subsp. aphylla (Schrad.) Gibbs Russell 126
subsp. ramosa 7, 126
Ehrharta rehmannii Stapf
subsp. filiformis (Stapf) Gibbs Russell 127
subsp. rehmannii 126, 127
subsp. subspicata (Stapf) Gibbs Russell 127
var. filiformis Stapf 127
Ehrharta rupestris Nees ex Trin.
subsp. dodii (Stapf) Gibbs Russell 127
subsp. rupestris 7, 127
subsp. tricostata (Stapf) Gibbs Russell 127
Ehrharta setacea Nees
subsp. disticha (Stapf) Gibbs Russell 127
subsp. scabra (Stapf) Gibbs Russell 128
subsp. setacea 128

subsp. uniflora (Burch, ex Stapf) Gibbs Russell 128
var. scabra Stapf 128
Ehrharta subspicata Stapf 127
Ehrharta thunbergii Gibbs Russell 126, 128
Ehrharta triandra Nees ex Trin. 125, 128
Ehrharta tricostata Stapf 127
Ehrharta uniflora Burch, ex Stapf 128
Ehrharta villosa Schult. f.
var. maxima Stapf 128
var. villosa 1, 122, 126, 128
Ehrharta virgata Launert 128
Elbow buffalo grass 242
Elephant grass 184, 250
Eleusine Gaertn. 1 1, 29, 129, 385, 419
Eleusine africana K. -O’Byrne 129
Eleusine coracana (L.) Gaertn.
subsp. africana (K. -O’Byrne) Hilu & De Wet 129 (fig.

75), 364 (pi. 68)

Eleusine indica (L.) Gaertn.
subsp. africana (K. -O’Byrne) S.M. Phillips 129
subsp. indica 130

Eleusine multiflora Rich. 130
Eleusine tristachva (Lam.) Lam. 130
Elionurus Willd. 30, 130, 388
Elionurus argenteus Nees 131
Elionurus glaber Phill. 131
var. villosus Phill. 131

Elionurus muticus (Spreng.) Kunth 43, 111, 130 (fig. 76),
131, 289, 364 (pi. 69)

Elionurus pretoriensis Phill. 131
Elionurus tripsacoides Willd. 131
Elsgras 219

Elymandra Stapf 1 1, 30, 131, 387

Elvmandra grallata (Stapf) Clayton vii, 1 3 1 (fig. 77), 132,
‘364 (pi. 70)

Elvtrigia Desv. 29, 132. 181, 292, 336. 382

Ely trigia repens (L.) Nevski 132 (fig. 78), 364 (pi. 71)

Elytrophorus Beauv. 29. 133, 384

Elytrophorus globularis Hack. 133 (fig. 79), 364 (pi. 72)

Elytrophorus spicatus (Willd.) A. Camus 133

Engler’s bristle grass 50

English bluegrass 170

Enneapogon Beauv. 10. 29, 134. 385, 399

Enneapogon brachystachyum (Jaub. & Spach) Stapf 135

Enneapogon cenchroides (Roem. & Schult.) C.E. Hubb.

80. 134 (fig. 80). 135, 136, 365 (pi. 73)

Enneapogon desvauxii Beauv. 5, 11, 135 (fig. 81)
Enneapogon filifolius (Pilg.) Stapf ex Garabedian 136
Enneapogon pretoriensis Stent 135
Enneapogon scaber Lehm.
var. scaber 135

var. (=De Winter & Hardy 8051) 135
Enneapogon scoparius Stapf 136
Enneapogon spathaceus Goossens 136
Enneapogon sp. (=E1 1 is 3208) 136
Enteropogon Nees 29, 136. 385

Enteropogon macrostachvus (A. Rich.) Benth. 136 (fig.
82), 137. 365 (pi. 74)

Enteropogon monostachyus (Vahl) K. Schum.

subsp. africanus Clayton 137
Enteropogon prieurii (Kunth) Clayton 137
Enteropogon rupestris (J.A. Schmidt) A. Chev. 137
Enteropogon simplex (Schumach. & Thonn.) A. Chev. 137
Entolasia Stapf 29, 137, 386

Entolasia imbricata Stapf vii, 138 (fig. 83), 365 (pi. 75)
Entolasia olivacea Stapf 138
Entoplocamia Stapf 10, 11, 29, 138, 385
Entoplocamia aristulata (Hack. & Rendle) Stapf 138 (fig.
84), 139, 365 (pi. 76)

Eragrostis Wolf 4, 5, 10, 11, 29, 104, 139, 315, 385, 386
Eragrostis abyssinica (Jacq.) Link 162
Eragrostis acraea De Winter 144
Eragrostis aethiopica Chiov. 144, 156, 159
Eragrostis annulata Rendle ex Scott Elliot 144, 146, 148
Eragrostis arenicola C.E. Hubb. 145, 148, 162, 163
Eragrostis aristata De Winter 145
Eragrostis aspera (Jacq.) Nees 145, 152
Eragrostis atherstonei Stapf 162
Eragrostis atrovirens auctt.. non Trin. ex Steud. 151
Eragrostis barbinodis Hack. 145. 149, 159
Eragrostis barrelieri Dav. 145, 146, 148, 154
Eragrostis bergiana (Kunth) Trin. 145, 163
Eragrostis bicolor Nees 146

Eragrostis biflora Hack, ex Schinz 146 (fig. 86), 150
Eragrostis brizantha Nees 146. 150
Eragrostis caesia Stapf 147, 149

Eragrostis capensis (Thunb.) Trin. 147 (fig. 87), 148, 365
(pi. 77)

Eragrostis chalcantha Trin. 158
Eragrostis chapelieri (Kunth) Nees 147, 150, 151
Eragrostis chloromelas Steud. 148, 149
Eragrostis cilianensis (All.) F.T. Hubb. 145, 148, 154
Eragrostis ciliaris (L.) R. Br. 145, 148, 162, 163
Eragrostis cimicina Launert 147, 148
Eragrostis congesta Oliv. 148
Eragrostis crassinervis Hack. 149
Eragrostis curvula (Schrad.) Nees 2, 4, 9, 139 (fig. 85),
148, 149, 152, 157, 365 (pi. 78)

427

Eragrostis cylindriflora Hochst. 149, 150, 155, 157, 162
Eragrostis cyperoides (Thunb.) Beauv. 86
Eragrostis denudata Hack, ex Schinz 154
Eragrostis desolata Launert 161
Eragrostis dinteri Stapf 149, 160
Eragrostis echinochloidea Stapf 146, 150, 155, 158
Eragrostis elatior Stapf 148, 150
Eragrostis gangetica (Roxb.) Steud. 150, 154
Eragrostis glandulosipedata De Winter 149, 150, 155
Eragrostis gummiflua Nees 150
Eragrostis habrantha Rendle 146, 150, 161
Eragrostis henrardii Jansen 162
Eragrostis heteromera Stapf 151, 160
Eragrostis hierniana Rendle 151, 152
Eragrostis homomalla Nees 151
Eragrostis horizontalis Peter 149
Eragrostis hygrophila C.E. Hubb. & Schweick. 151
Eragrostis inamoena K. Schum. 148, 151, 152
Eragrostis jeffreysii Hack. 151
Eragrostis kingesii De Winter 152, 157, 158
Eragrostis laevissima Hack. 152
Eragrostis lamprospicula De Winter 152, 154
Eragrostis lappula Nees 151, 152
Eragrostis leersiiformis Launert 145, 152
Eragrostis lehmanniana Nees
var. chaunantha (Pilg.) De Winter 152, 153
var. lehmanniana 6, 149, 153 (fig. 88), 159, 162
Eragrostis maerochlamys Pilg.
var. maerochlamys 153, 157

var. wilmaniae (C.E. Hubb. & Schweick.) De Winter 153
Eragrostis margaritacea Stapf 160
Eragrostis membranacea Hack, ex Schinz 150, 152, 154
Eragrostis micrantha Hack. 150, 152, 154, 161
Eragrostis minor Host 145, 146, 148, 154
Eragrostis moggii De Winter
var. moggii 154

Eragrostis namaquensis Nees ex Schrad. 104
Eragrostis nebulosa Stapf 157

Eragrostis nindensis Fical. & Hiern 154, 155 (fig. 89), 158
Eragrostis obtusa Munro ex Fical. & Hiern 6, 155, 158
Eragrostis omahekensis De Winter 149, 150, 155
Eragrostis pallens Hack. 155, 156 (fig. 90)

Eragrostis patens Oliv. 148. 155
Eragrostis patentissima Hack. 156
Eragrostis pilgeriana Dinter ex Pilg. 156, 162
Eragrostis pilosa (L.) Beauv. 144, 156, 159
Eragrostis plana Nees 157, 162, 31 1
Eragrostis planiculmis Nees 149, 157
Eragrostis poaeoides Beauv. ex Roem. & Schult. 154
Eragrostis porosa Nees 149, 157
Eragrostis procumbens Nees 152, 153, 154, 157
Eragrostis x pseud-obtusa De Winter 155, 157, 158 (fig.
91)

Eragrostis pseudosclerantha Chiov. 152, 158
Eragrostis pusilla Hack. 104
Eragrostis pygmaea De Winter 152, 158
Eragrostis racemosa (Thunb.) Steud. 154, 158, 161, 365
(pi. 79)

Eragrostis remotiflora De Winter 144, 156, 159
Eragrostis rigidior Pilg. 145, 149, 159 (fig. 92)
Eragrostis robusta Stent 149
Eragrostis rogersii C.E. Hubb. 149, 159
Eragrostis rotifer Rendle 151, 160
Eragrostis sabinae Launert 152, 160
Eragrostis sabulosa (Steud.) Schweick. 160
Eragrostis sarmentosa (Thunb.) Trin. 160
Eragrostis sclerantha Nees
subsp. sclerantha 158, 160, 161
subsp. villosipes (Jedw.) Launert 161
var. villosipes (Jedw.) De Winter 161
Eragrostis scopelophila Pilg. 161
Eragrostis spinosa (L. f.) Trin. 86
Eragrostis stapfii De Winter 140, 161
Eragrostis stenothyrsa Pilg. 161
Eragrostis superba Peyr. 9, 147, 156, 161, 365 (pi. 80)
Eragrostis tef (Zucc.) Trotter 162

