Grasses of Hawaii 9780824885182

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GRASSES OF HAWAII

GRASSES OF HAWAII

PETER P. ROTAR

UNIVERSITY OF HAWAII PRESS HONOLULU,

1968

HAWAII

COPYRIGHT 1 9 6 8 BY THE UNIVERSITY OF HAWAII PRESS

Library of Congress Catalog Card No. 68-13888

Printed in the United States of America Layout and composition by Cathay Press, Hong Kong

TO MEREDITH

PREFACE T h i s treatise is intended to fill the need for a modern, up-to-date classification of all native and introduced grasses (excluding bamboos) in Hawaii. T h e last comprehensive work on this subject (Whitney, Hosaka, and Ripperton, 1939) is outdated and in need of revision, since many introductions have been made since that time, and there have been a number of recent changes in the taxonomy of grasses. Research for this study was completed in March, 1965. Every attempt has been made to include in this listing all of the known introductions as well as the native grasses, but it is probable that a number of accessions have been inadvertently omitted, and a great deal of argument is anticipated over the synonymy and placement of genera and species in the different tribes. Likewise, among the introductions a number of items are recorded for which there are no living specimens in existence. It is possible that some of the missing species may exist in isolated localities, but it is assumed that most of them were completely unadapted to the introduction site and have failed to survive. Still others may have been wrongly named because of lack of sufficient information. M a n y of the grasses found in Hawaii at present are of little or no economic value and of only academic interest, although they may form an appreciable component of the prevailing land cover. A few are definitely important as objectionable weeds or useful as animal forage, turf cover, ornamentals, and industrial materials. N o listing is attempted of the many hybrids which have been introduced by the Hawaiian Sugar Planters' Association. M a n y of the latter are reported only as generic hybrids with no indication of the species involved. Some of the best established introductions are the least desirable and of these many are noxious weeds. A m o n g the weedy species notable examples are Andropogon virginicus (broom sedge), Cenchrus echinatus (sandbur), Panicum repens (torpedograss), Setaria geniculata (yellow foxtail), Sporobolus africanus (rattail), and Trichachne insularis (sourgrass). Some weedy grasses are on the noxious weed list of the State of Hawaii and their introduction is prohibited; among these are Avenafatua (wild oats), Imperata cylindrica, and Sorghum halepense (johnsongrass). A m o n g the distinctly beneficial grasses are Cynodon dactylon (bermudagrass), Digitaria decumbens (pangolagrass), Pennisetum clandestinum (kikuyugrass), Saccharum officinarum (sugar cane), Zea mays (maize), and Zoysia sp.

viii

GRASSES OF H A W A I I

Introduction of seeds and live vegetative parts of all plants including grasses is strictly controlled by the State Quarantine Board, as a means of excluding undesirable plants, plant diseases, and parasites. It is deplored that because of the ease of bringing in cuttings and seeds there is a constant danger that undesirable or harmful material will be introduced either purposely or accidentally through ignorance of the harmful potentials. In fact this is the way by which most of the current biological pests have gained entry. Although Hawaii's major islands comprise a mere 6,400 square miles of land located in the middle of the Pacific Ocean, Hawaii has received via man's, at times, thoughtless efforts more than 500 species of grasses representing over 100 genera. About 60 species are either native to Hawaii or were introduced by the early Hawaiians. T h e native grasses have not been the best adapted for pastures, and with the development of an extensive livestock industry, many introductions have been made by private individuals and public agencies. Other introductions have been made for the purpose of establishing ornamentals and for lawns and turf. In the preparation of this list all previous works and available information on grasses in Hawaii have been consulted and included where applicable. Major sources of material were publications of the Hawaii Agricultural Experiment Station; records of accessions and herbarium collections of the Department of Agronomy and Soil Science, College of Tropical Agriculture, University of Hawaii; records and herbarium collections of the Bernice P. Bishop Museum; the records of the Hawaii Board of Agriculture; the records of the United States Soil Conservation Service; the records of the Hawaiian Sugar Planters' Association; and the observations of private individuals, scattered through records on Hawaiian flora. The references used in the preparation of the taxonomic descriptions, keys and listings are presented in the bibliography. I wish to acknowledge the aid of Dr. N. L. Bor, formerly Assistant Director of the Royal Botanic Gardens, Kew, and Dr. C. E. Hubbard, Deputy Director of the Royal Botanic Gardens, Kew for their valuable time and generous assistance in reviewing the manuscript. M y debt to them both and to Dr. Bor in particular, for his permission to use his text as a guide, cannot be properly expressed. I am indebted to my colleague, Dr. George Gillett, Department of Botany, University of Hawaii, for his valuable assistance during the preparation of the manuscript. Without his suggestions and encourage-

ix

PREFACE

ment this project would never have been completed. Thanks are due to Dr. C. H. Lamoureux, Department of Botany, University of Hawaii, and to Dr. Otto Degener and Dr. Isa Degener, Mokuleia Beach, Waialua, Hawaii, for their aid in reviewing the manuscript. I wish to thank the Soil Conservation Service of the United States Department of Agriculture, Honolulu, Hawaii, for help in preparing the lists of introduced species; the Bernice P. Bishop Museum for permission to examine herbarium collections; Dr. G. D. Sherman, Associate Director, Hawaii Agricultural Experiment Station, for permission to undertake this project along with my regular duties; the University of Hawaii Research Council for funds to help defray costs of typing the manuscript ; and the Hawaii Agricultural Experiment Station for support in its publication. Drawings used to illustrate the manuscript were obtained from a number of sources. Acknowledgements are due to Dr. N. L. Bor, to the Secretary of State for Commonwealth Relations, England, and to the Ministry of Overseas Development, England, for permission to reprint illustrations from N. L. Bor, The Grasses of Burma, Ceylon, India and Pakistan; to the Regents of the University of California for permission to reprint illustrations from H. L. Mason's A Flora of the Marshes of California, to the Board of Trustees of the Grasses and Pastures of South Africa Book Fund for permission to reprint illustrations from the book edited by D. Meredith, The Grasses and Pastures of South Africa; to the Bishop Museum for permission to reprint illustrations from A. S. Hitchcock's The Grasses of Hawaii, to Dr. Otto Degener for permission to reprint illustrations from Flora Hawaiiensis; and to the United States Department of Agriculture for permission to reprint a number of their illustrations. Several illustrations are reprinted from Hawaii Agricultural Extension Service circulars and still others are from original drawings in the files of the Department of Agronomy and Soil Science, College of Tropical Agriculture, University of Hawaii. Special thanks are due to two undergraduate assistants, Jane Kuebitz and Betsy Lee Walton, who helped check references and proofread sections of the manuscript as they were completed. PETER P .

ROTAR

Honolulu, Hawaii

TABLE OF CONTENTS

The Grass Plant

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Taxonomy

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20

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23

Common Names, Origins, and Dates of First Record of Hawaiian Grasses .. .. .. .. .. ..

312

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Page Index to Tribes and Genera Gramineae

Glossar

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Bibliography Index

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343 345

THE GRASS PLANT Grasses are cosmopolitan. They dominate the vegetation over great expanses of the earth's surface and are versatile in their adaptation to habitat. They grow in water and in deserts, at sea level and on high mountain peaks, in dense tropical forests and on treeless plains. Wherever flowering plants can live, grasses will survive.

Duration and habit T h e duration of a plant is that length of time which represents its life span. T h e habit of a plant is the form of growth. Most grasses are either ANNUAL or PERENNIAL in duration. A n ANNUAL is a plant that completes

its life cycle within one year, whereas a PERENNIAL lives for more than two years. A few grasses are BIENNIAL, that is, they complete their life cycles in two years. In annual grasses, all or most of the shoots bear flower heads, and rhizomes and/or stolons are not developed. Perennial grasses have few to many leafy, vegetative shoots at the base of the plant, and rhizomes and/or stolons are developed frequently. T h u s perennials are different from annuals in habit, and the expression 'annual (or perennial) in habit* is occasionally used. T h e terms annual, biennial, and perennial are misleading in those geographic areas in which growth is not limited by a definite cold or dry period. In Hawaii a given species may grow as an annual or as a perennial in two different locations only a few miles apart. Because of restricted rainfall in one area it may grow as an annual; in the other area, with adequate moisture, it may live for several years and would then be classified as a perennial. Unless this is understood, one can become hopelessly confused in trying to classify the life span of a plant. T h e sterile shoots in perennial grasses form loose or dense bunches or tufts, and the plant is said to be TUFTED or CAESPITOSE. T h e word is applied also to annuals which have their stems in clusters.

Parts of the grass plant In describing the morphology of grasses, Bor (i960) states: 'The amazing diversity of shapes, sizes, texture and so on in the various parts that make up the vegetative and the reproductive shoot is alike the admiration and exasperation of the taxonomist. It can be said, I think, without fear of contradiction that in no other family of monocotyledons, or dicotyledons for that matter, has a relatively simple structure been so

2

GRASSES OF HAWAII

altered in the process of evolution as to produce the amazing wealth of forms known to us. It is as if Evolution, in an excess of exuberance, had added this, subtracted that, enlarged this, suppressed that, in fact tried out every possible combination and permutation of characters to produce a family as complicated and as difficult as, if not more difficult than, any other.' For an excellent discussion of this complexity, the reader is referred to Bor's study of the morphology of the grass plant. A typical grass plant consists of roots and shoots, the latter consisting of stems, leaves, and flower heads. [Fig. i] ROOTS

Grass roots are FIBROUS. They occur at the base of the plant and sometimes at the joints of the stems. STEMS

Besides the axis of the flower head, and the sterile, leafy shoots in perennials, three kinds of stems occur in grasses: CULMS, RHIZOMES, and STOLONS.

Culm T h e main aerial stem in grasses is known as the CULM; it is the axis on which the flower head as well as the leaves are borne. T h e culm is a cylindrical or somewhat flattened stem with one or more joints known as NODES. T h e part between each pair of nodes is known as an INTERNODE. T h e nodes are solid. Many tropical grasses have solid culms; temperate region grasses have hollow culms. In a few grasses, the basal internode is swollen and tuberous. In some cases the culm is SIMPLE (unbranched below the flower head), in others it is more or less BRANCHED. T h e branches arise at the nodes and produce shoots with or without flower heads. T h e culm may grow upright and straight (erect); it may be slightly or conspicuously bent at the nodes (geniculate and ascending); its lower part may lie along the ground and the upper part be erect (decumbent); or it may lie flat on the ground for the greater part of its length (procumbent or prostrate). Decumbent and prostrate culms frequently give rise to roots at those nodes in contact with the ground. Occasionally, roots develop at the lowest or lower nodes of erect culms, and are called proproots or stiltroots. Maize is a good example of a grass with proproots.

stigmas -AWA/

AMTHER-

flLAHEHr-

WARr -LEMMA LODICOLE'

HACHILLA•

AST.

GLUME

PEDUNCLE LÌGULE" PEDICEL SHEATH-

RHIZOME"

SrOLOA/

FIG. I. Characteristic grass plant parts. A, Flowers in a spikelet arranged on a central axis enclosed in two empty glumes. B, The grass flower. C , The developed caryopsis or grain. The grain is enclosed by the outer glumes, with the lemma both closely adhering and free. D, Spikelets arranged in a terminal spike. E, Spikelets arranged in a panicle. F, Spikelets in a raceme. G, A ligule, at the junction of the leaf blade and leaf sheath. H, /, Means by which the grass plant may be propagated or spread: stolon, rhizome, and bulb, respectively. (Photograph from U.S.D.A. Yearbook, 1948. Grasses.)