Eragrostis tenella (L.) Roem. & Schult. 145, 148, 162, 163
Eragrostis tenuifolia (A. Rich.) Steud. 157, 162
Eragrostis trichophora Coss. & Dur. 149, 162
Eragrostis truncata Hack. 146, 163
Eragrostis uniglumis Hack. 151
Eragrostis virescens Presl 163
Eragrostis viscosa (Retz.) Trin. 145, 148, 163
Eragrostis volkensii Pilg. 163, 344
Eragrostis walteri Pilg. 2, 140, 149, 163, 385, 386, 409
Eriochloa Kunth 29, 164, 386, 400
Eriochloa borumensis sensu Hack., non Stapf 165
Eriochloa borumensis sensu Stapf, non Hack. 165
Eriochloa fatmensis (Hochst. & Steud.) Clayton 164, 165
Eriochloa macclounii Stapf 164
Eriochloa meyeriana (Nees) Pilg.
subsp. grandiglumis (Stent & Rattray) Gibbs Russell 165
subsp. meyeriana 12, 164 (fig. 93), 165, 342, 365 (pi. 81)
Eriochloa nubica (Steud.) Hack. & Stapf ex Thell. 164
Eriochloa parvispiculata C.E. Hubb. 165
Eriochloa stapfiana Clayton 10, 164, 165
Eriochrysis Beauv. 30, 165, 387
Eriochrysis brachypogon (Stapf) Stapf 166
subsp. australis J.G. Anders. 166
Eriochrysis pallida Munro 165 (fig. 94), 166, 366 (pi. 82)
Eulalia Kunth 30, 166, 387
Eulalia aurea (Bory) Kunth 167
Eulalia geniculata Stapf 167

Eulalia villosa (Thunb.) Nees 166 (fig. 95), 167, 191, 366
(pi. 83)

Eustachys Desv. 10, 29, 167, 385
Eustachys mutica auett. 168

Eustachys paspaloides (Vahl) Lanza & Mattei 7, 167 (fig.
96), 168, 366 (pi. 84), 415

False barley 182
False couch finger grass 1 1 1
False signal grass 65
Famine grass 310
Fan-leaved bristle grass 295
Feathered chloris 84
Feathertop 25 1
Fern grass 79

Festuca L. 5, 10, 1 1, 29, 73, 168, 202, 203, 354, 382

Festuca africana (Hack.) Clayton 169, 318

Festuca arundinacea Schreb. 170

Festuca caprina Nees 169, 170

Festuca costata Nees 7, 169 (fig. 97), 170, 366 (pi. 85)

Festuca dracomontana Linder 170

Festuca elatior L. 170

Festuca killickii K. -O’Byrne 170

Festuca longipes Stapf 170

Festuca ovina L. 7, 409

Festuca scabra Vahl 10, 169, 170

Festuca vulpioides Steud. 170

Fibrous dropseed 312

Fine-bristled burgrass 80

Fine-leaved finger grass 1 13

Fine thatching grass 185

Finger grass 109

Fingerhuthia Nees 29, 171, 385

Fingerhuthia africana Lehm. 2, 7, 9, 171 (fig. 98), 366 (pi.
86)

Fingerhuthia sesleriiformis Nees 171
Flaccid finger grass 1 10
Fringed dropseed 31 1
Fog grass 35

Folded leaf tussock grass 297
Footpath love grass 158
Forest hood grass 285
Fountain grass 250
Foxtail chess 76
Foxtail millet 296
Fynblousaadgras 312
Fyntamboekiegras 185
Fyn vleigras 146

428

Garden bristle grass 297
Garssteekgras 50
Gastridium Beauv. 29, 172, 382

Gastridium phleoides (Nees & Meyen) C.E. Hubb. 172
(fig. 99), 366 (pi. 87)

Gause grass 78

Gazochloa scopulorum J.B. Phipps 102

Geelaartamboekiegras 187

Geelhoutpluimgras 151

Geeltamboekiegras 189

Gesteektegras 42

Gewone paspalum 246

Gaint crowfoot 101

Giant reed 58

Giant spear grass 337

Giant thatching grass 187

Giant trident grass 348

Giant turpentine grass 95

Golden beard grass 85

Gomgras 150

Gongoni-steekgras 5 1

Goose grass 1 30

Grootbewertjiegras 72

Grootpluimeragrostis 145

Grootwitbaardandropogon 41

Guineafowl grass 283

Guinea grass 239

Gulf barnyard grass 120

Gum grass 150

Haasgras 126, 194, 229, 287
Hackelochloa Kuntze 30, 172, 388, 400
Hackelochloa granularis (L.) Kuntze 173 (fig. 100), 366
(pi. 88)

Hainardia Greuter4, 29, 173, 381, 382
Hainardia cylindrica (Willd.) Greuter 173 (fig. 101), 174,
200, 203, 243, 366 (pi. 89)

Hairy bluegrass 40
Hairy blue thatching grass 184
Hairy chess 74
Harde steekgras 55
Harestail 194

Harige bloutamboekiegras 184
Harpochloa Kunth 29, 174, 385

Harpochloa falx (L. f.) Kuntze 6, 174 (fig. 102), 366 (pi.
90)

Hartjie-eragrostis 147

Helictotrichon Schult. 7, 10, 12, 29, 175, 382
Helictotrichon barbatum (Nees) Schweick. 175
Helictotrichon capense Schweick. 175, 176
Helictotrichon dodii (Stapf) Schweick. 175, 177
Helictotrichon galpinii Schweick. 176
Helictotrichon hirtulum (Steud.) Schweick. 175, 176 (fig.
103)

Helictotrichon leoninum (Steud.) Schweick. 176
Helictotrichon longifolium (Nees) Schweick. 176. 177
Helictotrichon longum (Stapf) Schweick. 177
Helictotrichon namaquense Schweick. 177
Helictotrichon natalense (Stapf) Schweick. 176, 177
Helictotrichon quinquesetum (Steud.) Schweick. 177
Helictotrichon turgidulum (Stapf) Schweick. 176, 177,
367 (pi. 91)

Helictotrichon sp. (=Ellis 4663) 177
Hemarthria R. Br. 10, 30, 178, 200, 388, 391, 400
Hemarthria altissima (Poir.) Stapf & C.E. Hubb. 12, 178
(fig. 104), 367 (pi. 92)

Herringbone grass 273
Heterocarpha schiemanniana Schweick. 70
Heteropogon Pers. 10, 12, 30, 178, 387
Heteropogon contortus (L.) Roem. & Schult. 2, 4, 7, 9,
12, 179 (fig. 105), 290, 335, 337, 349, 367 (pi. 93)
Heteropogon melanocarpus (Ell.) Benth. 2, 179
Hippo grass 353
Holcus L. vii, 10, 29, 180, 382
Holcus lanatus L. 180 (fig. 106), 367 (pi. 94)

Holcus mollis L. 180

Holcus setiger Nees 180
Hordeum L. 9, 29, 132, 180. 382
Hordeum capense Thunb. 181
Hordeum compressum Griseb. 182
Hordeum marinum Huds.

subsp. gussoneanum (Pari.) Thell. 181, 182
Hordeum murinum L.
subsp. glaucum (Steud.) Tzvel 182
subsp. leporinum (Link) Archangeli 182
subsp. murinum 7, 181 (fig. 107), 182, 367 (pi. 95)
Hordeum nodosum auctt., non L. 181
Hordeum stenostachys Godr. 182
Hordeum vulgare L. 1

Hvparrhenia Fourn. iii, 9, 10, 30, 182, 223, 387
Hyparrhenia anamesa Clayton 184, 185
Hyparrhenia aucta (Stapf) Stapf ex Stent 184
Hyparrhenia buchanani (Stapf) Stapf ex Stent 186
Hyparrhenia collina (Pilg.) Stapf 184, 186, 187
Hyparrhenia cymbaria (L.) Stapf 184. 186, 188
Hyparrhenia dichroa (Steud.) Stapf 184. 185, 187
Hyparrhenia dissoluta (Nees ex Steud.) C.E. Hubb. 189
Hy parrhenia dregeana (Nees) Stapf 184, 185, 186, 187
Hyparrhenia familiaris (Steud.) Stapf 186
Hyparrhenia filipendula (Hochst.) Stapf
var. filipendula 184, 185, 186
var. pilosa (Hochst.) Stapf 185
Hyparrhenia finitima (Hochst.) Anderss. ex Stapf 184,
' 185

Hyparrhenia gazensis (Rendle) Stapf 184, 185, 186
Hyparrhenia glauca Stent 1 87

Hvparrhenia hirta (L.) Stapf 4, 7, 9, 183 (fig. 108), 184,
' 185. 186, 367 (pi. 96), 414

Hyparrhenia newtonii (Hack.) Stapf
var. macra Stapf 185
var. newtonii 185, 186

Hyparrhenia nyassae (Rendle) Stapf 185, 186
Hyparrhenia pilgeriana C.E. Hubb. 186
Hyparrhenia pilosissima (Hack.) J.G. Anders. 184
Hyparrhenia poecilotricha (Hack.) Stapf 186, 187
Hyparrhenia quarrei Robyns 185, 186
Hyparrhenia rudis Stapf 184, 186, 187, 188
Hyparrhenia rufa (Nees) Stapf
var. rufa 184, 186, 187

Hyparrhenia schimperi (A. Rich.) Stapf 186, 187, 188
Hyparrhenia tamba (Steud.) Stapf 5, 184, 186, 187 (fig.
* 109)

Hyparrhenia umbrosa (Hochst.) Anderss. ex Clayton 184,
188

Hyparrhenia variabilis Stapf 184, 187, 188
Hyperthelia Clayton 30, 188, 387

Hyperthelia dissoluta (Steud.) Clayton 188 (fig. 1 10), 189,
367 (pi. 97)

Hypogynium Nees 5

Hypogynium schlechteri (Hack.) Pilg. 41
Imperata L. 30, 189. 387

Imperata cylindrica (L.) Raeuschel 6, 9, 11, 12, 189 (fig.
Ill), 190, 367 (pi. 98)
var. africana (Anderss.) C.E. Hubb. 190
var. major (Nees) C.E. Hubb. 190
Ischaemum L. 30, 190, 387

Ischaemum afrum (J.F. Gmel.) Dandy 191, 367 (pi. 99)
Ischaemum arcuatum (Nees) Stapf 191
Ischaemum brachyatherum (Hochst.) Hack. 191
Ischaemum fasciculatum Brongn. 167, 190 (fig. 1 12), 191
Ischaemum franskae J.M. Wood 267
Ischaemum glaucostachyum Stapf 191
Italian rye grass 202

Jakkalsstert 50
Job’s tears 89
Johnson grass 302
Jungle rice 1 19

Kalahari dune bushman grass 320
Kalahari sandkweek 291

429

Kalahari water grass 120
Kalkgras 135, 306
Kalkkweek 145
Kammetjiesgras 31 1

Kaokochloa De Winter 29, 191, 385, 399
Kaokochloa nigrirostris De Winter 191 (fig. 113), 192,
368 (pi. 100)

Kapokgras 40
Kappiegras 285

Karroochloa Conert & Tuerpe 5, 29, 192, 384, 400
Karroochloa curva (Nees) Conert & Tuerpe 7, 192 (fig.
114), 193

Karroochloa purpurea (L. f.) Conert & Tuerpe 193, 368
(pi. 101)

Karroochloa schismoides (Stapf ex Conert) Conert &
Tuerpe 193

Karroochloa tenella (Nees) Conert & Tuerpe 193

Karroo quick grass 97

Katstertsteekgras 49

Kattestertgras 116

Kentucky bluegrass 272

Kikuyu grass 248

Kinagras 349

Kleinbewertjiegras 72

Kleinrolblaar 281

Klitsgras 338

Klitssetaria 300

Klosgras 63

Klossiegras 84

Knietjiesgras 153

Knotroot 295

Koeleria Pers. vii, 29, 193, 204, 382, 400
Koeleria capensis (Steud.) Nees 34, 194 (fig. 115), 204,
288, 368 (pi. 102)

Koeleria cristata (L.) Pers. 194
Koeleria cristata auctt., non (L.) Pers.
var .cristata 194

var. brevifolia (Nees) C.E. Hubb. 194
var. convoluta (Steud.) C.E. Hubb. 194
Koeleria phleoides (Vill.) Pers. 205
Kokoma grass 283
Koperdraad 1 3 1
Koperdraadgras 49
Kophaargras 338
Kortbeenboesmangras 326
Kortblaarboesmangras 321
Kousklits 338
Kropaargras 332
Kruipgras 241
Kruipsinjaalgras 66
Krulblaar 159
Krulgras 85
Kuilgras 1 17

Kuruman finger grass 1 13
Kweekgras 97

Lagurus L. 29, 194, 382

Lagurus ovatus L. 194, 195 (fig. 1 16), 368 (pi. 103)
Lamarckia Moench. 14, 29, 195, 382
Lamarckia aurea (L.) Moench. 195 (fig. 117), 196, 368
(pi. 104)

Land grass 241

Langbaardswenkgras 355

Langbeenboesmangras 321

Langbeen paspalum 247

Langbeensteekgras 51

Langbeentwagras 321

Langnaaldbromus 74

Langnaaldsteekgras 53

Large carrot-seed grass 338

Large silver andropogon 41

Large woolly three-awn 55

Lasiochloa Kunth 5, 339

Lasiochloa echinata (Thunb.) Adamson 340

Lasiochloa longifolia (Schrad.) Kunth 340

Lasiochloa ohtusifolia Nees 341

Lasiochloa utriculosa Nees 341

Leersia Swartz. 29, 196, 383
Leersia denudata Launert 197
Leersia friesii Meld. 197

Leersia hexandra Swartz 7, 1 1, 196 (fig. 118), 197, 368
(pi. 105)

Leersia tisserantii (A. Chev.) Launert 197

Leeugras 324
Lehmann’s love grass 153
Lemoengras 95

Leptocarydion Stapf 29, 197, 385

Leptocarydion vulpiastrum (De Not.) Stapf 197 (fig.

119), 198, 368 (pi. 106)

Leptochloa Beauv. 29, 1 16, 198, 385, 391
Leptochloa chinensis (L.) Nees 199
Leptochloa panicea (Retz.) Ohwi 198 (fig. 120), 199, 368
(pi. 107)

Leptochloa uniflora A. Rich. 199
Lepturus R. Br. 29, 199, 385

Lepturus repens (G. Forst.) R. Br. vii, 174, 199 (fig. 121),
200, 368 (pi. 108)

Leucophrys Rendle 5, 29, 200, 386
Leucophrys mesocoma (Nees) Rendle 200 (fig. 122), 369
(pi. 109)

Limestone dropseed 306

Limpopo grass 120

Lintonia Stapf 29, 201, 385

Lintonia nutans Stapf 201 (fig. 123), 369 (pi. 1 10)

Litjiesinjaalgras 66

Little love grass 1 54

Little quaking grass 72

Lizardtail grass 173

Lolium L. 29, 168, 201, 381, 382

Lolium loliaceum (Bory & Chaup.) Hand.-Mazz 203

Lolium multiflorum Lam. 10, 202, 369 (pi. Ill)

Lolium multiflorum x L. perenne 202

Lolium perenne L. 202, 203

Lolium rigidum Gaudin 174, 203

Lolium temulentum L. 1,9, 202 (fig. 124), 203

Long-awned water grass 121

Long-plumed finger grass 1 14

Lophachme Stapf 29, 203, 385

Lophachme digitata Stapf 203 (fig. 125), 204, 369 (pi.
112)

Lophochloa Reichenb. vii, 29, 204, 382
Lophoehloa cristata (L.) Hyl. 205
Lophochloa pumila (Desf.) Bor 204 (fig. 126), 205, 369
(pi. 113)

Loudetia Steud. 10, 29, 205, 387

Loudetia anomala C.E. Hubb. & Schweick. 102

Loudetia densispica (Rendle) C.E. Hubb. 205

Loudetia filifolia Schweick. 206

Loudetia flavida (Stapf) C.E. Hubb. 206 (fig. 127)

Loudetia lanata (Stent & Rattray) C.E. Hubb. 206

Loudetia pedicellata (Stent) Chippind. 206

Loudetia ramosa (Stapf) C.E. Hubb. 102

Loudetia simplex (Nees) C.E. Hubb. 207, 369 (pl.l 14)

Loudetia superba De Not. 348

Mahem’s crest 281
Maize 1

Marram grass 38
Meadow fescue 170
Meadow grass 272
Mediterranean barley 181
Megaloprotachne C.E. Hubb. 29, 207, 386
Megaloprotachne albescens C.E. Hubb. 207 (fig. 128),
208,369 (pi. 115)

Megaloprotachne glabrescens Roiv. 208
Megastachya Beauv. 10, 29, 208, 381, 383
Megastachya mucronata (Poir.) Beauv. 7,"208 (fig. 1 —9),
369 (pi. 116)

Melica L. 7, 10, 1 1, 29, 209, 330, 381, 382, 400
Melica bolusii Stapf 209
Melica brevifolia Stapf 209

Melica decumbens sensu Gordon-Gray, non Thunb. 209

Melica decumbens Thunb. 209
Melica neesii Stapf 209

430

Melica ovalis Nees 209
Melica pumila Stapf 209

Melica racemosa Thunb. 10, 209 (fig. 130), 369 (pi. 1 17)
Melinis Beauv. 5, 29, 1 10, 201, 210, 386, 418
Melinis ambigua Hack. 213

Melinis drakensbergensis (C.E. Hubb. & Schweick.)
Clayton 211

Melinis kallimorpha (Clayton) Zizka 211
Melinis longiseta (A. Rich.) Zizka
subsp. bellespicata (Rendle) Zizka 211
subsp. longiseta 211

Melinis macrochaeta Stapf & C.E. Hubb. 211, 212 (fig.
132)

Melinis minutiflora Beauv. 210, 212, 370 (pi. 1 18)
Melinis nerviglumis (Franch.) Zizka 212
Melinis repens (Willd.) C.E. Hubb
subsp. grandiflora (Hochst.) Zizka 212, 213
subsp. repens 210 (fig. 131), 212, 370 (pi. 1 19)

Melinis scabrida (K. Schum.) Hackel 211, 213
Melinis subglabra Mez 213
Melinis tenuinervis (Stapf) Stapf 212
Melinis tenuissima Stapf 213
Merxmuellera Conert 2, 4, 5, 8, 29, 213, 384, 400
Merxmuellera arundinacea (Berg.) Conert 215, 370 (pi.
120)

Merxmuellera aureocephala (J.G. Anders.) Conert 215.

217

Merxmuellera cincta (Nees) Conert 215
Merxmuellera davyi (C.E. Hubb.) Conert 215, 217
Merxmuellera decora (Nees) Conert 215, 217, 218
var. tricostachya Stapf 215
Merxmuellera disticha (Nees) Conert 10, 213, 216
Merxmuellera drakensbergensis (Schweick.) Conert 213
(fig. 133), 216, 218

Merxmuellera dura (Stapf) Conert 216 (fig. 134)
Merxmuellera guillarmodiae Conert 216
Merxmuellera lupulina (Thunb.) Conert 217, 218
Merxmuellera macowanii (Stapf) Conert 215, 217
Merxmuellera papposa (Nees) Conert 217
Merxmuellera rangei (Pilg.) Conert 217
Merxmuellera rufa (Nees) Conert 217
Merxmuellera stereophylla (J.G. Anders.) Conert 216,

218

Merxmuellera stricta (Schrad.) Conert 216, 218, 370 (pi.
121)

Merxmuellera sp. (=Ellis 2500) 218
Mfufu 250

Microchloa R. Br. 29, 218, 281, 385
Microchloa caffra Nees 4, 7, 69, 218 (fig. 135), 219, 370
(pi. 122)

Microchloa indica (L. f.) Beauv. 219
Microchloa kunthii Desv. 219
Microchloa setacea R. Br. 219
Microlaena R. Br. 5, 14, 29, 219, 383
Microlaena stipoides R. Br. 11, 220, 370 (pis. 123 & 124)
Microstegium Nees 30, 220, 387
Microstegium capense (Hochst.) A. Camus 220
Microstegium nudum (Trin.) A. Camus 220 (fig. 136), 370
(pi. 125)

Milanje finger grass 1 1 1
Millet 1, 129, 249
Miscanthidium Stapf 221
Miscanthidium capense (Nees) Stapf
var. capense 221
var. villosa Stapf 221

Miscanthidium erectum Stent & C.E. Hubb. 221
Miscanthidium junceum Stapf 221
Miscanthidium sorghum (Nees) Stapf 221
Miscanthidium teretifolium (Stapf) Stapf 221
Miscanthus Anderss. 30, 221, 387

Miscanthus capensis (Nees) Anderss. 6, 221 (fig. 137),
370 (pi. 126)

Miscanthus junceus (Stapf) Pilg. 7, 221
Misty dropseed 3 1 2
Mnesithea Kunth 88, 172
Molasses grass 212

Molineriella Rouy 265
Monelytrum Hack. 29, 222, 385
Monelytrum annuum Goossens 222

Monelytrum luederitzianum Hack. 6, 222 (fig. 138). 371
(pk 127)

Monerma P. Beauv. 199

Monerma cylindrica auct., non (Willd.) Coss & Dur. 174
Monocymbium Stapf 30, 222, 387

Monocvmbium ceresiiforme (Nees) Stapf vii, 8, 41, 223
(fig. 139), 371 (pi. 128)

Mosdenia Stent 29, 223, 385

Mosdenia leptostachvs (Fical. & Hiem) Clayton 223 (fig.