4

GRASSES

OF

HAWAII

Rhizomes and stolons Rhizomes and stolons are modified stems that in most cases provide the means by which grasses spread. Grasses with rhizomes or stolons, or both, tend to form a sward or turf rather than tufts or bunches. RHIZOMES are stems that occur below the surface of the ground. T h e y are distinguished from roots by being jointed and by having reduced, bladeless leaves (scales) arising from the joints. Rhizomes are frequently branched and usually creep, either slightly or extensively, their presence or absence and their size depending on the age of the plant or conditions of growth. S T O L O N S are stems which creep along—not below—the surface of the ground. T h e y are basal branches of the culm and develop roots and shoots at some or all of the nodes. T h e y may be confused with decumbent or prostrate culms which root from the nodes. T h e reduced bladeless leaves of rhizomes are brownish or translucent, whereas the leaves from stolons are green and have blades; stolons are usually green or purplish, while rhizomes are normally whitish or yellowish. LEAVES

Grass leaves are of two kinds: ( i ) green leaves consisting of a sheath and blade, and (2) variously colored, reduced leaves consisting of a sheath only. T h e ordinary vegetative leaves are borne singly at the nodes (with some exceptions) on alternate sides of the culms. T h i s arrangement is sometimes difficult to see because of the shortening of the culm internodes and consequent overlapping of the leaves. Those at the base of the culm are known as basal leaves, those scattered along the culm as cauline leaves. Reduced leaves of at least one kind occur in all grasses, as they form the bracts which enclose the flowers (glumes, lemmas, and paleas). In a few grasses all or most of the vegetative leaves are replaced by bladeless sheaths, while many grasses have one or two kinds of bladeless sheaths in addition to perfect leaves. These are scales (scalelike leaves, developed at the base of the culm and on rhizomes) and spathes (reduced leaves occurring on the flower head, especially in the Andropogoneae). T h e leaves are marked by few to many longitudinal parallel veins. When these are conspicuous and prominent, the leaves are striate; when they are raised, the leaves are ribbed. In some grasses, especially bamboos and forest and aquatic grasses, there are numerous short transverse

T H E GRASS

PLANT

5

veins between the longitudinal veins. In a few tropical forest grasses (for example, Pharus and Leptaspis) the main veins run obliquely from the base of the midnerve to the margins. A perfectly developed grass leaf consists of a lower portion known as the LEAF SHEATH, and an upper portion known as the LEAF BLADE. On the inner (upper) surface of the leaf there is nearly always, between the sheath and the blade, an outgrowth known as the LIGULE. The zone at the junction of the sheath and blade on the outer (lower) surface is known as the COLLAR. Leaf sheath The leaf sheath protects the young shoot. It encircles the culm where it arises at the node and often for its entire length. Sometimes successive sheaths overlap and are longer than the internodes, so that the culm is hidden. Overlapping basal sheaths, especially when they are flattened and keeled, are often arranged in a fan-shaped manner and are flabellate. In most grasses, the margins of the sheath overlap without being joined to each other, but in a few, for example species of Bromus, they are joined (connate), so that the sheath is like a tube. Tubular sheaths are found in Glyceria, Melica, Festuca, and a number of other genera, particularly of temperate region grasses. The marginal zone of the sheath where it joins the blade is known as the sheath mouth. In some grasses the margins at the apex produce appendages that are often ear-shaped and are called AURICLES. Leaf blade The blade is the portion of the leaf above the sheath. In most grasses it is linear in shape, that is, several times longer than wide and with the margins parallel to each other except, usually, towards the apex and sometimes the base. In a number of grasses, however, the leaf blade is lanceolate, being widest in the lowest third and gradually narrowed upwards. A few grasses, especially those that grow in shady places, have ovate blades, and intermediate shapes between linear, lanceolate, and ovate frequently occur. In some grasses the blade is widened into a rounded lobe on either side at the base, a condition referred to as cordate. Some grasses produce a pointed appendage at the base of the blade at each side; such a base is termed saggitate.

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GRASSES OF

HAWAII

T h e blade margins are sometimes thickened, in which case they are frequently crinkled as well. T h e y may be smooth to the touch, or rough (SCABRID) due to the presence of a fringe of minute, spiny hairs. In other species the blade margins are fringed with glands. While still enclosed in the bud, leaf blades are either folded or rolled. When they emerge, they may: (a) open out, a condition called EXPANDED ; (b) remain compressed with the midrib forming a sharp, longitudinal keel, and the margins pressed together or at least almost meeting— referred to as FOLDED; (C) remain ROLLED, with one margin wrapped around the other (convolute), or with the margins rolled inwards towards each other (involute). W h e n blades are wiry and bristle-like they are SETACEOUS; when thread-like they are FILIFORM. Setaceous and filiform blades are usually rolled or folded, rarely expanded. Prophyll T h e prophyll (prophyllum) is a 2-keeled scalelike leaf occurring between the culm and an axillary bud or shoot. It is sometimes much reduced. Ligule T h e ligule consists of either a flap of tissue known as a membrane or a fringe of hairs. All gradations between the two occur. Membranous ligules are either thin, skinlike, translucent and white, or they are firmer in texture, papery, dry, opaque and brownish. T h e ligule, whether a membrane or a fringe of hairs, is greatly reduced in some grasses, and may be represented by a short and inconspicuous rim. Rarely, it is absent, with the junction between the sheath and blade being marked by a darker colored zone. Collar T h e collar in most grasses is marked by a slightly discolored rather inconspicuous zone on the outer (lower) surface of the leaf at the junction of sheath and blade. It may be marked by a line or beard of hairs. INFLORESCENCE

(FLOWER HEAD)

T h e flower head bears the flowers and later the seed. In grasses it is known as the INFLORESCENCE which terminates the culm and the flowering branches of the culms. T h e inflorescence is made up of smaller units known as SPIKELETS (rarely a single spikelet), borne on one or more axes. There is a wide range of variation in structure, size, and shape.

T H E GRASS

PLANT

7

When the spikelets are carried on the divisions of a branched axis, the inflorescence is known as a PANICLE. When they are carried directly on an unbranched axis, they form a SPIKE if they are SESSILE (without stalks), or a RACEME if all or some of them are PEDICELLED (stalked). T h e unbranched axis is known as the RACHIS. A panicle, then, is an inflorescence having a central axis with few to many branches that are often profusely divided. T h e panicle may be open and loose, or contracted, or spikelike and dense. A spikelike, dense panicle is often known as a 'false spike' especially when the branches are so short that they are concealed by the spikelets. In some grasses, the panicle is made up of a number of false spikes. The divisions of the primary (main) branches are short, and the spikelets are crowded together on them. Often the branches are undivided and without spikelets on the lower part, such that they form stalks to the false spikes. These units are sometimes called 'peduncled false spikes.' T h e primary branches of the panicle are arranged on the central axis either singly (described as solitary), or in pairs or groups. When they are in groups with those of each group in a circle around the axis, they are said to be in WHORLS or WHORLED. In the same panicle, some branches may be in whorls while others are solitary or in pairs. In many grasses the inflorescence is made up of one or more spikes or racemes arranged in one of the following ways: (i) a single (solitary) spike or raceme terminates the culm; (2) a single pair of spikes terminates the culm or one of the pair is placed some distance below the other; or (3) there are three or more spikes or racemes arranged either (a) DIGITATELY, that is, radiating like the fingers of the hand in a whorl which terminates the culm (a DIGITATE inflorescence) or (b) along a central axis, singly, in pairs, or in groups. In some grasses, especially genera of the Andropogoneae, the inflorescence is composed of a large number of racemes, arranged either singly or in pairs, and often with spathes at the base. These may be small and inconspicuous or enlarged and brightly colored. T h e whole inflorescence is known as a spatheate panicle. Racemes occur more frequently than spikes, but they are often spikelike, since the spikelet stalks (pedicels) are short and inconspicuous and the racemes resemble spikes. When the inflorescence is made up of two or more racemes, which do not always bear spikelets right from the base, the lower part of the rachis forms a stalk or peduncle.

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GRASSES OF H A W A I I

Spikelets are borne either singly on the rachis (solitary) or in pairs or clusters. When they are in pairs, those of each pair are usually either unequally pedicelled, or one of the pair is sessile, the other pedicelled. The spikelets are often borne on one side of the rachis only, and are then usually arranged in one, two, or four regular ranks. Spikes and racemes of this type are said to be one-sided or secund. Spikelet The spikelet, the unit of the grass inflorescence, consists of bracts surrounding the floral organs and later the 'seeds.' There is a wide range of variation in structure, size, and shape, and the spikelet is of great value in identification of grasses. A typical spikelet consists of an axis, known as the RACHILLA, bearing at the base two empty bracts called GLUMES, and above them, on alternate sides of the rachilla, one or more FLORETS. A floret consists of a flower borne in the axil of two bracts. The outer (lower) of these bracts is known as the LEMMA, the inner (upper) as the PALEA. The lemma has also been termed flowering glume, lower palea (pale), or valve, and the palea has been called pale, upper palea and valvule. At the base of the floral organs (in most grasses) there are two small, inconspicuous scales known as LODICULES, which probably represent the perianth in grasses. The spikelet may have from one to many flowers. Glumes

The two glumes are borne at slightly different levels on the rachilla— hence the terms LOWER GLUME and UPPER GLUME. In size, the glumes vary greatly in relation to the rest of the spikelet. In some grasses both glumes are greatly reduced and are represented by a rim or minute scales, or the lower glume may be absent or minute while the upper is developed, as in some species of Paspalum and Digitaria. The glumes are frequently unequal in size and nervation (veining) with the lower usually shorter and narrower and with fewer nerves. In texture they vary from thin and delicate to hard, thick, and indurated. They may also vary in shape, with the presence or absence of outgrowths such as hairs, bristles, tubercles, glands, and awns. Floret

A perfectly developed floret consists of a lemma, palea, and flower. In grasses with more than one flower per spikelet, one or more florets

THE GRASS PLANT

9

are sometimes reduced to lemmas, or to a lemma and palea without a flower. T h e flower is either PERFECT (bisexual), with both stamens and a pistil; or it is male, with stamens only; or, rarely, functionally female, with a pistil and rudimentary stamens. Lemma A much wider range of variation is found in the structure of the lemma than in the glumes or paleas. In grasses with spikelets having florets all of one sex, the lemmas are usually all alike. When spikelets or florets are not bisexual, however, the lemmas enclosing bisexual flowers are usually different in form from those which have male flowers or are sterile. In a large number of grasses, the lemma is awned. An AWN is a bristlelike extension of a nerve, or a bristle formed by splitting of the apex of the lemma. The awn may project from an undivided apex, or it may arise on the back of the lemma, or from between segments known as LOBES, which occur on a lemma which is variously toothed at the apex. T h e awn may be straight or wavy, or when dry it may bend at about the middle and become twisted below the bend. In some grasses it is flattened either throughout or in the lower part only, or it may be entirely rounded. Besides the diversity of awns found in different grasses, lemmas vary, like the glumes, in size, shape, nervation, and the presence or absence of hairs and other outgrowths. In some grasses with hairy lemmas, the hairs are carried in tufts or fringes on the nerves or between the nerves. T h e lemma may be thickened at the base to form a CALLUS, which may be blunt, sharp, or needle-shaped. Palea T h e palea is more uniform in structure than the lemma. In most grasses it is slightly or much shorter than the lemma and more delicate in texture, and has a sharp keel near the margin on either side. In some grasses the palea bears variously shaped appendages. Flower T h e flower in most grasses consists of lodicules, stamens, and a pistil. There may be two, rarely three, lodicules, or occasionally there are none. Fertilization is usually facilitated by the swelling of the lodicules to force the lemma and palea apart exposing the stamens and pistil. T h e stamens are usually three in number, although rarely there are one, two, or six. T h e single pistil bears at the top of the ovary two (rarely three or one) styles and feathery or brushlike stigmas.