140), 224, 371 (pi. 129)

Mosdenia phleoides (Hack.) Stent 224
Motwortelterpentyngras 95
Mountain andropogon 40
Mountain bristle grass 296
Mountain brome grass 75
Muiswildegars 182
Munnik fescue 170
Munnik-swenkgras 170
Muskusgras 95

Namib dune bushman grass 326
Napier fodder 250
Narrow heart love grass 158
Nassella Desv. 29, 224, 384, 400, 415
Nassella trichotoma (Nees) Hack, ex Arech. 224 (fig. 141),
318. 371 (pi. 130)

Nassella tussock 224
Natal buffalo grass 240
Natalbuffelsgras 240
Natal crowfoot 100
Natal grass 250
Natal red top 212
Nile grass 32

Oats 1, 59

Odontelytrum Hack. 8, 29, 225. 386, 400
Odontelytrum abvssinicum Hack, vii, 225 (fig. 142), 371
(pi. 131)

Odvssea Stapf 29, 225, 385

Odyssea paucinervis (Nees) Stapf 226 (fig. 143), 371 (pi.
132)

Old man's beard 40

Olifantsgras 184

Olive dropseed 307

Olyra L. 11,29, 226, 382,383

Olyra latifolia L. 6, 7, 10, 226, 227 (fig. 144), 371 (pi. 133)
One finger grass 1 1 1
Oortjiesgras 308

Oplismenus Beauv. 2, 29, 227, 278, 386
Oplismenus burmannii (Retz.) Beauv. 227
Oplismenus hirtellus (L.) Beauv. 2, 227, 228 (fig. 145),
371 (pi. 134)

Oplismenus undulatifolius (Ard.) Roem. & Schult. 228
Orchard grass 99
Oropetium Trin. 29, 228, 385

Oropetium capense Stapf 9, 229 (fig. 146), 371 (pi. 135)

Oryza L. 29, 229, 383

Oryza barthii auctt., non A. Chev. 229

Oryza barthii A. Chev. 229

Oryza longistaminata A. Chev. & Roehr. 10, 11, 229, 230
(fig. 147), 372 (pi. 136)

Oryza punctata Steud. 230
Oryza sativa L. 1, 229
Oryzidium C.E. Hubb. 29, 230, 386
Oryzidium barnardii C.E. Hubb. & Schweick. 230, 231
(fig. 148), 372 (pi. 137)

Osgras 129, 130
Oulandsgras 149

Oxyrhachis Pilg. 8, 30, 231, 386, 387, 388
Oxyrhachis gracillima (Bak.) C.E. Hubb. 231 (fig. 149),
232, 372 (pi. 138)

Oxytenanthera Munro 29, 232, 383
Oxytenanthera abyssinica (A. Rich.) Munro vii, 232 (fig.
150)

431

Panicum L. 2, 3, 5, 7, 10, 11, 29, 233, 278, 342, 386, 409

Panicum aequinerve Nees 235
Panicum aphanoneurum Steud. 237
Panicum arbusculum Mez 235
Panicum arcurameum Stapf 235, 236, 240
Panicum atrosanguineum A. Rich. 235, 236, 240
Panicum bechuanense Brem. & Oberm. 236, 242
Panicum brevifolium L. 238
Panicum chusqueoides Hack. 65
Panicum coloratum L.
var. coloratum 236, 239, 242
var. makarikariense Goossens 236
Panicum comorense Mez 236
Panicum deustum Thunb. 10, 236
Panicum dewinteri J.G. Anders. 238
Panicum dregeanum Nees 237
Panicum ecklonii Nees 2, 237
Panicum filiculme Schinz 238
Panicum fluviicola Steud. 237
Panicum ful gens auctt., non Stapf 240
Panicum genuflexum Stapf 237
Panicum gilvum Launert 237, 238, 242
Panicum glabrescens Steud. 242
Panicum glandulopaniculatum Renvoize 238
Panicum heterostachyum Hack. 238
Panicum hians Ell. 238
Panicum hymeniochilum Nees 238
var. glandulosum Nees 238
var. hymeniochilum 238
Panicum impeditum Launert 238, 242
Panicum inaequilatum Stapf & C.E. Hubb. 235
Panicum infestum Peters 238, 239
Panicum kalaharense Mez 239
Panicum laevifolium Hack,
var. con trac turn Pilg. 237
var. laevifolium 241
Panicum lanipes Mez 239
Panicum laticomum Nees 2, 239

Panicum maximum Jacq. 9, 233 (fig. 151), 239, 372 (pi.

139), 418

Panicum merkeri Mez 236, 239
Panicum meyerianum Nees
var. grandeglume Stent & Rattray 165
var. meyerianum 1 65
Panicum miliaceum L. 235
Panicum monticola Hook. f. 236, 239
Panicum natalense Hochst. 9, 240 (fig. 152), 372 (pi. 140)
Panicum novemnerve Stapf 235, 236, 240, 243
Panicum obumbratum Stapf 240
Panicum parvifolium Lam. 241, 242
Panicum pearsonii Bol. f. 239
Panicum phragmitoides Stapf 239
Panicum pilgerianum (Schweick.) Clayton 241
Panicum radula Mez 239
Panicum repens L. 241
Panicum repentellum Napper 241
Panicum schinzii Hack. 241, 243
Panicum stapfianum Fourc. 236, 239, 241, 242
Panicum subalbidum Kunth 238, 241, 242
Panicum subflabellatum Stapf 241, 242
Panicum trichonode Launert & Renvoize 242
Panicum volutans J.G. Anders. 242
Panicum sp. aff. P. hippothrix K. Schum. 242
Panicum sp. 1 (=Smook 3463) 238, 242
Panicum sp. 2 (=Giess 8605) 241, 242
Pan dropseed 309
Pankweek 313

Parapholis C.E. Hubb. 29, 243, 382

Parapholis incurva (L.) C.E. Hubb. vii, 174, 243 (fig.

153) , 372 (pi. 141)

Paratheria Griseb. 14, 29, 243, 386
Paratheria prostrata Griseb. 244, 372 (pi. 142)
Paspalidium Stapf 29, 244, 386, 400
Paspalidium geminatum (Forssk.) Stapf 244, 245
Paspalidium obtusifolium (Del.) Simpson 7, 244 (fig.

154) , 245, 372 (pi. 143)

Paspalidium platyrrhachis C.E. Hubb. 245

Paspalum L. 12, 29, 245, 386, 400
Paspalum commersonii Lam. 246

Paspalum dilatatum Poir. 245 (fig. 155), 246, 247, 372 (pi
144)

Paspalum distichum L. 245, 246, 247
Paspalum notatum Fluegge 245, 246, 247
Paspalum orbiculare Forst.

Paspalum paspalodes (Michx.) Scribn. 246
Paspalum polystachyum R. Br. 246
Paspalum scrobiculatum L. 245, 246
Paspalum urvillei Steud. 246, 247
Paspalum vaginatum Swartz 245, 246, 247
Pearl millet 249

Pennisetum Rich. 5, 10, 29, 247, 386, 415, 416
Pennisetum albicauda Stapf & C.E. Hubb. 249
Pennisetum americanum (L.) Leeke 1
subsp. americanum 249

Pennisetum clandestinum Chiov. 1,5, 11, 247, 248, 249
(fig. 157)

Pennisetum echinurus (K. Schum.) Stapf & C.E. Hubb. 249

Pennisetum foermerianum Leeke 80, 248
Pennisetum glaucodadum Stapf & C.E. Hubb. 248, 249
Pennisetum glaucum (L.) R. Br. I, 249, 250
Pennisetum macrourum Trin. 248, 249, 250
Pennisetum mezianum Leeke 249
Pennisetum natalense Stapf 248, 249
Pennisetum nigritarum (Schlecht.) Dur. & Schinz 249
Pennisetum purpureum Schumach. 250
Pennisetum setaceum (Forssk.) Chiov. 250 (fig. 158), 373
(pi. 145)

Pennisetum sphacelatum (Nees) Dur. & Schinz 9, 248
(fig. 1 56), 250
var. sphacelatum 250
var. tenuifolium (Hack.) Stapf 250
Pennisetum stapfianum F. Bol. 249
Pennisetum thunbergii Kunth 248, 250
Pennisetum typhoides (Burm. f.) Stapf & C.E. Hubb. 249
Pennisetum unisetum (Nees) Benth. 14, 250
Pennisetum villosum R. Br. ex Fresen. 1, 251
Pentameris Beauv. 4, 8, 11, 29, 251, 278, 384, 413
Pentameris dregeana Stapf 252
Pentameris longiglumis (Nees) Stapf 252
Pentameris macrocalycina (Steud.) Schweick. 251 (fig.
159), 252, 253

Pentameris obtusifolia (Hochst.) Schweick. 252, 253
Pentameris speciosa Nees 252
Pentameris squarrosa Stapf 252
Pentameris thuarii Beauv. 7, 253, 373 (pi. 146)
Pentameris sp. 1 (=Esterhuysen 11115) 253
Pentameris sp. 2 (=Ellis 2546) 253
Pentaschistis Stapf iii, vii, 2, 4, 5, 12, 14, 15,253,384.413
Pentaschistis acinosa Stapf 256
Pentaschistis airoides (Nees) Stapf
subsp. airoides 253 (fig. 160), 256
subsp. jugorum (Stapf) Linder 256
Pentaschistis alticola Linder 256
Pentaschistis ampla (Nees) McClean 257, 263
Pentaschistis angulata sensu Adamson, non Nees 257
Pentaschistis angustifolia (Nees) Stapf 262
var. albescens Stapf 262
var. cirrhulosa (Nees) Stapf 258
var. micrathera (Nees) Stapf 260
Pentaschistis argentea Stapf 217, 257
Pentaschistis aristidoides (Thunb.) Stapf 257
Pentaschistis aristifolia Schweick. 257
Pentaschistis aspera (Thunb.) Stapf 257
Pentaschistis aurea (Steud.) McClean
subsp. aurea 257

subsp. pilosogluma (McClean) Linder 257
Pentaschistis bachmannii McClean 259
Pentaschistis barbata (Nees) Linder
subsp. barbata 257
subsp. orientalis Linder 258
Pentaschistis basutorum Stapf 216, 258
Pentaschistis brachyanthera Stapf 264
Pentaschistis burchellii Stapf 258

432

Pentaschistis calcicola Linder
var. calcicola 258
var. hirsuta Linder 258
Pentaschistis capensis (Nees) Stapf 258
Pentaschistis capillaris (Thunb.) McClean 258
Pentaschistis caulescens Linder 258
Pentaschistis chippindalliae Linder 258
Pentaschistis cirrhulosa (Nees) Linder 258
Pentaschistis colorata (Steud.) Stapf 259
var. polytricha Stapf 259

Pentaschistis curvifolia (Schrad.) Stapf 259 (fig. 161), 373
(pi. 147)

Pentaschistis densifolia (Nees) Stapf 259
Pentaschistis densifolia (Nees) Stapf
var. intricata Stapf 259
Pentaschistis ecklonii (Nees) McClean 259
Pentaschistis elegans (Nees) Stapf 260
Pentaschistis eriostoma (Nees) Stapf 260
Pentaschistis euadenia Stapf 262
Pentaschistis exserta Linder 260
Pentaschistis fibrosa Stapf 264
Pentaschistis filiformis (Nees) Stapf 262
Pentaschistis galpinii (Stapf) McClean 260, 373 (pi. 148)
Pentaschistis glandulosa (Schrad.) Linder 260, 263
Pentaschistis heptamera (Nees) Stapf 260
Pentaschistis heterochaeta Stapf 262
Pentaschistis holciformis (Nees) Linder 260
Pentaschistis imperfecta Stapf 262
Pentaschistis involuta sensu Adamson & Chippindall 257
Pentaschistis jugorum Stapf 256
Pentaschistis juncifolia Stapf 260
Pentaschistis leucopogon Stapf 257
Pentaschistis lima (Nees) Stapf 260
Pentaschistis longipes Stapf 261

Pentaschistis malouinensis (Steud.) Clayton 261 (fig. 162)
Pentaschistis microphylla (Nees) McClean 261
Pentaschistis montana Linder 261
Pentaschistis natalensis Stapf 261
Pentaschistis nutans (Nees) Stapf 264
Pentaschistis oreodoxa Schweick. 261, 263
Pentaschistis pallescens (Schrad.) Stapf 262
Pentaschistis pallida (Thunb.) Linder 4, 262
Pentaschistis papillosa (Steud.) Linder 262
Pentaschistis patula (Nees) Stapf 262
Pentaschistis patula (Nees) Stapf
var. acuta Stapf 262
var. glabrata Stapf 256
Pentaschistis patuliflora Rendle 258
Pentaschistis pilosogluma McClean 257
Pentaschistis praecox Linder 262
Pentaschistis pseudopallescens Linder 262
Pentaschistis pungens Linder 262
Pentaschistis pusilla (Nees) Linder 14, 262, 373 (pi. 149)
Pentaschistis pyrophila Linder 263
Pentaschistis reflexa Linder 263
Pentaschistis rigidissima Pilg. ex Linder 263
Pentaschistis rosea Linder
subsp. purpurascens Linder 263
subsp. rosea 263

Pentaschistis rupestris (Nees) Stapf 263, 265
Pentaschistis scandens Linder 263
Pentaschistis setifolia (Thunb.) McClean 263
Pentaschistis steudelii McClean 261
Pentaschistis subulifolia Stapf 262
Pentaschistis sylvatica Adamson 262
Pentaschistis thunbergii sensu Stapf 262
Pentaschistis tomentella Stapf 264
Pentaschistis tortuosa (Trin.) Stapf 264
Pentaschistis triseta (Thunb.) Stapf 264
Pentaschistis tysonii Stapf 260, 264 (fig. 163)
Pentaschistis velutina Linder 265
Pentaschistis veneta Linder 263, 265
Pentaschistis viscidula (Nees) Stapf 217, 265
Pentaschistis zeyheri Stapf 262
Perdegras 178
Perennial rye grass 203

Periballia Trin. 14, 29, 265, 382
Periballia minuta (L.) Asch. & Graebn. 36, 265
Perotis Aiton 29, 265, 385
Perotis leptopus Pilg. 266

Perotis patens Gand. 7, 266 (fig. 164), 373 (pi. 150)

Perotis vaginata Hack. 266

Pers steekgras 53

Phacelurus Griseb. 30, 266, 388

Phacelurus franksae (J.M. Wood) Clayton 267 (fig. 165),

373 (pi. 151)

Phalaris L. 10, 29, 267, 381, 382
Phalaris angusta Nees ex Trin. 268
Phalaris aquatica L. 268 (fig. 166), 373 (pi. 152)
Phalaris arundinacea L. 34, 268
var. picta L. 268
Phalaris canariensis L. 269
Phalaris minor Retz. 269
Phalaris nodosa L. 268
Phalaris paradoxa L. 269
var. praemorsa Cross. & Dur. 269
Phalaris tuberosa L. 268
Phragmites Adans. 10, 29, 58, 269, 384, 399
Phragmites australis (Cav.) Steud. 1, 6, 7, 12, 269, 270
(fig. 167), 373 (pi. 153)

Phragmites communis Trin. 269
Phragmites mauritianus Kunth 7, 270
Pincushion grass 219
Pinhole grass 63

Plagiochloa Adamson & Spraque 339
Plagiochloa acutiflora (Nees) Adamson & Sprague 340
Plagiochloa alternans (Nees) Adamson & Sprague 340
Plagiochloa brachystachya (Nees) Adamson & Sprague
340

Plagiochloa ciliaris (Stapf) Adamson & Sprague 340
Plagiochloa oblitera (Hemsl.) Adamson & Sprague 341
Plagiochloa uniolae (L. f.) Adamson & Sprague
var. uniolae 341

var. villosa (Stapf) Adamson 341
Pluimsteekgras 50
Poa L. 29, 270, 382. 401
Poa annua L. 271 (fig. 168), 374 (pi. 154)

Poa atherstonei Stapf 271
Poa bidentata Stapf 272
Poa binata Nees 271, 272
Poa bulbosa L. 271
Poa divaricata Gouan 304
Poa heterogama Hack. 271
Poa leptoclada A. Rich. 272
Poa pratensis L. 271, 272
Poa trivialis L. 271, 272
Poa vivipara (L.) Willd. 271
Poagrostis Stapf 5, 253, 254, 384
Poagrostis pusilla (Nees) Stapf 262
Pogonarthria Stapf 29, 272, 385
Pogonarthria fleckii (Hack.) Hack. 272, 273
Pogonarthria leiarthra Hack. 273
Pogonarthria squarrosa (Roem. & Schult.) Pilg. 272, 273
(fig. 169), 374 (pi. 155)

Pointed russet grass 206

Polevansia De Winter 29, 274, 385

Polevansia rigida De Winter 274 (fig. 170), 374 (pi. 156)

Polgras 185

Polswenkgras 169

Polypogon Desf. 10, 12, 29, 33, 274, 382
Polypogon griquensis (Stapf) Gibbs Russell ined. 275
Polypogon minutiflorus Pilg. 275

Polypogon monspeliensis (L.) Desf. 4, 274 (fig. 171), 275,

374 (pi. 157)

Polypogon strictus Nees 276

Polypogon viridis (Gouan.) Breistr. 27, 275 (fig. 172), 276

Polypogon semiverticillatus (Forssk.) Hyl. 276

Potamophila prehensilis (Nees) Benth. 277

Predikantluis 74

Prickly brack grass 226

Prionanthium Desv. 29, 276, 384

Prionanthium dentatum (L.f.) Henr. 276

433

Prionanthium ecklonii (Nees) Stapf 276
Prionanthium pholiuroides Stapf 276 (fig. 173), 277, 374
(pi. 158)

Prionanthium rigidum Desv. 276
Pronkgras 250
Proso 235

Prosphytochloa Schweick. 29, 277, 383, 400
Prosphytochloa prehensilis (Nees) Schweick. 6, 277 (fig.

174) , 374 (pi. 159)

Pseudechinolaena Stapf 29, 277, 386
Pseudechinolaena polystachya (Kunth) Stapf 12, 278 (fig.

175) , 374 (pi. 160)

P seudohrachiaria Launert 5, 63, 65
P seudohrachiaria deflexa (Schumach.) Launert 65
Pseudobromus K. Schum. 5, 168
Pseudobromus africanus (Hack.) Stapf 169
Pseudobromus silvaticus K. Schum. 169
Pseudopentameris Conert 29, 278, 384
Pseudopentameris brachyphylla (Stapf) Conert 7, 278,
279 (fig. 176)

Pseudopentameris macrantha (Schrad.) Conert 279, 374
(pi. 161)

Psilochloa Launert 233, 241

Psilochloa pilgerana (Schweick.) Launert 241

Puccinellia Pari. 7, 29, 279, 382

Puccinellia acroxantha Smith & C.E. Hubb. 280, 374 (pi

162)

Puccinellia angusta (Nees) Smith & C.E. Hubb. 280

Puccinellia distans (L.) Pari. 280

Puccinellia fasciculata (Torr.) Bickn. 280 (fig. 177)

Purple brome 76

Purple finger grass 1 14

Purple hood grass 286

Purple plume grass 62

Purple spike grass 266

Pylgras 179

Quinine grass 349

Ratstail fescue 355
Red autumn grass 290
Red brome 76
Reddingsgras 74
Red honey grass 346
Red quick grass 97
Red Rhodes grass 168
Red swamp grass 178
Red top grass 67
Reed Canary grass 268
Reed panicum 236
Reengrassie 151
Rendlia Chiov. 29, 281, 385

Rendlia altera (Rendle) Chiov. 281 (fig. 178), 375 (pi.
163)

Rendlia nelsonii (Stapf) Chiov. 281
Rescue grass 74
Reuse kweekgras 96, 97
Reuse pylgras 337
Reuse terpentyngras 95
Rhodesian bluegrass 41
Rhodes grass 84
Rhodesiese andropogon 41
Rhynchelytrum Nees 5, 210, 386, 418
Rhychelytrum bellespicatum (Rendle) Stapf & C.E. Hubb
211 '

Rhynchelytrum brevipilum (Hack.) Chiov. 212
Rhynchelytrum costatum Stapf & C.E. Hubb. 212
Rhynchelytrum drakensbergense C.E. Hubb. & Schweick.
'211

Rhynchelytrum grandiflorum Hochst. 212
Rhynchelytrum kallimorphon Clayton 21 1
Rhynchelytrum longisetum (A. Rich.) Stapf & C.E. Hubb.

211

Rhynchelytrum minutiflorum (Rendle) Stapf & C.E. Hubb.

var. melinoides (Stent) Stapf & C.E. Hubb. 21 1
Rhynchelytrum nerviglume (Franch.) Chiov. 212

Rhynchelytrum nyassanum (Mez) Stapf & C.E. Hubb. 212
Rhynchelytrum ramosum Stapf & C.E. Hubb. 212
Rhynchelytrum repens (Willd.) C.E. Hubb. 212
Rhynchelytrum rhodesianum (Rendle) Stapf & C.E. Hubb.
212

Rhynchelytrum scabridum (K. Schum.) Chiov. 213
Rhynchelytrum setifolium (Stapf) Chiov. 212
Rhynchelytrum suberostratum Stapf & C.E. Hubb. 213
Rhynchelytrum subglabrum (Mez) Stapf & C.E. Hubb. 213
Rhynchelytrum villosum (Pari.) Chiov. 212
Rhytachne Desv. 30, 281, 388
Rhytachne latifolia Clayton 282
Rhytachne robusta Stapf 282 (fig. 179)

Rhytachne rottboellioides Desv. 283, 375 (pi. 164)
Ribbon bristle grass 297
Rice 1

Richmond and Wynberg finger grass 1 10

Rietgras 41, 57

Riffelblaarsetaria 297

Ripgut brome 74, 76

Riverbank pennisetum 249

River bushman grass 325

River grass 57

Robies Cocks foot 332

Rock bushman grass 322

Rock powder grass 102

Rock three awns 54

Rolling grass 242, 343

Rooidekgras 290

Rooigras 308, 335

Rooikweek 178

Rooisaadgras 348

Rooisinjaalgras 67

Rooivleigras 191

Rottboellia L.f. 1 1, 30, 283, 388, 400, 410
Rottboellia cochinchinensis (Lour.) Clayton 12, 283 (fig.