IO

GRASSES OF H A W A I I

Grass 'seed' T h e fruit of grasses is usually a caryopsis, in which the single seed is grown fast to the pericarp, forming a seedlike grain, but in a few genera the seed is free from the pericarp. T h e caryopsis may be free from the lemma and palea or it may be permanently enclosed. T h e embryo can be seen as an oval depression of the side of the caryopsis next to the lemma. T h e hilum is the dot or line opposite the embryo which marks the point of attachment of the seed to the ovary wall.

TAXONOMY T h e taxonomy of grasses has had a long history of development involving much controversy over placement of grasses into groupings which show natural relationships. T h e early taxonomist had the very important job of identifying and placing the newly described grasses into workable categories. T h i s early classification into tribes, genera, and species was based on similarities of gross morphological structure, type of inflorescence, number of florets per spikelet, etc. T h i s system was surprisingly accurate considering the problems which the grass taxonomist encountered and which still plague him. With the increased knowledge of the anatomy, cytology, genetics, ecology, etc., of grasses, the serious faults of the early systems of classification have become increasingly apparent. According to Bor (i960), the modern taxonomist, in determining the relationships of a species, genus, or tribe, should consider, besides gross morphological characteristics, the following topics: (1) chromosome number and size; (2) leaf anatomy, internal as well as external; (3) first seedling leaf, size and shape; (4) number, size, and shape of lodicules; (5) the nature and size of the embryo; (6) the size and shape of the hilum; (7) the nature and position of the root hairs; (8) the nature of the starch grain, whether simple or compound; (9) the physiological make-up; (10) the nucleoli; and (11) the nature of the shoot apex. T o this list should be added: (a) ecological studies of clonal materials in different environments; (b) the ability of the individuals to hybridize; and (c) the fertility of the hybrids. T h e problem of classifying grasses and determining their relationships becomes complex and involves many disciplines. Cytology, Cytogenetics, Ecology, Anatomy, Physiology, and Paleobotany must be united with T a x o n o m y in determining plant relationships. A s man continues to change the features of the landscape to suit his whims, vegetation becomes destroyed and parts of the puzzle are lost. With the shifting of vegetation and the continuing search for new germ plasm, hybrid cominations are made which defy interpretation, thus adding to the complexity of the problem. In order to illustrate the complexity of the situation, a number of the previously mentioned characteristics are elaborated further. 1.

T h e basic number and size, whether small, intermediate, or large should be considered.

CHROMOSOMES.

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GRASSES OF H A W A I I

2.

LEAF ANATOMY. Six types of leaf anatomy (in transverse section) have been described. Brown (1958) described them as follows:

a.

The festucoid type of leaf exhibits a well-developed mestome sheath of thick-walled cells, surrounded by an indistinct parenchymatous sheath of thin-walled cells containing chloroplasts similar to those of the loose irregular chlorenchyma.

b.

The bambusoid type also has a mestome sheath with thick walls, a parenchymatous sheath of circular to elliptical cells with thick walls containing typical chloroplasts and a somewhat specialized chlorenchyma. The arundinoid type has a poorly developed endodermis but is surrounded by a sheath of greatly enlarged parenchyma cells. The panicoid type may (Paniceae) or may not (Andropogoneae) have a mestome sheath and a parenchymatous sheath in which specialized starch plastids may or may not be present. The chlorenchyma is somewhat radially arranged but this arrangement is not so marked as in the chloridoid type.

c.

d.

e.

f.

The aristidoid type has no mestome sheath but has a parenchymatous sheath two cells thick, the walls of which are thicker than usual. The chlorenchyma is radially arranged. The chloridoid type has a mestome sheath around the larger bundles, and a single parenchymatous sheath. The chlorenchyma consists of long narrow radially arranged cells forming one layer.

3.

FIRST SEEDLING LEAF.

This may be erect and narrow in the Festucoid group and broad and spreading in the Panicoid group.

4.

THE EMBRYO.

The detailed structure of the embryo has proven to be of great importance. Reeder (1957) considered that four embryonic characters had taxonomic significance: (a) the course of the vascular system (whether the traces to the scutellum and embryonic leaves diverge at about the same point, or are separated by a more or less elongated internode), (b) epiblast (whether present or absent), (c) lower part of the scutellum (whether free from the coleorhiza or fused to it), and (d) the cross-section of the embryonic leaf (whether having many bundles and over-

TAXONOMY

lapping margins, or few bundles and margins which merely meet). From these characters he recognized the following types: a. Festucoid type which has a festucoid vascular supply, presence of an epiblast, no cleft between coleorhiza and scutellum, and a primary leaf in which there are few bundles and with margins that do not overlap. b. Panicoid type which has a panicoid vascular supply, no epiblast, a definite cleft between scutellum and coleorhiza, a primary leaf which overlaps at the margins and has many vascular bundles. c. Chloridoid-eragrostoid type which has a panicoid vascular supply, a cleft between the scutellum and coleorhiza, an epiblast and primary leaves of which the margins do not overlap. d. Bambusoid type which has panicoid characters except for the presence of an epiblast. e. Oryzoid-olyroid type which has a festucoid type of vascular supply and a panicoid primary leaf. An epiblast and/or a cleft between the scutellum and the coleorhiza may or may not be present. f. Arundinoid-danthonioid type which has mainly panicoid characters and differs in the embryonic leaf which has few vascular bundles and margins which only meet and do not overlap. It was soon realized during the preparation of this text that any short, simple classification scheme would be impossible to adapt for use here because of the tremendous diversity of grasses introduced into Hawaii. The scheme used by Hitchcock and Chase in the Manual of Grasses of the United States, 2nd ed. (1950), could not be used without considerable difficulty. They recognized only 13 tribes of grasses and admitted that part of the scheme was rather awkward and arbitrary. Hubbard, in Hutchinson (1959), recognized 27 tribes which show more nearly natural relationships among the members of each group. Bor's (i960) scheme is an expansion of Hubbard's and he recognized 39 tribes excluding the bamboos. The tribes of grasses as recognized by Hitchcock and Chase, Hubbard (Hutchinson), and Bor are listed in Table 1. An alphabetical listing of the Hawaiian grass genera with their tribal placement according to Hitchcock and Chase, Hubbard, and Bor are presented in Table 2.

J

4

GRASSES OF

TABLE

HAWAII

I

T H E G R A S S T R I B E S A C C O R D I N G TO B O R , H U B B A R D , AND HITCHCOCK AND CHASE BOR*

HUBBARD ( H U T C H I N S O N , 1 9 5 9 )

Group I (Panicoideae) Andropogoneae Maydeae Paniceae Group II (Pooideae) Aeluropodeae Agrostideae Aristideae Arundineae Arundinelleae Aveneae Brachypodieae Bromeae Centotheceae Chlorideae Danthonieae Ehrharteae Eragrosteae Festuceae Garnotieae Glycerieae Hubbardieae Isachneae Leptureae Lygeeae Meliceae Milieae Monermeae Oryzeae Pappophoreae Perotideae Phaenospermeae Phalarideae Phareae Pommereulleae Sporoboleae Stipeae Streptogyneae Thysanolaeneae Triticeae Zoysieae * Does not list Bambuseae

Group I (Panicoideae) Andropogoneae Maydeae Paniceae Group II (Pooideae) Agrosteae Anomochloeae Arundineae Arundinelleae Aveneae Bambuseae Chlorideae Eragrosteae Festuceae Hordeeae Leptureae Lygeae Nardeae Olyreae Oryzeae Pappophoreae Parianeae Phalarideae Phareae Sporoboleae Stipeae Streptochaeteae Thysanolaeneae Zoysieae

HITCHCOCK A N D CHASE

Group I (Panicoideae) Andropogoneae Melinideae Paniceae Tripsaceae Group II (Festucoideae) Agrostideae Aveneae Bambuseae Chlorideae Festuceae Hordeae Oryzeae Phalarideae Zizanieae Zoysieae

TAXONOMY

15

TABLE 2 COMPARISON OF TRIBAL ASSIGNMENTS OF HAWAII GRASS GENERA ACCORDING TO HITCHCOCK AND CHASE, HUBBARD, AND BOR TRIBAL PLACEMENT GENERA

HITCHCOCK AND CHASE

HUBBARD

BOR

Agropogon Agropyron Agrostis Aira Alloteropsis Alopecurus Ammophila Andropogon Anthoxanthum Apera Aristida Arrhenatherum Arundinella Arundo Astrebla Avena Axonopus

Agrostideae Hordeae Agrostideae Aveneae Paniceae Agrostideae Agrostideae Andropogoneae Phalarideae Agrostideae Agrostideae Aveneae Festuceae Festuceae Chlorideae Aveneae Paniceae

Agrosteae Hordeae Agrosteae Aveneae Paniceae Agrosteae Agrosteae Andropogoneae Phalarideae Agrosteae Stipeae Aveneae Arundinelleae Arundineae Aveneae Aveneae Paniceae

Agrostideae Triticeae Agrostideae Aveneae Paniceae Agrostideae Agrostideae Andropogoneae Aveneae Agrostideae Aristideae Aveneae Arundinelleae Arundineae Aveneae Aveneae Paniceae

Bechmanttia Bothriochloa Bouteloua Brachiaria Brachypodium Briza Bromus Buchloé

Chlorideae Andropogoneae Chlorideae Paniceae Festuceae Festuceae Festuceae Chlorideae

Festuceae Andropogoneae Chlorideae Paniceae Festuceae Festuceae Festuceae Chlorideae

Festuceae Andropogoneae Chlorideae Paniceae Brachypodieae Festuceae Bromeae Chlorideae

Calamagrostis Cenchrus Chloris Chrysopogon Coix Cortaderia Cymbopogon Cynodoti Cynosurus

Agrostideae Paniceae Chlorideae Andropogoneae Tripsaceae Festuceae Andropogoneae Chlorideae Festuceae

Agrosteae Paniceae Chlorideae Andropogoneae Maydeae Arundineae Andropogoneae Chlorideae Festuceae

Agrostideae Paniceae Chlorideae Andropogoneae Maydeae Arundineae Andropogoneae Chlorideae Festuceae

Dactylis Dactyloctenium Danthonia

Festuceae Chlorideae Aveneae

Festuceae Eragrosteae Aveneae

Festuceae Eragrosteae Danthonieae

i6

GRASSES OF H A W A I I

TABLE 2

(continued) TRIBAL PLACEMENT

GENERA

HITCHCOCK AND CHASE

HUBBARD

BOR

Deschampsia Dichanthium Dichelachne Digitarla Dissochondrus

Aveneae Andropogoneae Agrostideae Paniceae Paniceae

Aveneae Andropogoneae Agrosteae Paniceae Paniceae

Aveneae Andropogoneae Agrostideae Paniceae Paniceae

Echinochloa Ehrharta Eleusine Elymus Enneapogon Enteropogon Eragrostis Eremochloa Erianthus Eriochloa Euchlaena Eulalia

Paniceae Phalarideae Chlorideae Hordeae Festuceae Chlorideae Festuceae Andropogoneae Andropogoneae Paniceae Tripsaceae Andropogoneae

Paniceae Phalarideae Eragrosteae Hordeae Pappophoreae Chlorideae Eragrosteae Andropogoneae Andropogoneae Paniceae Maydeae Andropogoneae

Paniceae Ehrharteae Eragrosteae Triticeae Pappophoreae Chlorideae Eragrosteae Andropogoneae Andropogoneae Paniceae Maydeae Andropogoneae