180), 375 (pi. 165)

Rottboellia exaltata L.f 283
Rough-leaved bushman grass 323
Roughstalk bluegrass 272
Ruigtegras 221
Ruspergras 174
Rye 1

Rye bushman grass 324
Rytidosperma 5, 213

Sacciolepis Nash 2, 29, 284, 386, 393, 415, 418
Sacciolepis africana C.E. Hubb. & Snowden 285
Sacciolepis auriculata Stapf 285
Sacciolepis chevalieri Stapf 285, 286
Sacciolepis cinereo-vestita ( Pilg. ) C.E. Hubb. 286
Sacciolepis curvata (L.) Chase 285
Sacciolepis glaucescens Stapf 286
Sacciolepis huillensis (Rendle) Stapf 285
Sacciolepis indica (L.) A. Chase 285, 286
Sacciolepis interrupta (Willd.) Stapf 285
Sacciolepis rigens (Mez) A. Chev. 285, 286
Sacciolepis typhura (Stapf) Stapf 284 (fig. 181), 285, 286,
375 (pi. 166)

Sandveldpluimgras 155
Sartidia De Winter 29, 286, 385

Sartidia angolensis (C.E. Hubb.) De Winter vii, 286 (fig.

182) , 287, 375 (pi. 167)

Sartidia jucunda (Schweick.) De Winter 287

Sartidia sp. (=Muller 2174) 287

Sawtooth love grass 161

Schismus Beauv. 29, 287, 384, 400

Schistous barbatus (Loefl. ex L.) Thell. 12, 287, 288 (fig.

183) , 375 (pi. 168)

Schismus inermis (Stapf) C.E. Hubb. 287
Schismus pleuropogon Stapf 288
Schismus scaberrimus Nees 288
Schizachyrium Nees 30, 223, 288, 387
Schizachyrium brevifolium (Swartz) Buese 289
Schizachyrium exile (Hochst.) Pilg. 289
Schizachyrium inclusum Stent 289

434

Schizachyrium jeffreysii (Hack.) Stapf 131, 289
Schizachyrium rupestre (K. Schum.) Stapf 289
Schizachyrium sanguineum (Retz.) Alst. vii, 179, 289
(fig. 184), 290, 335, 375 (pi. 169)

Schizachyrium semiberhe Nees 290
Schizachyrium ursulus Stapf 290
Schmidtia Steud. 29, 290, 385, 399
Schmidt ia bulhosa Stapf 291
Schmidtia kalihariensis Stent 291

Schmidtia pappophoroides Steud. 6, 136, 290 (fig. 185),
291, 375 (pi. 170)

Schoenefeldia Kunth 29, 291, 385

Schoenefeldia transiens (Pilg.) Chiov. vii, 291 (fig. 186),

375 (pi. 171)

Scleropoa Griseb. 78
Scleropoa rigida (L.) Griseb. 79
Seaside rush grass 313
Seashore paspalum 247
Secale L. 29, 132, 181, 292, 382

Secale africanum Stapf 292 (fig. 187), 376 (pi. 172)

Secale cereale L. 1, 292
Sedge-stemmed love grass 86
Sehima Forssk. 30, 292, 387

Sehima galpinii Stent 2, 293 (fig. 188), 376 (pi. 173)
Sehima ischaemoides Forssk. 2, 293
Sekelgras 273
Serrated tussock 224

Setaria Beauv. 2, 5, 29, 244, 293, 386, 415, 416
Setaria almaspicata De Wit 298
Setaria anceps Stapf ex Massey 298
Setaria appendiculata (Hack.) Stapf 7, 295. 298
Setaria cana De Wit 298

Setaria chevalieri Stapf ex Stapf & C.E. Hubb. 297
Setaria decipiens De Wit 299
Setaria eylesii Stapf & C.E. Hubb. 296
Setaria fiabellata Stapf
subsp .flabellata 299
subsp. natalensis De Wit 299
Setaria flabelliformis De Wit 298
Setaria Finita Launert 295
Setaria geniculata (Lam.) Beauv. 295
Setaria gerrardii Stapf 296
Setaria holstii Herr. 296
Setaria homblei De Willd. 299
Setaria homonyma (Steud.) Chiov. 295
Setaria incrassata (Hochst.) Hack. 6, 296, 297
Setaria insignis De Wit 297
Setaria italica (L.) Beauv. 295, 296
Setaria lindenbergiana (Nees) Stapf 296 (fig. 190), 297,

376 (pi. 174)

Setaria lutescens (Wiegel) F.T. Hubb. 249
Setaria megaphylla (Steud.) Dur. & Schinz 8, 297
Setaria neglecta De Wit 299

Setaria nigrirostris (Nees) Dur. & Schinz 296, 297
Setaria obscura De Wit 297

Setaria pahularis Stapf 296

Setaria pallide-fusca (Schumach.) Stapf & C.E. Hubb.

295, 297, 300
Setaria palustris Stapf 296
Setaria perberbis De Wit 296
Setaria perennis Hack. 299
Setaria phillipsii De Wet 296
Setaria phragmitoides Stapf 296
Setaria plicatilis (Hochst.) Engl. 296, 297
Setaria porphyrantha Stapf 296
Setaria pseudaristata (Peter) Pilg. 297
Setaria rigida Stapf 298
Setaria rudifolia Stapf 296

Setaria sagittifolia (A. Rich.) Walp. 7, 295, 298 (fig. 191)
Setaria sphacelata (Schumach.) Moss
subsp. aquamontana De Wit 299
subsp. nodosa De Wit 298
subsp. pyropea De Wit 298
var. sericea (Stapf) Clayton 298, 299
var. sphacelata 9, 294 (fig. 189), 298, 299
var. splendida (Stapf) Clayton 298, 299
var. stolonifera De Wit 299
var. torta (Stapf) Clayton 299, 376 (pi. 175)

Setaria splendida Stapf 299
Setaria stenantha Stapf 299
Setaria tenuiseta De Wit 297
Setaria torta Stapf 299
Setaria ustilata De Wit 295, 297 . 300
Setaria verticillata (L.) Beauv. 12. 298, 299 (fig. 192),
300. 376 (pi. 176)

Setaria woodii Hack,
subsp. bechuanica De Wit 296
var. fonssalutis De Wit 296
var. woodii 296
Shattercane 302
Sickle grass 93
Silky bushman grass 328
Silky grass 131, 250
Silver finger grass 108
Silverspike 190
Silver thread grass 40
Skurwe steekgras 53
Smalhartjie-eragrostis 158
Small Canary grass 269
Small carrot-seed grass 338
Small dropseed 308
Smooth brome 75
Snowflake grass 40
Soetkweek 97
Soft brome 75
Soft grass 180

Sorghastrum Nash 30. 300. 301, 387, 400
Sorghastrum friesii (Pilg.) Pile. 300 (fig. 193). 301. 376
(pi. 177)

Sorghastrum rigidifolium (Stapf) Chippind. 301
Sorghastrum stipoides (Kunth) Nash 7. 301
Sorghum Moench 30, 301, 300, 387, 400
Sorghum almum Parodi 302
Sorghum bicolor (L.) Moench 1
Sorghum bicolor (L.) Moench
subsp. arundinaceum (Desv.) De Wet & Harlan 302, 376
(pi. 178)

subsp. drummondii (Steud.) De Wet 302
Sorghum halepense (L.) Pers. 6, 7, 301 (fig. 194), 302
Sorghum sudanense (Piper) Stapf 302
Sorghum versicolor Anderss. 6, 302
Sorghum verticilliflorum (Steud.) Stapf 302
South African bent grass 34
Spaanseriet 58
Spade grass 198
Spanish brome 75
Spartina Schreber 29, 302, 385
Spartina capensis Nees ex Trin. 303
Spartina maritima (Curtis) Fernald 302 (fig. 195), 303,
376 (pi. 179)

Sphenopus Trin. 8, 29, 303, 382

Sphenopus divaricatus (Gouan) Reichb. 303 (fig. 196),
304, 376 (pi. 180)

Spiderweb chloris 84
Spike bushman grass 324
Spiny love grass 86

Sporobolus R. Br. 10, 11, 29, 146, 304, 385, 419
Sporobolus acinifolius Stapf 306. 312, 313
Sporobolus africanus (Poir.) Robyns & Tournay 304 (fig.

197), 306, 309, 310, 311
Sporobolus albicans Nees 306, 307, 312, 313
Sporobolus argutus (Nees) Kunth 308
Sporobolus artus Stent 309
Sporobolus baumianus Pilg. 313
Sporobolus bechuanicus Goossens 306, 312, 313
Sporobolus capensis (Willd.) Kunth 306
Sporobolus centrifugus (Trin.) Nees 307 (fig. 198), 310,
312, 377 (pi. 181)

Sporobolus congoensis Franch. 307, 310, 312
Sporobolus conrathii Chiov. 307, 313
Sporobolus consimilis Fresen. 308
Sporobolus cordofanus (Steud.) Coss. 308
Sporobolus coromandelianus (Retz.) Kunth 308, 309, 310
Sporobolus discosporus Nees 308 (fig. 199), 377 (pi. 182)
Sporobolus engleri Pilg. 308, 310, 312
Sporobolus eylesii Stent & Rattray 307

435

Sporobolus festivus A. Rich. 308, 310, 312
\ar.fibrosus Stent 308

Sporobolus fimbriatus (Trin.) Nees 306, 309, 377 (pi .
183), 409

var. latifolius Stent 309.

Sporobolus fourcadii Stent 306, 309
Sporobolus ioclados (Trin.) Nees 309, 312
var. usitatus (Stent) Chippind. 309
Sporobolus kentrophyllus (K. Schum.) Clayton 309
Sporobolus lampranthus Pilg. 310
Sporobolus ludwigii Hochst. 308, 309, 310
Sporobolus macranthelus Chiov. 309
Sporobolus macrothrix Pilg. 313
Sporobolus marginatus Hochst. ex A. Rich. 309
Sporobolus mauritianus (Steud.) Dur. & Schinz 307, 310,
312

Sporobolus molleri Hack. 310

Sporobolus natalensis (Steud.) Dur. & Schinz 306, 309,

310

Sporobolus nebulosus Hack. 7, 308, 310, 312
Sporobolus nervosus Hochst. 310
Sporobolus nitens Stent 7, 308, 309, 310, 31 1 (fig. 200)
Sporobolus panicoides A. Rich. 310
Sporobolus pectinatus Hack. 311
Sporobolus pellucidus Hochst. 311
Sporobolus pyramidalis Beauv. 157, 306, 309, 311
Sporobolus pyramidatus sensu Chippind., non (Lam.)
Hitchc. 308

Sporobolus rangei Pilg. 309, 312
Sporobolus rhodesiensis Stent & Rattray 312
Sporobolus robustus sensu Chippind., non Kunth 308
Sporobolus salsus Mez 306, 312, 313
Sporobolus sanguineus Rendle 307, 310, 312
Sporobolus schlechteri Schweick. 307
Sporobolus sladenianus Bol. f. 310
Sporobolus smutsii Stent 309
Sporobolus spicatus (Vahl) Kunth 312, 313
Sporobolus stapfianus Gand. 308, 310, 312
Sporobolus stolzii Mez 312

Sporobolus subtilis Kunth vii, 307, 312, 313 (fig. 201)

Sporobolus tenellus (Spreng.) Kunth 306, 312, 313

Sporobolus uniglumis Stent & Rattray 308

Sporobolus usitatus Stent 309

Sporobolus verdcourtii Napper 309

Sporobolus virginicus (L.) Kunth 160, 306, 307, 312, 313

Sporobolus welwitschii Rendle 313

Sporobolus sp. (-Smook 3429) 313

Spotted signal grass 67

Spreading prickle grass 48

Spreading steekgras 50

Squirreltail fescue 354

Star grass 96, 97

Steekgras 47

Steekriet 86, 226

Steekrietboesmangras 325

Stemless bushman grass 327

Stenotaphrum Trin. 29, 314, 386

Stenotaphrum dimidiatum (L.) Brongn. 314

Stenotaphrum secundatum (Walt.) Kuntze 6, 314 (fig.

202), 377 (pi. 184)

Stereochlaena Hack. 29, 315, 386

Stereochlaena cameronii (Stapf) Pilg. 315 (fig. 203), 377
(pi. 185)

Sterretjiegras 101
Stiburus Stapf 5, 29, 315, 385

Stiburus alopecuroides (Hack.) Stapf 316 (fig. 204), 377
(pi. 186)

Stiburus conrathii Hack. 316
Sticky love grass 163
Stingelgras 207
Stinkgras 148

Stipa L. 21, 29, 52, 224, 225, 316, 320, 384, 394

Stipa capensis Thunb. 316, 317
Stipa clandestina Hack. 317
Stipa dregeana Steud.
var. dregeana 169, 317 (fig. 205)
var. elongata (Nees) Stapf 317, 377 (pi. 187)

Stipa namaquensis Pilg. 320
Stipa neesiana Trin. & Rupr. 318
Stipa papposa Nees 318
Stipa parvula Nees 52
Stipa tenuissima Trin. 225, 318
Stipa tortilis Desf. 317
Stipa trichotoma Nees 224
Stipa variabilis Hughes 318

Stipagrostis Nees iii, 2, 4, 9, 11, 29, 318, 385, 413, 414
Stipagrostis amabilis (Schweick.) De Winter 320, 325
Stipagrostis anomala De Winter 320, 321 (fig. 207), 319,
378 (pi. 188)

Stipagrostis brevifolia (Nees) De Winter 321, 325
Stipagrostis capensis Nees 329
Stipagrostis ciliata (Desf.) De Winter
var. capensis (Trin. & Rupr.) De Winter 5, 319, 321, 327,
329

Stipagrostis damarensis (Mez) De Winter 321
Stipagrostis dinteri (Hack.) De Winter 322
Stipagrostis dregeana Nees 322
Stipagrostis fastigiata (Hack.) De Winter 322
Stipagrostis garubensis (Pilg.) De Winter 322
Stipagrostis geminifolia Nees 7, 322
Stipagrostis giessii Kers 323, 324
Stipagrostis gonatostachys (Pilg.) De Winter 323
Stipagrostis hermannii (Mez) De Winter 323, 327
Stipagrostis hirtigluma (Trin. & Rupr.) De Winter
subsp. hirtigluma 323, 324, 327, 328
subsp. patula (Hack.) De Winter 323, 324, 327
subsp. pearsonii (Henr.) De Winter 323, 324
Stipagrostis hochstetteriana (Beck ex Hack.) De Winter
var. hochstetteriana 323, 324
var. secalina (Henr.) De Winter 324
Stipagrostis lanipes (Mez) De Winter 324
Stipagrostis lutescens (Nees) De Winter
var. lutescens 324, 325
var. marlothii (Hack.) De Winter 324
Stipagrostis namaquensis (Nees) De Winter 320, 321,
322, 325 (fig. 208)

Stipagrostis namibensis De Winter 325
Stipagrostis obtusa (Del.) Nees 323, 324, 326 (fig. 209)
Stipagrostis proxima (Steud.) De Winter 326
Stipagrostis ramulosa De Winter 326
Stipagrostis sabulicola (Pilg.) De Winter 326
Stipagrostis schaeferi (Mez) De Winter 321, 327
Stipagrostis subacaulis (Nees) De Winter 323, 327
Stipagrostis uniplumis (Licht.) De Winter
var. intermedia (Schweick.) De Winter 327
var. neesii (Trin. & Rupr.) De Winter 323, 327, 328, 377
(pi. 189)

var. uniplumis 318 (fig. 206), 327, 328
Stipagrostis zeyheri (Nees) De Winter
subsp. barbata (Stapf) De Winter 328, 329
subsp. macropus (Nees) De Winter 321, 329
subsp. sericans (Hack.) De Winter 328 (fig. 210), 329
subsp. zeyheri 328, 329, 378 (pi. 190)

Stippelgras 63
Strandkoringgras 336
Strandkweek 303
Streblochaete Pilg. 29, 329, 382

Streblochaete longiarista (A. Rich.) Pilg. vii, 329 (fig.

211), 330, 378 (pi. 191)

Struikpanicum 235

Styppeiochloa De Winter 8, 29, 330, 384
Styppeiochloa gynoglossa (Goossens) De Winter 7, 330
(fig. 212)„ 378 (pi. 192)

Sudan grass 302
Suurbuffelsgras 249
Suurpol 131
Swamp grass 1 17
Swartsaadgras 302
Swaziland finger grass 1 10
Sweet signal grass 66
Sweet tanglehead 179
Sweet vernal grass 45
Sygras 190

436

Taaipol 306, 31 1
Taaipoleragrostis 157
Tall bushman grass 321
Tall three-awned grass 53
Tamboekiegras 95
Tanglehead 179
Tarentaalgras 283
Tarigidia Stent 29, 331, 386

Tarigidia aequiglumis (Goossens) Stent 43, 331 (fig. 2 1 3),
378 (pi. 193)

Teff 162

Tetrachne Nees 29, 331, 385

Tetrachne dregei Nees 332 (fig. 214), 378 (pi. 194)

Tetrapogon Desf. 29, 332, 385

Tetrapogon mossambicensis (K. Schum.) Chippind. ex.
Fisher 84

Tetrapogon tenellus (Roxb.) Chiov. 332, 333 (fig. 215),
378 (pi. 195)

Thamnocalamus Munro 1 1, 29, 333, 383
Thamnocalamus tessellatus (Nees) Soderstrom & Ellis 7,
333 (fig. 216), 378 (pi. 196)

Thelepogon Roem. & Schult. 30, 334, 387
Thelepogon elegans Roem. & Schult. vii, 334 (fig. 217),
378 (pi. 197)

Themeda Forssk. 10, 30, 334, 387, 409, 416
Themeda triandra Forssk. 4, 7, 8, 1 79, 290, 335 (fig. 2 1 8),

378 (pi. 198), 379 (pi. 199)
var. burchellii (Hack.) Stapf 335
var. hispida (Nees) Stapf 335

var. imberbis (Retz.) A. Camus 335
var. trachyspathea Goossens 335
var. vulgaris auctt., non Hack. 335
Thimble grass 171

Thinopyrum A. Loeve 29, 132, 336, 382
Thinopyrum distichum (Thunb.) Loeve 336 (fig. 219),

379 (pi. 200)

Thunberg’s pennisetum 250
Thread-leaved andropogon 1 15
Three-awned rolling grass 48
Tick grass 150
Torro-boesmangras 320
Towoomba Canary grass 268
Trachypogon Nees 30, 336, 387

Trachypogon spicatus (L. f.) Kuntze 6, 179, 293, 337 (fig.

220), 349, 379 (pi. 201)

Trachypogon capensis (Thunb.) Trin. 337
Tragus Haller 29, 337, 385

Tragus berteronianus Schult. 338 (fig. 221), 379 (pi. 202)
Tragus koelerioides Aschers. 338
Tragus pedunculatus Pilg. 338
Tragus raeemosus (L.) All. 12, 338
Transvaalkweek 97
Transvaal quick grass 97
Tribolium Desv. 5, 29, 339, 384
Tribolium acutiflorum (Nees) Renvoize 340
Tribolium alternans (Nees) Renvoize 340
Tribolium amplexum Renvoize 340
Tribolium brachystachyum (Nees) Renvoize 340
Tribolium ciliare (Stapf) Renvoize 340
Tribolium echinatum (Thunb.) Renvoize 340
Tribolium hispidum (Thunb.) Renvoize 340
Tribolium obliterum (Hemsl.) Renvoize 341
Tribolium obtusifolium (Nees) Renvoize 341
Tribolium uniolae (L. f.) Renvoize 339 (fig. 222), 341, 379
(pi. 203)

Tribolium utriculosum (Nees) Renvoize 341
Tricholaena Schrad. 29, 341, 386

Tricholaena arenaria Nees 341
var. glauca (Hack.) Stapf 341
Tricholaena capensis (Licht. ex Roem. & Schult.) Nees
subsp. arenaria (Nees) Zizka 341, 342
subsp. capensis 342

Tricholaena monachne (Trin.) Stapf & C.E. Hubb. 342
(fig. 223), 379 (pi. 204)

Trichoneura Anderss. 29, 342, 385
Trichoneura eleusinoides (Rendle) Ekman 343, 344

Trichoneura grandiglumis (Nees) Ekman 4, 12, 343 (fig.
224), 379 (pi. 205)
var. grandiglumis 343
var. minor 343

Trichoneura schlechteri Ekman 344
Trichoneura sp. (=Codd 5325) 344
Trichopteryx Nees 29. 344, 387