Festuca

Festuceae

Festuceae

Festuceae

Garnotia Gastridium Glyceria

Agrostideae Agrostideae Festuceae

Agrosteae Agrosteae Festuceae

Garnotieae Agrostideae Glycerieae

Hackelochloa Heteropogon Hilaria Holcus Hordeum Hyparrhenia

Andropogoneae Andropogoneae Zoysieae Aveneae Hordeae Andropogoneae

Andropogoneae Andropogoneae Zoysieae Aveneae Hordeae Andropogoneae

Andropogoneae Andropogoneae Zoysieae Aveneae Triticeae Andropogoneae

Imperata Isachne Ischaemum Ixophorus

Andropogoneae Paniceae Andropogoneae Paniceae

Andropogoneae Paniceae Andropogoneae Paniceae

Andropogoneae Isachneae Andropogoneae Paniceae

Lamarckia Leptochloa Lepturus Lolium Lycurus

Festuceae Chlorideae Hordeae Hordeae Agrostideae

Festuceae Eragrosteae Leptureae Festuceae Agrosteae

Festuceae Eragrosteae Leptureae Festuceae Agrostideae

Melica Melinis

Festuceae Melinideae

Festuceae Paniceae

Meliceae Paniceae

TAXONOMY

TAB L E 2

(continued) TRIBAL PLACEMENT

GENERA

HITCHCOCK AND CHASE

HUBBARD

BOR

Microlaena Muhlenbergia

Phalarideae Agrostideae

Ehrharteae Agrosteae

Phalarideae Agrostideae

Narenga

Andropogoneae

Andropogoneae

Andropogoneae

Oplismenus Oryza Oryzopsis

Paniceae Oryzeae Agrostideae

Paniceae Oryzeae Stipeae

Paniceae Oryzeae Stipeae

Panicum Paspalidium Paspalum Pennisetum Phalaris Phleum Poa Polypogon Polytoca Polytrias

Paniceae Paniceae Paniceae Paniceae Phalarideae Agrostideae Festuceae Agrostideae Tripsaceae Andropogoneae

Paniceae Paniceae Paniceae Paniceae Phalarideae Agrosteae Festuceae Agrosteae Maydeae Andropogoneae

Paniceae Paniceae Paniceae Paniceae Phalarideae Agrostideae Festuceae Agrostideae Maydeae Andropogoneae

Rhynchelytrum

Paniceae

Paniceae

Paniceae

Saccharum Sacciolepis Schedonnardus Schizachyrium Sclerostachya Secale Sehima Setaria Sorghastrum Sorghum Spartirla Sphenopholis Spodiopogon Sporobolus Stenotaphrum Stipa

Andropogoneae Paniceae Chlorideae Andropogoneae Andropogoneae Hordeae Andropogoneae Paniceae Andropogoneae Andropogoneae Chlorideae Aveneae Andropogoneae Agrostideae Paniceae Agrostideae

Andropogoneae Paniceae Chlorideae Andropogoneae Andropogoneae Hordeae Andropogoneae Paniceae Andropogoneae Andropogoneae Chlorideae Aveneae Andropogoneae Sporoboleae Paniceae Stipeae

Andropogoneae Paniceae Chlorideae Andropogoneae Andropogoneae Triticeae Andropogoneae Paniceae Andropogoneae Andropogoneae Chlorideae Aveneae Andropogoneae Sporoboleae Paniceae Stipeae

Themeda Tragus Trichachne Trichoneura

Andropogoneae Zoysieae Paniceae Chlorideae

Andropogoneae Zoysieae Paniceae Eragrosteae

Andropogoneae Zoysieae Paniceae Eragrosteae

18

GRASSES OF

TABLE

2

HAWAII

(continued) TRIBAL PLACEMENT

GENERA

HITCHCOCK AND CHASE

HUBBARD

BOR

Tridens Tripsacutn Triraphis Trisetum Triticum

Festuceae Tripsaceae Eragrosteae Aveneae Hordeae

Eragrosteae Maydeae Eragrosteae Aveneae Hordeae

Eragrosteae Maydeae Eragrosteae Aveneae Triticeae

Urochloa

Paniceae

Paniceae

Paniceae

Vulpia

Festuceae

Festuceae

Festuceae

Zea Zizania Zoysia

Tripsaceae Zizanieae Zoysieae

Maydeae Oryzeae Zoysieae

Maydeae Oryzeae Zoysieae

Bor's arrangement of the systematic work on Indian grasses is strictly practical, using two subfamilies, Panicoideae and Pooideae, with the tribes in each subfamily, genera in each tribe, and species in each genus following in alphabetical order. There is no suggestion of a phylogenetic arrangement in the position of any tribe or lessor taxon. He arranged in groups (tribes) genera which seem to be related because of their morphological characteristics, foliar anatomy, chromosome numbers, etc., as far as they were known. T h e Hawaiian grasses included in this listing, however, are placed into six subfamilies as outlined by Stebbins and Crampton (1959) in order to show a more natural grouping of the tribes: Subfamily Subfamily Tribe Tribe Subfamily Tribe Tribe Tribe Tribe Tribe

i. 2.

i. 2.

33456. 7-

Bambusoideae Oryzoideae Ehrharteae Oryzeae Arundinoideae Aristideae Arundineae Arundinelleae Danthonieae Garnotieae

Subfamily Tribe Tribe Tribe Tribe Tribe Tribe Tribe Tribe Tribe

48. 9-

10. 11. 12.

14.

i516.

Festucoideae Agrostideae Aveneae Brachypodieae Bromeae Festuceae Glycerieae Isachneae Leptureae Meliceae

TAXONOMY

Tribe Tribe Tribe Subfamily Tribe Tribe Tribe

17. 18. 19. 5. 20. 21. 22.

Phalarideae Stipeae Triticeae Eragrostoideae Chlorideae Eragrosteae Pappophoreae

Tribe Tribe Subfamily Tribe Tribe Tribe

19 23. 24. 6. 25. 26. 27.

Sporoboleae Zoysieae Panicoideae Andropogoneae Maydeae Paniceae

T h e citations for publication of the species names have been checked against originals whenever possible and with the listing presented by Chase and Niles (i960). In a number of instances there is a great deal of conflict as to nomenclature. N o complete listing of synonymy has been attempted, but, insofar as possible, all references in the literature pertaining to Hawaiian plants have been included. T h e r e are several large gaps in our knowledge of our native and naturalized grasses. A number of genera, in particular our native Panicums and Deschampsias, need careful study before classification to species can be accomplished. It would be of great help to grow all of them clonaly in a number of different environments to determine if we have true species differences or merely phenotypic modifications by environmental interaction. Spatial isolation on different islands does not necessarily mean that the different morphological forms are species differences. K e y s to the tribes (adapted from N . L . Bor, i960) are presented immediately after the taxonomic description of the family Gramineae. Each tribal description is followed by a key to its genera (if more than one are found in Hawaii). Genus descriptions are followed by a listing of the species and their synonyms. N o keys or descriptions are given for the species within each genus. In the following pages the family is divided into five subfamilies (excluding Bambusoideae) : Oryzoideae, Arundinoideae, Festucoideae, Eragrostoideae, and Panicoideae. Within each subfamily, the tribes, genera, and species are arranged in alphabetical succession.

PAGE INDEX TO TRIBES AND GENERA

Tribes

Tribes

Agrostideae

Garnotieae

49

key

51 51

Glycerieae

111

.. key

188 188

Isachneae

114

Aristideae

38

Leptureae

116

Arundineae

41

Maydeae

223

Andropogoneae

..

key

41

Arundinelleae

44

Aveneae

73 key

Brachypodieae

90

Bromeae Chlorideae

73

key Meliceae Oryzeae

23i

Phalarideae

119

Sporoboleae

178

key key

3o 30 151 151

Triticeae

key

95 95

Zoysieae

key

Festuceae

231

175

46 46

Eragrosteae

35 35

Pappophoreae

134 134

Ehrharteae

key

key

key

Danthonieae

118

Paniceae

91

223

Stipeae

key

122 122

key

124 124

key

182 182

P A G E I N D E X TO T R I B E S A N D G E N E R A

Genera

Agropogon Agropyron Agrostis Aira Alloteropsis Alopecurus Ammophila Andropogon Anthoxanthum Apera Aristida Arrhenatherum Arundinella Arundo Astrebla Avena Axonopus Bechmannia Bothriochloa Bouteloua Brachiaria Brachypodium Briza Bromus Buchloe Calamagrostis Cenchrus Chloris Chrysopogon Coix Cortaderia Cymbopogon Cyrtodon Cynosurus Dactylis

21

Genera

..

..

•

••

53

•

••

125

•

••

53

•

• •

74

• .

•• ..

.

..

235 60 60 J

•

••

• .

•• ..

75 60

.

..

38

•

••

77

• .

•• ..

44 41

.

..

46

•

••

77

•

••

235

.

..

96

•

••

195

Echinochloa Ehrharta Eleusine Elymus Enteropogon Enneapogon Eragrostis Eremochloa Erianthus Eriochloa Euchlaena Eulalia

• .

•• ..

135 236

Festuca

•

••

9°

• .

•• ..

97 91

•

••

137

.

..

62

..

238

..

138

• • .

93

Dactyloctenium Danthonia Deschampsia Dichanthium Dichelachne Digitaria Dissochondrus

197

..

42

••

I

..

142

•• ••

99 97 99

Garnotia Gastridium Glyceria Hackelochloa Heteropogon Hilaria Holcus Hordeum Hyparrhenia Imperata Isachne Ischaemum Ixophorus

• •

152

•• ..

47 80

..

201

..

62

..

246

••

253

• •

255

••

3°

••

153 146

••

175

•• ..

155 202

•• ..

203 258

• •

225

• •

203

••

99

•• ..

49 65 n i

• • ..

203 204

..

182

..

84

..

126

..

206

..

208

..

114

..

208

..

260

GRASSES OF HAWAII

22

Genera

Genera

Lamarckia Leptochloa Lepturus Lolium Lycurus

..

Melica Melinis Microlaena Muhlenbergia

..

IOI

..

169

..

..

116

..

..

103

..

..

66

..

..

118

. . . .

31

. . . .

66

Narenga Oplismenus Oryza Oryzopsis Panicum Paspalidium Paspalum Pennisetum Phalaris Phleum Poa Polypogon Polytoca Polytrias Rhynchelytrum Saccharum Sacciolepis Schedonnardus

..

..

262

•• ..

•• ..

35 122

..

..

263

..

..

285

..

..

285

..

..

293

..

..

119

. . . . ..

..

. . . . ..

..

..

68 105

Sporobolus Stenotaphrum Stipa

•

• .

•• ..

213 214

..

129

..

214

•• ..

3°2 215

..

216

.

..

149

,

..

87

..

220

..

178

••

3°6

••

123

..

220

..

183 308

Themeda Tragus Trichachne Trichoneura Tridens Tripsacum Triraphis Trisetum Triticum

..

87

..

131

Urochloa

••

311

Vulpia

..

109

Zea Zizania Zoysia

..

228

169 ..

171

•• ..

225 174

68

..

225

..

212

..

295

..

212

..

..

300

..

..

149

..

Schizachyrium .. Schlerostachya Secale Sehima Setaria Sorghastrum Sorghum Spartina Sphenopkolis Spodiopogon

..

36

..

186

GRAMINEAE GRAMINEAE A. L. de Jussieu, Gen. PI. 28 (1789) Annual or perennial herbs, rarely shrubs or trees with woody stems often reaching a considerable height (some Bambuseae); stems jointed, erect or prostrate, and creeping at the base and then usually rooting at the nodes, less often erect upon stiltroots (the latter usually developed in forest or shade-loving species and those living in swamps), cylindrical, rarely flattened or quadrangular, hollow in the internodes, solid at the nodes, sometimes solid all through; underground root system often well developed in perennials, extending sometimes to considerable distances by means of rhizomes; stolons, which are sometimes tough and woody, are developed by some species and spread to considerable distances. In branched culms, at the point where the branching occurs, with its back to the main stem, is developed a 2-keeled organ called the prophyllum. Leaves arranged alternately in two ranks on the stems and consisting of sheath, ligule, and blade; sheaths surrounding the culm with overlapping free or sometimes connate margins, often swollen at the base, ribbed or smooth; ligule situated at the junction of sheath and blade, membranous or chartaceous, a fringe of bristles or hairs, rarely absent; blades usually long and narrow, rarely broad, ovate, sometimes filiform or setaceous, joined to or jointed upon the sheath with a narrowed, rounded, or clasping base, rarely petioled. Inflorescence various, rarely consisting of 1 spikelet, more often in spikes, few- or many-spiculate panicles, racemes or false racemes in which a sessile spikelet is accompanied by a pedicelled spikelet or its rudiment. Flowers mostly bisexual, sometimes unisexual, usually consisting of stamens and pistil and 2 or 3 minute, hyaline scales (lodicules) representing the perianth or the latter absent, enclosed and sessile between 2 bracts (lemma and palea), the whole called a floret. Florets, if more than one, alternate in one plane on the opposite sides of a jointed or tough axis (rachilla), rarely subspirally arranged with 2 empty bracts (glumes) at the base of the spikelet, rarely one or both of the glumes absent. Stamens hypogynous, 1-6 (very numerous in Pariana), consisting of long delicate filaments and 2-locular anthers, opening usually by longitudinal slits, occasionally by pores; ovary 1-celled, with 1 anatropous, hemitropous, or campylotropous ovule usually attached to a point or line on the adaxial side of the carpel, the point of attachment being visible on the grain as the hilum; styles

24

G R A S S E S OF H A W A I I

usually 2, rarely i or 3; stigmas mostly plumose; fruit 1-seeded, indehiscent, rarely with a mucilage-forming pericarp. KEY TO THE TRIBES OF GRAMINEAE

A.

Spikelets 2-flowered, falling entire at maturity, usually with the upper floret bisexual and the lower male or sterile and if the latter, often reduced to the lemma or rarely the lemma entirely absent, all alike or more often differing in size, shape and structure, frequently dorsally compressed B

A.

Spikelets 1- to many-flowered, breaking up at maturity above the more or less persistent glumes, or if falling entire, then not 2flowered with the lower floret male or sterile and the upper bisexual, usually more or less laterally compressed or terete, or if 2-flowered and falling entire, the glumes and lemmas all membranous and with a globose wrinkled seed which splits the palea when mature . C

B i . Male and female spikelets in separate inflorescences or in different parts of the same inflorescence and of different appearance; lemmas hyaline or membranous and thinner than the glumes Maydeae p. 223 B i . Spikelets all bisexual, or with male or sterile and bisexual spikelets mixed in the same inflorescence and so arranged that a male or sterile spikelet is near a bisexual spikelet, or if unisexual then the lemma of the fertile floret indurated 2.

Spikelets often paired, with one sessile and the other pedicelled, those of each pair similar or more often dissimilar, rarely solitary and all alike; glumes as long as the spikelet and enclosing the florets, more or less rigid and firmer than the lemmas which are both hyaline or membranous; upper lemma usually awned Andropogoneae p. 188

2.

Spikelets solitary or paired, more or less similar; glumes usually membranous, the lower usually smaller or sometimes suppressed; lower lemma mostly resembling the upper glume in texture; upper lemma papery to very tough and rigid, usually awnless Paniceae p. 231

GRAMINEAE - KEY TO TRIBES

25

C i . Treelike or shrubby bamboos; culms woody, persistent; leaf blades usually with a petiole-like base, often articulated with the sheath; lemmas 5- to many-nerved . . . . Bambuseae C i . Perennial or annual herbs, including reeds and reedlike plants, not treelike, rarely shrubby; culms usually neither woody nor persistent; leaf blades mostly sessile and not articulated with the sheaths; lemmas 1- to many-nerved 2.

Spikelets borne in open or contracted or spikelike panicles, less often in racemes or spikes 3.

Spikelets usually with 2 or more fertile florets, or if with 1 fertile floret then with sterile reduced florets above it* 4.

Lemma and rachilla glabrous or hairy, but with the hairs not enveloping the lemma, or if so, then the lemma with a geniculate awn; low or moderately tall grasses 5.

Glumes usually shorter then the lowest floret and with the upper floret distinctly exserted, rarely longer and then with firm dull margins like the lemmas, or if with hyaline margins then the lemma firmly keeled; lemmas awnless or with a straight or curved awn from the entire or bifid apex, or several-awned or -lobed 6.

Lemmas usually 5- to many-nerved, awnless or awned; awn usually simple, not kneed or twisted below the knee 7.

Lemmas if awned, with 1 awn (if more-awned, tip of ovary hairy), if awnless, lemmas not 7-9-nerved or strongly compressed 8.

8. * See Chlorideae

Ovary with a hairy appendage at the apex; styles borne laterally below the appendage 9.

Inflorescence of racemes or in panicles; awn inserted below the entire or 2-lobed tip Bromeae p. 91

9.

Inflorescence a raceme; awn terminal Brachypodieae p. 90 Ovary glabrous at the apex, or if hairy at least without a hairy appendage

26

GRASSES OF

HAWAII

10. Lemmas dull, membranous to coriaceous, exserted from the glumes; glumes 2 or i (upper glume only present in Lolium) . Festuceae P- 95 10. Lemmas membranous and shining, the upper represented by a bundle of empty, rudimentary lemmas; glumes 2 . . . . Meliceae p. 118 7.

6.

Lemmas 9-awned or if awnless, lemmas 7-9-nerved, strongly compressed or rounded; ovary not hairy 11. Lemmas 9-nerved; the nerves produced as 9 awns Pappophoreae P- 175 11. Lemmas 7-9-nerved; nerves not produced as awns; spikelets several-flowered with conspicuously nerved lemmas; ligules membranous Glycerieae p. h i Lemmas 1-3-nerved; inflorescence of panicles or racemes

12. Inflorescence of panicles; if racemes or spikes the spikelets not secund Eragrosteae P- J 5i 12. Inflorescence of racemes or panicles of racemes, the spikelets secund . . . . Chlorideae P- 134 5. Glumes usually as long as or longer than the lowest floret, often as long as the spikelet and enclosing the florets, if shorter than the lowest floret, then the palea with 2 awns 13. Lemmas awnless or awned from the back, rarely from the tip; awn usually kneed; ligules membranous Aveneae P- 73 13. Lemmas awned from the sinus between the two terminal lobes or no trace of an awn, often only a mucro; ligule a hairy rim . . . . Danthonieae p. 46

GRAMINEAE - KEY TO TRIBES

4.

3.

27

Lemmas or rachilla joints bearing long silky hairs which envelop the lemma (at least in fertile florets); lemmas awnless or with a straight awn from the tip, often thin; tall grasses with usually large plumelike panicles Arundineae p. 41 Spikelets with 1 fertile floret (male or female in unisexual 1-flowered spikelets), with or without 1 or 2 male or sterile florets below it, the latter often much reduced

14. Glumes minute (usually represented by 2 semicircular inconspicuous lips) or suppressed; fertile lemma and palea very similar in shape and texture; spikelets firmly compressed; stamens usually 6; florets bisexual or unisexual Oryzeae P- 35 14. Glume well developed; fertile lemma and palea not similar 15. Spikelets with 3 florets, the lower two represented by empty lemmas which are often minute and occasionally one of them absent 16. Lower florets represented by 2 more or less welldeveloped lemmas, one or both of which are transversely ridged or flanged, if smooth or hairy, not golden brown or black in color . . . . Ehrharteae P- 3° 16. Lower florets well developed or minute and sometimes one absent, never transversely rugose or flanged, sometimes silky, golden brown to almost black Phalarideae p. 119 15. Spikelets with 1 or 2 florets 17. Spikelets with 2 florets, the lower male or sterile, the upper bisexual; spikelets falling with part of the pedicel attached or leaving the glumes attached to the pedicel; grains elliptic to oblong or terete, not globose

28

GRASSES OF H A W A I I

18. Glumes equal in size, obtuse or mucronate; upper lemma unawned . . . . Isachneae p. 114 18. Glumes unequal, the lower acute, the upper acuminate; upper lemma awned with a perfect awn or at least a bristle Arundinelleae p. 44 17. Spikelets with 1 fertile floret 19. Spikelets cylindrical, falling entire; base of the glumes bearded; lemma awned (see also Agrostis semiverticillata, Polypogon spp. of the Agrostideae) Garnotieae P- 49 19. Spikelets cylindrical, ellipsoid-acute or definitely laterally compressed, not bearded at the base; rachilla disarticulating above the more or less persistent glumes 20. Spikelets very rarely falling entire and then with firmly membranous to coriaceous, awned, or 5-nerved lemmas 21. Lemmas hyaline or membranous at maturity, rarely indurated and then laterally compressed, awnless or awned from low down on the back, or from the middle, or from the entire or bifid tip 22. Lemmas usually 3-5-nerved, frequently awned; glumes longer and firmer than the hyaline lemma or when the lemma is longer than the latter, indurated; grains usually with an adhering pericarp . Agrostideae P- 5i 22. Lemmas i-3-nerved, awnless; glumes and lemmas very similar in texture, hyaline or thinly membranous, shining; grain usually with a free pericarp . Sporoboleae p. 178

G R A M I N E A E - KEY TO TRIBES

29

21. Lemmas indurated or rigid at maturity, terete or dorsally compressed 23. Awns trifid .

Aristideae P- 38 23. Awns simple, sometimes flanked by 2 bristles Stipeae p. 122 .

.

.

20. Spikelets falling entire at maturity, either singly or in clusters from the axis of slender spikelike panicles or racemes; lemma delicate, 1-3-nerved; spikelets fascicled or solitary; glumes not awned Zoysieae p. 182 2.

Spikelets sessile or shortly pedicelled along one side of the rachis of solitary, digitate, or scattered spikes, spikelike racemes (with 1 fertile floret and 1-3-nerved lemmas), or on opposite sides of the rachis of solitary spikes or racemes (care must be taken that certain genera of the Andropogoneae in which the pedicelled spikelet is suppressed or reduced are not placed in this section) 24. Spikelets on opposite sides of the rachis of solitary spikes or spikelike racemes 25. Ovary hairy, especially at the apex; grain often longitudinally grooved, with the linear hilum as long as the grain; starch grains simple; lodicules hairy; lemmas 3-5- or more nerved Triticeae p. 124 25. Ovary glabrous; grain not grooved; hilum small, basal or subbasal; starch grains compound or simple; lodicules glabrous; lemmas 1-3-nerved; lemma of the first floret with its back adjacent to the rachis; lower glume completely absent Leptureae p. 116 24. Spikelets secund on a tough rachis; 1-2 florets bisexual with sterile florets above; inflorescence of digitatt or solitary spikes or panicles or spikes Chlorideae P- *34

GRASSES OF

3°

HAWAII

EHRHARTEAE Nevski, Act. Inst. Bot. Acad. Sci. U.R.S.S., ser. i , fasc. 4:227 (1937). Spikelets 1-3-flowered with the two lower florets reduced to the lemmas, the uppermost bisexual. Glumes equal or unequal, usually shorter than the spikelet; rachilla disarticulating above the glumes, not between the florets. Sterile lemmas awned or awnless, varying in size, equal or unequal, one or both ridged, wrinkled, or flanged, the upper of the two often with a basal appendage; fertile lemma slightly to much shorter; palea 2-nerved, 2-keeled; keels close together. Stamens 6, 4, 3, 2, or 1. Styles 2; stigmas plumose. Grain elliptic, compressed; hilum linear, almost the length of the grain; embryo small; starch grains compound. Annual or perennial grasses. Leaf blades linear to lanceolate, flat or rolled, sometimes much reduced, with festucoid anatomy; 2-celled microhairs present, elongated; silica bodies cross-shaped, nodular or dumbbell-shaped. Ligules membranous. Inflorescence a panicle or raceme. Chromosomes small; basic number 6. Genera:

Ehrharta Microlaena KEY TO THE GENERA OF EHRHARTEAE

1.