Trichoptervx dregeana Nees 7, 344 (fig. 225), 379 (pi.
206)

Trichopteryx ramosa Stapf 102
Trident grass 348

Tripogon Roem. & Schult. 29, 345. 385
Tripogon abyssinicus sensu Chippind., non Nees 345
Tripogon minimus (A. Rich.) Steud. 345 (fig. 226), 379
(pi. 207)

Triraphis R. Br. 29. 345, 385

Triraphis andropogonoides (Steud.) Phill. 346 (fig. 227),
347, 380 (pi. 208)

Triraphis elliottii Rendle 347
Triraphis fleckii Hack. 346
Triraphis pumilio R. Br. 346
Triraphis purpurea Hack. 346
Triraphis ramosissima Hack. 347
Triraphis schinzii Hack. 346, 347
Triraphis schlechteri Pilg. ex Stent 347
Triraphis welwitschii Rendle 346
Trisetaria Forssk. 204
Trisetaria pumila (Desf.) Maire 205
Trisetum pumilum (Desf.) Kunth 205
Tristachya Nees 9, 29, 206, 347. 387
Tristachya biseriata Stapf 348
Tristachya eylesii Stent & Rattray 348
Tristachya hitchcockii (C.E. Hubb.) Conert 348
Tristachya hispida (L. f.) K. Schum. 348
Tristachya leucothrix Nees 347 (fig. 228), 348, 380 (pi.
209)

Tristachya longispiculata C.E. Hubb. 348
Tristachya lualabaensis (De Wild.) J.B. Phipps 348
Tristachya nodiglumis K. Schum. 348
Tristachya rehmannii Hack. 348

Tristachya superba (De Not.) Schweinf. & Aschers. 348

Triticum aestivum L. 1

Tropical finger grass 108

Tuinsetaria 297

Tumble weed 242

Turfgras 191

Tussock fescue 169

Tweevingergras 40, 191

Twisted leaf bristle grass 299

Urelytrum Hack. 30, 349, 388, 400
Urelytrum agropyroides (Hack.) Hack. 9, 179, 293, 337,
349 (fig. 229), 380 (pi. 210)

Urelytrum squarrosum Hack. 349
Urochlaena Nees 29, 350, 384

Urochlaena pusilla Nees 12, 350 (fig. 230), 380 (pi. 21 1)
Urochloa Beauv. 5, 29, 350, 386
Urochloa holbodes (Steud.) Stapf 351
Urochloa brachyura (Hack.) Stapf 351, 352
Urochloa engleri Pilg. 352

Urochloa mosambicensis (Hack.) Dandy 9, 351 (fig. 231 ),
352, 380 (pi. 212)

Urochloa oligotricha (Fig. & De Not.) Henr. 351

Urochloa panicoides Beauv. 6, 352

Urochloa pullulans Stapf 351

Urochloa rhodesiensis Stent 351

Urochloa ruschii sensu Chippind., non Pilg. 352

Urochloa stolonifera (Goossens) Chippind. 351, 352

Urochloa trichopus (Hochst.) Stapf 351, 352

Vaalgras 291
Vaalsteekgras 48
Varkstertgras 349
Vasey grass 247
Velvet grass 180
Vertakte borseltjiegras 43

437

Vetiveria Bory 30, 352, 387

Vetiveria nigritana (Benth.) Stapf 352 (fig. 232), 353, 380
(pi. 213)

Vetiveria zizanioides (L.) Nash 353

Vingerhoedgras 171

Vinkagrostis 34

Vlei finger grass 105

Vleigras 308, 311, 344

Volstruisdoring 86

Vossia Wall. & Griff. 30, 353, 388

Vossia cuspidata (Roxb.) Griff, vii, 353 (fig. 233), 380 (pi.

214)

Vulpia C. Gmel. 12, 29, 168, 354, 381, 382

Vulpia bromoides (L.) S.F. Gray 354, 355

Vulpia fasciculata (Forssk.) Samp. 355

Vulpia muralis (Kunth) Nees 354, 355

Vulpia myuros (L.) C. Gmel. 354 (fig. 234), 355, 380 (pi.

215)

Waaigras 343
Watergras 246
Water grass 121
Weeping love grass 149
Weeluiseragrostis 161
Wheat 1

White buffalo grass 236
Whorled finger grass 112
Wildebeestegras 131
Wildegraansorghum 302

Wildehawergras 223
Wilderog 292
Wild oatgrass 223
Wild rice 229
Wild rye 292

Willkommia Hack. 29, 274, 355, 385

Willkommia annua Hack. 356, 380 (pi. 216)

Willkommia newtonii Hack. 356

Willkommia sarmentosa Hack. 355 (fig. 235), 356

Wire grass 51, 131

Wiry signal grass 65

Witbuffelgras 236

Withaargras 200

Witkafferkoring 302

Witsteekgras 48

Wolvoet panicum 239

Wondergras 135

Wool grass 43

Woolly bushman grass 324

Woolly leaved bushman grass 327

Woolly love grass 148

Woolly russet grass 206

Wurmsinjaalgras 67

Yellow thatching grass 189
Yorkshire fog 180
Ystergras 49

Zea mays L. 1,5, 7, 10, 1 1

while the years have lengthened and the grass has grown.

J.R.R. Tolkein

MEMOIRS OF THE BOTANICAL SURVEY OF SOUTH AFRICA
MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

The following memoirs are out of print:/Die volgende memoirs is uit druk: Nos. 3, 4, 6, 9, 10, 14, 15, 16, 20-22, 25, 26, 28, 30, 35-38
and/en 40. Still available are:/Nog beskikbaar is:

1. Phanerogamic flora of the Divisions of Uitenhage and Port Elizabeth. 1919. S. Schonland. 40 c.

2. Botanical survey of Natal and Zululand. 1921. R.D. Aitken & G.W. Gale. 20 c.

5. Researches on the vegetation of Natal. 1923. J.W. Bews & R.D. Aitken. 40 c.

7. The native timber trees of the Springbok Flats. 1925. E.E. Galpin. 40 c.

8. Researches on the vegetation of Natal. 1925. J.W. Bews & R.D. Aitken. 40 c.

11. A revised list of plant diseases occurring in South Africa. 1931. E.M. Doidge & A.M. Bottomly. 60 c.

12. Botanical survey of the Springbok Flats (Transvaal). E.E. Galpin. 40 c.

13. The vegetation of the Riversdale Area, Cape Province. 1929. J. Muir. 40 c.

17. The vegetation of the Division of Albany and Bathurst. 1937. R.A. Dyer. 40 c.

18. Notes on the vegetation of the Kamiesberg. 1938. R.S. Adamson. 40 c.

19. The value of botanical survey and the mapping of vegetation as applied to farming systems in South Africa. 1938. J.A. Pentz. 40 c.

23. The vegetation of Weenen County, Natal. 1951. O. West. 80 c.

24. An ecological account of the vegetation of the Potchefstroom Area. 1951. W.J. Louw. 60 c.

27. A botanical survey of the Keiskammahoek District. 1951. R. Story. R 1 , 10 ; overseas Rl,80.

29. The wheel-point method of survey. 1955. C.E.M. Tidmarsh & C.M. Havenga. 80 c; overseas Rl,50.

31. Studies of the vegetation of parts of the Bloemfontein and Brandfort Districts. 1958. J.W.C. Mostert. R2,00; overseas R3,30.

32. An account of the plant ecology of the Table Mountain area of Pietermaritzburg, Natal. 1959. D.J.B. Killick. R2,00; overseas R3,30.

33. The vegetation of the Districts of East London and King William’s Town, Cape Province. 1962. D.M. Comins. R2,45; overseas R3,60.

34. An account of the plant ecology of the Cathedral Peak area of the Natal Drakensberg. 1963. D.J.B. Killick. R3,00; overseas R3,80.

39. Flora of Natal. 1973. J.H. Ross. R4,30; overseas R5.45.

41. The biostratigraphy of the Permian and Triassic. Part 3. A review of Gondwana Permian palynology with particular reference to the northern
Karoo Basin, South Africa. 1977. J.M. Anderson. R5.00; overseas R6,50.

42. Vegetation of Westfalia Estate on the north-eastern Transvaal escarpment. 1977. J.C. Scheepers. R8,00; overseas R9,95.

43. The bryophytes of southern Africa. An annotated checklist. 1979. R.E. Magill & E.A. Schelpe. R7,00; overseas R8,70.

44. A conspectus of the African Acacia species. 1979. J.H. Ross. R15.20; overseas R17,00.

45. The plant ecology of the Isipingo Beach area. Natal, South Africa. 1980. C.J. Ward. R5,60; overseas R7,00.

46. A phytosociological study of the Upper Orange River Valley. 1980. M.J.A. Werger. R3,80; overseas R4,70.

47. A catalogue of South African green, brown and red algae. 1984. S.C. Seagrief. R5,10; overseas R6,40.

48. List of species of southern African plants. 1984. G.E. Gibbs Russell, the staff of the National Herbarium & P. Gonsalves. R5,00; overseas
R6.20.

49. Pattern analysis in savanna-woodlands at Nylsvley, South Africa. 1984. R.H. Whittaker, J.W. Morris & D. Goodman. R3,20; overseas R4,00.

50. A classification of the mountain vegetation of the Fynbos Biome. 1985. B.M. Campbell. R7,60; overseas R9,60.

51. List of species of southern African plants. Edn 2, Part 1. 1985. G.E. Gibbs Russell, C. Reid, J. van Rooy & L. Smook. R5,30; overseas R6,30.

52. A plant ecological bibliography and thesaurus for southern Africa up to 1975. 1986. A.P Backer, D.J.B. Killick & D. Edwards. R15.50;
overseas R19,40.

53. A catalogue of problem plants in southern Africa, incorporating the National Weed List of South Africa. 1986. M.J. Wells, A. A. Balsinhas,
H. Joffe, V.M. Engelbrecht, G. Harding & C.H. Stirton. R20,10; overseas R25,20.

54. Biomes of southern Africa — an objective categorization. 1986. M.C. Rutherford & R.H. Westfall. R4,35; overseas R5,40.

55. Barrier plants of southern Africa. 1987. L. Henderson. R7,00; overseas R8,50.

56. List of species of southern African plants. Edn 2, Part 2. 1987. G.E. Gibbs Russell, W.G. Welman, E. Retief, K.L. Immelman, G. Germishuizen,
B.J. Pienaar, M. van Wyk & A. Nicholas. R20,85; overseas R26.00.

57. Veld types of South Africa. 3rd edn. 1988. J.P.H. Acocks. R6,05; overseas R7,55. Map: R2,60; overseas R3,25.

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ISBN 0 620 14846 2

© and published by the National Botanic Gardens/Botanical Research Institute, Private Bag X101, Pretoria 0001, South Africa. Obtainable from
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