Palea linear in outline or boat-shaped almost as long as the fertile lemma, keeled, finely 2-nerved. Stamens 6 or 3, rarely 1 Ehrharta P- 30

2.

Palea narrow-lanceolate, shorter than the fertile lemma, 1-nerved. Stamens 4 or 2 Microlaena P- 3i

Ehrharta Thunb., Vet. Acad. Handl. Stockh. 216 (1779) nom. cons. SPIKELETS solitary or sometimes clustered, pedicelled, awned or awnless, not falling entire, FLOWER I . GLUMES persistent, sometimes widely diverging, unequal or subequal, lanceolate or ovate-lanceolate in profile; obtuse or acute, sometimes shortly mucronate, rarely ciliate, membranous, pubescent or glabrous, LEMMAS 3 ; the two lower lemmas equal or unequal, ovate-lanceolate or linear-oblong in profile, obtuse or produced into a long acumen, sometimes 2-lobed with a short awn from the sinus, transversely rugose, tuberculate or smooth, pilose or villous,

ORYZOIDEAE -

EHRHARTEAE

31

scabrid or glabrous, stiffly membranous or coriaceous, empty; the lower shorter or as long as the glumes, sometimes membranous; the upper sometimes shortly mucronate, with 2 unequally lobed appendages at the base; awn, when present, longer or shorter than the body of the lemma, usually scabrid. FERTILE LEMMA elliptic, oblong-linear or ovate in profile, obtuse, more rarely truncate, sometimes tuberculed and scabrid, membranous to coriaceous or almost hyaline, sometimes with a knoblike appendage at the base forming a hinge with the appendage of the upper empty lemma, usually glabrous, more rarely long-villous, subtending a palea and a bisexual flower, PALEA linear in profile or boat-shaped, almost as long as its lemma, keeled, finely 2-nerved. LODICULES 2. STAMENS 6 or 3, very rarely i. OVARY obovoid; styles free, plumose or brushlike above, exserted above the base, GRAIN elliptic in outline; embryo about 1/5 the length of the grain. Annuals or perennials; culm striate or ribbed, glabrous; leaf sheath longer or shorter than the internode of the culm, pubescent or glabrous; ligule scarious or membranous, or reduced to a mere rim, sometimes ciliate; blade sometimes much reduced, stiff or flaccid; flat and linear or linear-lanceolate, or folded or setaceous, or filiform, with a distinct midrib and distinct lateral veins, smooth or scabrid, pubescent or glabrous; inflorescence an open or contracted panicle or raceme; primary branches of the inflorescence often panicled at the nodes. 1.

Ehrharta calycina J.E. Sm., PI. Icon. Ined., pi. 33 (1789). [Fig. 2]

2.

E. erecta Lamk., Encycl. Meth. Bot. 2:347 (1786).

Microlaena R. Br., Prodr. Fl. Nov. Holl. 210 (1810). SPIKELETS solitary, pedicelled, laterally compressed, awned, not falling entire, the rachilla disarticulating above the glumes, not, or obscurely produced, FLOWER I . GLUMES persistent, unequal, membranous; the lower very small, ovate, entire, 1-nerved; the upper almost twice as long, otherwise similar, LEMMAS 3, heteromorphous, the two lower empty, the uppermost subtending a bisexual floret, STERILE LEMMAS much longer than the glumes, cartilaginous, laterally compressed, scabrid along the keel and tapering into a long terminal slender scabrid awn, the lower strongly 5-nerved or -ribbed, the upper 7-9-nerved and grooved; callus rather acute, bearded with white silky hairs, FERTILE LEMMA much shorter than the sterile lemmas, strongly laterally compressed, cartilaginous, acutely keeled, tapering into an acute point or short

32

and Pastures of South Africa.)

GRASSES OF H A W A I I

ORYZOIDEAE -

EHRHARTEAE

33

scabrid awn and more or less strigose-ciliate along the keel in the upper half, entire, 5-nerved. PALEA narrow-lanceolate, thinly membranous, shorter than the fertile lemma, 1-nerved, LODICUI.ES 2 , ovate, hyaline, acute, STAMENS 4 or 2 . STYLES distinct, GRAIN oblong-linear in outline, compressed, enclosed within the flowering glume and palea but free from them. Wiry perennials of loose or tufted habit. Leaf blade flat or convolute; ligule reduced to a short hyaline rim fringed with a few long silky hairs which fall early. Inflorescence a simple narrow or subracemose panicle or simple raceme. 1.

Microlaena stipoides (Labill.)

R. Br., Prodr. Fl. Nov. Ho 11.

2 1 0 (1810). [Fig. 3]

Ehrharta stipoides (Labill.), Fl. Nov. Holl. PI. 1 : 9 1

(1804).

GRASSES OF H A W A I I

3. Microlaena

stipoides R . Br. (From A . S. Hitchcock, T h e Grasses of Hawaii.)

ORYZOIDEAE -

ORYZEAE

35

ORYZEAE Dumort., Obs. Gram. Belg. 83, 135 (1823). Spikelets all alike, bisexual, 1- rarely 3-flowered, with the terminal floret bisexual and the two lower reduced to scalelike lemmas; rachilla disarticulating above the glumes. Glumes very minute or suppressed, sometimes represented by two flattened minute lips; lemmas of two kinds; the lower scalelike and empty or entirely suppressed; the upper, in the bisexual spikelets, membranous or coriaceous, 3-9-nerved, awnless or awned from the tip, similar in texture and shape to the palea. Lodicules 2, 2-lobed or entire. Stamens 6 or 3 or 1 ; filaments long. Ovary with 2 styles and 2 laterally exserted stigmas. Grain with a welldeveloped linear hilum; starch grains compound. Annual or perennial herbs, usually aquatic; leaf blades narrow or broad; chlorenchyma evenly distributed between the vascular strands; silica bodies dumbbell-shaped, placed transversely; microhairs elongate. The first foliage leaf of the seedling is flat, broad, and horizontal. Ligule membranous, eventually splitting and becoming lacerate, rarely absent. Inflorescence of pedicelled spikelets in open or contracted panicles. Chromosomes small; basic number 12. Genera: Oryza Zizania KEY TO THE GENERA OF ORYZEAE

i.

Spikelets unisexual; upper branches bearing female spikelets; the lower bearing male spikelets; tall aquatic annuals or perennials, reedlike Zizania P- 36 1. Spikelets bisexual Oryza P- 35 Oryza L., Sp. PI. 333 (1753). SPIKELETS solitary, pedicelled, falling entire; pedicels linear, somewhat cup-shaped at the apex, FLOWER I. GLUMES equal, scale- or bristle-like, acuminate, sometimes suppressed, LEMMA boat-shaped, obtuse or acute, awned or awnless, strongly 3-nerved, rigidly ciliate, scabrid, rugose, chartaceous to almost crustaceous, subtending a palea and a bisexual flower; awn, when present, 1 \-2.\ times as long as the body of the lemma, scabrid. PALEA as long as the lemma and similar in texture.

36

GRASSES OF H A W A I I

LODICULES 2-lobed, rarely entire, STAMENS 6. STYLES short, free, plumose above, laterally exserted. GRAIN narrow, free or adnate to the lemma and palea. Annuals (?); culm striate, glabrous; leaf sheath prominently ribbed, glabrous; ligule long, scarious; blade flat, long-linear, closely ribbed, glabrous; inflorescence a narrow panicle; primary branches of the inflorescence erect.

i.

Oryza sativa L., Sp. PL 333 (1753). [Fig. 4]

Zizania L., Sp. PL 991 (1753). unisexual, i-flowered, disarticulating from the pedicel. obsolete, represented by a small collar-like ridge, PISTILLATE SPIKELET terete, angled at maturity, LEMMA chartaceous, 3-nerved, tapering into a long slender awn. PALEA 2-nerved, closely clasped by the lemma, GRAIN cylindric, 1 to 2 cm long, STAMINATE SPIKELET soft. LEMMA 5-nerved, membranous, linear, acuminate or awn-pointed, PALEA about as long as the lemma, 3-nerved. STAMENS 6. Tall aquatic annuals or perennials, with flat blades and large terminal panicles, the lower branches ascending or spreading, at maturity erect, bearing appressed pistillate spikelets, the staminate spikelets early deciduous, the pistillate spikelets tardily deciduous. SPIKELETS

GLUMES

1.

Zizania latifolia (Griseb.) Stapf, Kew Bull. 1909:385 (1909).

FIG. 4. Oryza sativa L. Plant, X 1/2; spikelet, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U.S.D.A. photograph.)

38

GRASSES

OF

HAWAII

ARISTIDEAE C. E. Hubb. ex Bor, Grasses of Burma, Ceylon, India and Pak. 685 (i960). Spikelets all alike, bisexual, 1-flowered; rachilla disarticulating above the glumes, not produced beyond the floret. Glumes usually persistent, scarious, 1-3-nerved, equal or unequal, rarely the lower longer than the upper, sometimes awned, usually muticous or mucronate; lemma convolute or involute, at first chartaceous, becoming firmer, cylindric, faintly 3-nerved, entire or bifid at the tip, with a bearded pungent or rarely forked callus, awned, palea small, oblong, hyaline, 2-nerved; awn nearly always 3-partite from the base or above the base, jointed to the tip or to a column or continuous with the lemma; column straight or twisted, bristles plumose or the lateral plumose, or all naked. Lodicules 2. Stamens 3 or 1. Ovary glabrous; styles distinct, short; stigmas plumose, laterally exserted. Grain slender, terete or oblong-terete, tightly embraced by the lemma; hilum linear, almost as long as the grain; embryo short or long; starch grains compound. Annual or more often tufted perennial grasses. Leaf blades narrow, often convolute, with panicoid anatomy; silica bodies dumbbell-shaped or oblong; microhairs, when present, 1-celled, rarely 2-celled, and then elongate, slender; ligule a fringe of hairs. Spikelets pedicelled in effuse or contracted panicles. Chromosomes small; basic number 11 or 12. Genus:

Aristida

Aristida L., Sp. PI. 82 (1753). SPIKELETS solitary, pedicelled, not falling entire, FLOWER I . GLUMES persistent, subequal or unequal, shorter than or as long as the lemma, linear in outline, acute, subacute, or obtuse, sometimes bifid, acuminate, mucronate, or short- to long-awned, often scabrid on the keel, membranous, glabrous or minutely pubescent; lower glume shorter than the upper, C A L L U S pungent, villous, L E M M A I , often convolute, sometimes oblong and bilobed and hardly enveloping the flower, gradually tapering to tip, sometimes jointed at or above the middle, 3-nerved, awned, subtending a palea and the bisexual flower; awn nearly always 3-partite from the base or above the simple base (stipitate), very rarely simple, continuous with the lemma or disarticulating from it or deciduous with a portion of the lemma, straight or twisted at the foot, with the lobes equal or the middle lobe the longest, sometimes the two side awns

ARUNDINOIDEAE -

ARISTIDEAE

39

hairlike, as long as to about 6 times as long as the body of the lemma, smooth, scabrid, or the middle lobe only or all the lobes plumose, PALEA shorter than the lemma, 2-nerved or nerveless, hyaline, LODICULES 2, finely nerved, STAMENS 3. OVARY glabrous; styles free, plumose above, laterally exserted. GRAIN slender, cylindrical, sometimes grooved, tightly embraced by the lemmas. Annuals or perennials, sometimes caespitose, frequently with thick woody roots; culm finely striate or ribbed, sometimes subwoody, glabrous, rarely villous at the nodes; ligule usually a line of very short hairs; blade rolled, deeply channelled or flat and narrow-linear, sometimes setaceous, smooth or rough, glabrous; inflorescence a lax or contracted panicle, more rarely a raceme or spikelike. 1.

Aristida adscensionis L., Sp. PI. 82 (1753). [Fig. 5] A. divaricata Jacq., Eclog. Gram. Rar. 7 (1813).

4°

GRASSES OF

HAWAII

V \I

WK

A.R-C

F i g . 5. Aristida adscensionis L . J, Plant, x 1/2; 2, spikelet f r o m side, x 4. 3, L o w e r g l u m e ; 4, u p p e r g l u m e ; 5, l e m m a ; 6, stamens, ovary, and l o d i c u l e s ; all X 6. ( F r o m N . L . Bor, T h e G r a s s e s of B u r m a , C e y l o n , India and Pakistan.)

ARUNDINOIDEAE -

ARUNDINEAE

41

ARUNDINEAE Dumort., Obs. Gram. Belg. 82, 124 (1823) as Arundinaceae. Spikelets 2-io-flowered, somewhat laterally compressed, bisexual or unisexual, dioecious; rachilla disarticulating above the glumes and between the florets, sometimes penicillate with long white hairs; florets all male or all bisexual or the lowest male or sterile in a few genera. Glumes hyaline or membranous, acuminate, persistent; lemmas somewhat similar to the glumes, 1-3-nerved, acuminate, awnless or awned from the tip, often bearing long white hairs from the back or sides or lowest quite glabrous; palea short, linear-oblong, 2-keeled. Lodicules 2 or 3, 2-3-nerved, obovate, glabrous. Stamens 2 or 3. Ovary glabrous; styles 2; stigmas 2, plumose. Grain loosely enclosed by lemma and palea, oblong, terete; hilum oblong, short, basal. Starch grains compound. Perennial reeds. Leaf blades broad, long, often auriculate, with usually panicoid anatomy; silica bodies cross-shaped; microhairs 2-celled, long, narrow; ligule a row of stiff hairs. Spikelets pedicelled, arranged in large plumose panicles. Chromosomes small; basic number 12(F). Genera:

Arundo Cortaderia KEY TO THE GENERA OF ARUNDINEAE

i.

Plants dioecious; leaves crowded at the base of the stems Cortaderia p. 42

1.

Plants not dioecious; spikelets bisexual; leaves distributed along the culms Arundo p. 41 Arundo L., Sp. PI. 81 (1753). SPIKELETS solitary, pedicelled, not falling entire, FLOWERS 4. GLUMES persistent, equal, similar, enclosing the lemmas, lanceolate, acuminate, subobtuse, 3-nerved, membranous, glabrous, LEMMAS 4, decreasing in size upwards, similar, lanceolate, acuminate, 2-lobed, with a short awn from the sinus, membranous, dorsally long-villous, each lemma subtending a palea and a bisexual flower, PALEAS shorter than the lemmas, more or less oblong, truncate, 2-keeled, hyaline, ciliate on the keels, LODICULES 2. STAMENS 3. STYLES distinct, plumose above, GRAIN oblong, enclosed by the lemma and palea, glabrous; embryo large.

42

GRASSES OF

HAWAII

A robust perennial u p to 15 f t high; culm smooth, hollow, manynoded, simple or scantily branched; leaf sheath slightly exceeding the internode of the culm, smooth; ligule short, membranous; blade up to 2 ft long, broad, linear-lanceolate from a broad base, tapering to a very fine point, smooth; inflorescence a large contracted or effuse panicle. 1.

Arundo donax L., Sp. PI. 81 (1753). [Fig. 6]

2.

A. donax var. versicolor (Miller) (1812).

Stokes, Bot. Mat. Med. 1:160

Cortaderia Stapf, Gard. Chron. ser. 3, 22 (pt. 2)1396 (1897) nom. cons. Moorea Lemaire, 111. Hort. 2: Misc. 15 (1854) in obs. Dioecious or semidioecious; SPIKELETS solitary, not falling entire; rachilla disarticulating near the base of the internodes, the lower shorter part glabrous, the upper longer part bearded and forming a stipe to the floret, the rachilla and lemmas clothed with long silky hairs, GLUMES longer than the lowest lemma (excluding the awn), sometimes longer than all the lemmas, LEMMAS several, tapering into a slender point or awn, or awned from between the slender teeth of the bifid apex. Mostly large tussocky grasses, the leaves generally crowded at the bases of the culms, the blades long and narrow, tapering into a slender point, harshly scabrid on the margins; inflorescence a silvery-silky panicle usually large and plumelike. 1.

Cortaderia selloana (Schult.) Aschers. and Graebn., Syn. Mitteleur. Fl. 2:325 (1900). C. argentea (Nees) Stapf, Gard. Chron. ser. 3, 22:396 (1897).

ARUNDINOIDEAE -

ARUNDINEAE

43

FIG. 6. Arundo donax L. i, Portion of stem, X 1/3 ; 2, ligule, X 1 ; 3, inflorescence, X 1/4. 4, Spikelet; 5, upper glume; 6, lower glume; 7, lemma; all X 4. 8, Palea, X 6; 9, flower and lodicules, X 6. (From N. L. Bor, The Grasses of Burma, Ceylon, India and Pakistan.)

44

GRASSES

ARUNDINELLEAE Stapfm

OF

HAWAII

Dyer, Fl. Cap. 7:314 (1898).

Spikelets all alike, 2-flowered, with the lower floret male or sterile and the upper bisexual; rachilla disarticulating between the florets. Glumes persistent, unequal, the lower smaller, usually acute, the upper acuminate, membranous to coriaceous; lower lemma persistent, similar in texture to the upper glume, 3-9-nerved; palea absent; upper lemma more or less terete, smaller than the lower, firmer in texture, awnless or with a very short bristle or more often awned from the minutely 2-lobed tip, rarely accompanied by 2 bristles, one on either side at the tip; awn kneed and twisted below the knee; palea delicate, 2-keeled. Lodicules 2, cuneate. Stamens 3. Ovary glabrous with 2 distinct styles and plumose stigmas. Grain oblong or ellipsoid, tightly embraced by lemma and palea; hilum punctiform or linear; embryo about half the length of the grain; starch grains compound. Annual or perennial herbs. Leaf blades linear to lanceolate, frequently large, usually flat, with panicoid anatomy; silica bodies quadrate, crossshaped or dumbbell-shaped; microhairs 1- or 2-celled; ligules very short, membranous or ciliate. Inflorescence of pedicelled spikelets often paired, arranged in large, effuse or contracted panicles. Chromosomes small; basic number 7. Genus:

Arundinella

Arundinella Raddi, Agrost. Bras. 36 (1823). SPIKELETS solitary on very short pedicels or in pairs and unequally pedicelled, when in pairs then both spikelets alike or the shorter pedicelled one sterile and consisting of a scabrid glume, FLOWERS usually 2; the upper bisexual; the lower, when present, male or rarely bisexual. GLUMES persistent, membranous; lower ovate, acuminate, acute, 3 6-nerved, somewhat keeled and scabrid on the keel; upper similar to the lower, L E M M A S 2; lower boat-shaped, falcate, obtuse, 3-7-nerved, membranous, subtending a palea and a male (rarely bisexual) flower, sometimes flower absent; upper entire or minutely bifid, 3-7-nerved, minutely scaberulous, with the lobes sometimes produced into fine bristles about twice as long as the lemma, PALEA linear, 2-keeled, with the flaps more or less auricled. L O D I C U L E S 2, cuneate, rather fleshy. S T A M E N S 3. O V A R Y oblong in outline; styles distinct or connate at the base, plumose above, laterally exserted. G R A I N oblong in outline.

ARUNDINOIDEAE -

ARUNDINELLEAE

45

Perennials; culm finely ribbed, glabrous; leaf sheath as long as or shorter than the internodes of the culm, ribbed, glabrous; ligule extremely short, ciliate; blade linear, produced into a long fine point, with a distinct or prominent midrib, and close but distinct lateral veins, sometimes scabrid, otherwise glabrous. i.

Arundinella holcoides (Kunth) Trin., Bull. Acad. Sci. Petersb. 1 : 7 1 (1836). A. agrostoides Trin., Sp. Gram. Ic. 3 (1836). According to Hitchcock (1922), 'Arundinella agrostoides Trin. is included doubtfully by Hillebrand (1888) but should be omitted as the locality "Oahu" is probably an error.'

46

GRASSES

OF

HAWAII

DANTHONIEAE Nevski, Act. Inst. Bot. Acad. Sci. U.R.S.S. ser. i, fasc. 4:223 (1937). Spikelets 2-io-flowered, all alike, bisexual, uppermost florets reduced; rachilla disarticulating above the glumes and between the florets, produced and crowned with a rudimentary floret. Glumes equal or subequal to or longer than the lowest lemma, hyaline to scarious or sub-herbaceous, 3-11-nerved; lemmas firmly membranous to coriaceous, 5-11-nerved, more or less hairy, 2- or 3-lobed, awned in the sinus or the middle lobe awned; lobes aristate or not; awn geniculate, with a twisted column; palea 2-keeled, hyaline, as long as the lemmas or shorter. Lodicules 2, large or small. Stamens 3, sometimes minute. Ovary glabrous; styles 2, slender, long. Grain oblong, obovoid or ellipsoid; hilum obscure, linear. Starch grains compound. Perennials, rarely annuals; leaf blades narrowly linear or setaceous, rolled or flat, with festucoid anatomy; silica bodies rounded or dumbbellshaped; microhairs slender, 2-celled. Spikelets arranged in panicles, racemes, or spikes. Ligule a fringe of hairs. Chromosomes small; basic numbers 6, 9. Genera:

Astrebla Danthonia KEY TO THE GENERA OF DANTHONIEAE

1.

Lemmas 3-lobed, the central tapering into an awn

2.

Lemmas 2-lobed, awned in the sinus

.

.

Astrebla p. 46 Danthonia

P- 47 Astrebla F. Muell. in Benth., Fl. Austral. 7:602 (1878). SPIKELETS solitary, sessile or subsessile, laterally compressed or subterete, distichously imbricate, not falling entire; rachilla disarticulating above the glumes but not between the lemmas, FLORETS 2-9, the lowest 2 - 4 bisexual, the uppermost frequently reduced, GLUMES slightly unequal, linear to elliptical, strongly compressed or keeled, usually acute or acuminate, the lower 2-9-nerved, the upper 7 - 1 3 nerved. L E M M A S 2-9, the lowest 2-4 alone fertile, exserted from the glumes, coriaceous, rounded on the back above the middle, silky-villous at the base not always so on the dorsal surface, deeply 3-lobed, the middle lobe tapering from a broad base into a tough straight, curved

ARUNDINOIDEAE -

DANTHONIEAE

47

or hooked bristle or awn; the lateral lobes similar to the middle lobe or broader and flattened, varying from linear to lanceolate or semi-ovate, toughly coriaceous, or scarious and shining, 1-5-nerved; callus obsolete. PALEA shorter than the lemma, lanceolate to elliptical in outline, acuminate, dorsally compressed, firmly membranous to chartaceous, 2-keeled, the keels ciliate. LODICULES 2, truncate, STAMENS 3. OVARY glabrous; styles distinct, short, terminal; stigmas plumose, terminally exserted. GRAIN oblong or elliptical in outline, dorsally compressed, loosely enclosed by the hardened lemma and palea; hilum basal; embryo small. Tufted perennials; leaf blade narrow; ligule reduced to a ciliate rim; inflorescence of terminal or paired terminal spikes or spikelike racemes, the rachis continuous, triquetrous, tough. 1.

Astrebla elymoides Bailey et Muell. in F. M. Bailey, Grasses Queens. 1:8 (1888).

2.

A. lappacea (Lindl.) Domin., Bibl. Bot. 85:372 (1915).

3.

A. pectinata (Lindl.) F. Muell. in Benth., Fl. Austral. 7:602 (1878).

Danthonia DC. in Lamk. et DC., Fl. Fr. 3:32 (1805). solitary, pedicelled or subsessile, not falling entire; pedicels 2-8; the lower bisexual; the upper usually male. GLUMES persistent, equal or subequal, longer or shorter than the lemmas, linear or lanceolate, more rarely ovate, sometimes long-acuminate, obtuse or acute, keeled, 3-9-(rarely i-)nerved, often with transverse veins, hyaline to scarious or subherbaceous, glabrous, LEMMAS 2-8, varying much in shape, usually decreasing in size upwards, 2-lobed, usually awned from the sinus and sometimes with a fine awn from each lobe; the uppermost lemmas sometimes much reduced, or empty, or subtending male flowers, NORMAL LEMMAS firmly membranous to chartaceous, 7-11-(rarely 5-)nerved, sometimes the 2 lobes reduced to minute teeth and the awn to a mucro, hairy, often with the hairs partly gathered in variously arranged tufts, very rarely glabrescent, subtending a palea and a bisexual flower; lobes of the lemma, when present, free, rarely more or less adnate to the awn, with or without a bristle at the tip; central awn, when present, 1/2 to 3 times as long as the body of the lemma and the lateral awns, when present, slightly longer than the body of the lemma, PALEAS lanceolate or oblong or spathulate, obtuse, 2-keeled, ciliate on the keels, subhyaline. LODICULES 2, usually SPIKELETS

scabrid.

FLOWERS

48

GRASSES OF H A W A I I

large and ciliate. STAMENS 3. OVARY glabrous; styles free, plumose above, laterally exserted. GRAIN oblong, obovate, or elliptic in outline; embryo 1/3-1/2 the length of the grain. Perennials, more rarely annuals, often villous at the base; culm ribbed, glabrous; leaf sheath ribbed, glabrous; ligule ciliate; blade long or short, narrow-linear, often curled, usually rolled, more rarely flat, ribbed or almost smooth, sometimes rough, glabrous, more rarely pilose, pubescent on the ribs; inflorescence a contracted panicle, sometimes compact and more or less globose, more rarely a dense distichous spike. 1.

Danthonia pilosa R. Br., Prodr. Fl. Nov. Holl. 177 (1810).

2.

D. racemosa R. Br., Prodr. Fl. Nov. Holl. 177 (1810).

3.

D. semiannularis (Labill.) R. Br., Prodr. Fl. Nov. Holl. 177 (1810).

ARUNDINOIDEAE -

GARNOTIEAE

49

GARNOTIEAE Tateoka, Journ. Jap. Bot. 32:277 (1957). Spikelets strictly 1-flowered, lanceolate, cylindrical or ellipsoid in shape, falling entire from the pedicels; rachilla not produced beyond the solitary floret. Glumes as long as the spikelet, strongly 3-nerved, bearded or not at the base, beaked, acute or rarely awned; lemmas membranous or hyaline, 3-5-nerved, entire or bifid at the tip, if the latter then an awn issuing from between the lobes which are rarely awned; palea 2-nerved, narrow, hyaline. Lodicules 2, ovate-lanceolate or cuneate. Stamens 3. Ovary glabrous at the top; styles 2; stigmas plumose. Grain narrow, free between lemma and palea; hilum basal, oblong or punctiform; embryo about 1/3 the length of the grain. Erect, perennial, rarely annual, grasses, with narrow, acuminate or acute leaves; silica bodies variable, sometimes saddle-shaped or dumbbell-shaped; microhairs 2-celled, threadlike; ligules a ciliate rim or a narrow ciliate membrane. Spikelets borne in panicles, falling entire from the pedicels. Chromosomes small; basic number 10. Genus:

Garnotia

Garnotia Brongn. in Duperrey, Voy. Coq. Bot. 132 (1832). SPIKELETS 1-flowered, the pedicel disarticulating below the glumes; in unequally pedicelled pairs; spikelets nearly terete, base of spikelet pilose, GLUMES narrow, about equal, firm, somewhat scaberulous on the nerves and back; the first 1-nerved, the second 3-nerved. LEMMA firm, usually awned at the apex, PALEA about as long as the lemma, 2-nerved, awnless, usually with 2 auricles at the base, GRAIN enclosed in the slightly stiffened lemma and palea. Annual or perennial grasses with flat blades and narrow panicles.

1.

Garnotia sandwicensis Hillebr., Fl. Haw. Is. 513 (1888). [Fig. 7]

50

GRASSES OF H A W A I I

51

FESTUCOIDEAE - AGROSTIDEAE

AGROSTIDEAE Dumort., Obs. Gram. Belg. 83, 125 (1823). Spikelets all alike, bisexual, 1-flowered, rarely falling attached to the pedicel; rachilla disarticulating above, rarely below the glumes, sometimes produced beyond the floret as a glabrous or penicillately hairy bristle; glumes sometimes persistent, usually as long as or longer than the lemma, sometimes shorter; lemmas hyaline or membranous, similar in texture to the glumes or thinner, 3-5-nerved, awnless or awned from the back or from the 2-lobed tip; awn geniculate or straight; palea usually delicate, often small or almost absent, 2-nerved. Lodicules 2, rarely o. Stamens 3, 2, or 1. Grain often tightly enclosed between the lemma and palea; hilum linear, short or up to 2/3 the length of the grain; embryo small; starch grains compound. Annual or perennial herbs, usually with slender culms. Leaf blades most often narrow, with festucoid anatomy; silica bodies oblong or rounded; 2-celled microhairs absent; ligule membranous, glabrous. First foliage leaf of the seedling erect, narrow. Spikelets in open or contracted, sometimes spikelike panicles. Chromosomes large; basic number 7. Genera:

Agropogon

Dichelachne

Agrostis

Gastridium

Alopecurus

Lycurus

Ammophila

Muhlenbergia

Apera

Phleum

Calamagrostis

Polypogon

KEY TO THE GENERA OF AGROSTIDEAE

x.

Glumes saccate at the base

1.

Glumes not saccate 2.

Gastridium p. 65

Glumes long or shortly awned 3.

Glumes long-awned; spikelets deciduous

Polypogon p. 68

3.

Glumes shortly awned; spikelets persistent .

Agropogon P- 53

52

GRASSES OF H A W A I I

2.

Glumes not awned 4.

Inflorescence a dense spikelike panicle with overlapping spikelets 5. Spikelets large (10-16 mm long); a stout tall grass with strong rhizomes; leaf blades pungent, tightly inrolled Ammophila p. 60 5.

4.

Spikelets much smaller, herbaceous grasses; leaf blades flat 6.

Spikelets in pairs, one male, one biseuxal, on deciduous branchlets Lycurus p. 66

6.

Spikelets all alike, bisexual 7.

Glumes acute; lemma awned

7.

Glumes truncate; lemma awnless

.

.

Alopecurus p. 60

.

.

Phleum p. 68

Inflorescence often effuse if dense, not cylindrical or spikelike 8.

Glumes lanceolate-acuminate, much longer than the lemmas 9.

Lemma long awned from just below the tip Dichelachne p. 62

9.

Lemma shortly awned from the back or tip Calamagrostis p. 62

8.

Glumes not acuminate 9.

Lemma indurated, awned or not from the tip Muhlenbergia p. 66

9.

Lemmas not indurated 10. Lemma awned from the tip

.

.

.

Apera p. 60

10. Lemma awnless or very shortly awned from the back Agrostis P- 53

FESTUCOIDEAE -

AGROSTIDEAE

53

Agropogon P. Fourn., Quatre Fl. France, 50 (1934) in clavi. One species described as follows: Agropogon littoralis (Smith) C. E. Hubb., Journ. Ecol. 33:333 (1946). [This is an intergeneric hybrid between Polypogon monspeliensis (L.) Desf. and Agrostis stolonifera L.] SPIKELETS persistent, narrowly oblong, or gaping, 2-3 mm long, laterally compressed, i-flowered. GLUMES similar, narrowly oblong or elliptic, narrowed upwards, minutely notched at the tip, rounded on the back below, keeled above, membranous, 1-nerved, rough-pointed, especially in the lower half, awned from the tip, with the fine straight awn up to about 2 mm long, L E M M A between 1 ¡2 and 2/3 the length of the glumes, elliptic oblong, with a very blunt minutely toothed tip, smooth, very thin, obscurely 5-nerved, awned from the back just below the tip, with the awn up to 3 mm long, PALEA about 3/4 the length of the lemma, finely 2-nerved. ANTHERS about 1 mm long. Chromosome number 2n = 28. Perennial, loosely tufted or creeping, 8-60 cm high. Culms kneed at the base and then erect, or ascending from a prostrate many-noded base, usually branched in lower part, slender, smooth. Leaves hairless, grayish-green, sheaths smooth; ligules blunt, becoming toothed, 3-7 mm long, membranous; blades finely pointed, flat, 3-20 cm long, 2-11 mm wide, rough towards the tip, or all over. Panicles erect, moderately to very dense, lanceolate to narrowly ovate, or oblong, more or less lobed, 2-18 cm long, 0.6-7 c m wide, green or purplish; branches closely divided, rough; pedicels very short.

Agrostis L„ Sp. PI. 6 (1753). SPIKELETS solitary, pedicelled, not falling entire, GLUMES persistent, equal or subequal, as long as to almost twice as long as the lemma, lanceolate, linear, or oblong-lanceolate, sometimes acuminate, acute or obtuse, usually ciliate on the keel, sometimes scabrid, membranous glabrous, L E M M A I , lanceolate or linear, acute or obtuse, sometimes truncate and notched at the apex, awned from the back or awnless, 5-(rarely 3-)nerved, with the side nerves evanescent below and often excurrent into fine mucros or bristles, delicately membranous, glabrous or hairy, PALEA usually shorter than the lemma, sometimes much reduced or obsolete, 2-nerved, delicately hyaline, L O D I C U L E S 2, lanceolate, delicately hyaline, STAMENS 3. O V A R Y ellipsoid; styles free, very short, plumose above, laterally exserted. G R A I N free, oblong in outline, enclosed

54

GRASSES OF H A W A I I

in the scarcely altered lemma and palea; embryo short. Annuals or perennials of varying habitat; culm 2-many-noded, sometimes the lower nodes rooting, smooth or slightly striate or more or less rough below the nodes; leaf sheath ribbed, rough, glabrous; ligule long (may be very short, e.g., A. tenuis), membranous; blade narrowlinear, flat or rarely folded, acuminate, with a distinct midrib, scabrid or smooth, glabrous; inflorescence an open, sometimes effuse, panicle, more rarely spikelike; primary branches of the inflorescence often fascicled on the main rachis, naked at, or bearing spikelets almost to, the base, often scabrid. 1. 2.

3.

4. 5. 6. 7. 8.

9. 10. 11. 12.

13. 14.

Agrostis alba L., Sp. PI. 62 (1753). [Fig. 8] Agrostis avenacea Gmel., Syst. Nat. 2:171 (1791). A. retrofracta Willd., Enum. PI. 94 (1809). Calamagrostis retrofracta (Willd.) Link, Hort. Berol. 2:247 (1833). Deyeuxia forsteri Kunth, Rev. Gram. 1:77 (1829). D. retrofracta (Willd.) Kunth, Rev. Gram. 1:77 (1829). C. forsteri Steud., Nom. Bot. ed. 2, 1:250 (1840). A. borealis Hartm., Handb. Skand. Fl. ed. 3, 17 (1838). ? A. rubra L., Sp. PI. 62 (1753). (Hitchc. and Chase, 804 [1950], 'Identity uncertain.') A. canina L., Sp. PI. 62 (1753). A. capillaris L., Sp. PI. 62 (1753). A. exarata Trin., Gram. Unifl. 207 (1824). [Fig. 9] A. fallax Hillebr., Fl. Haw. Is. 516 (1888). [Fig. 10] A. microphylla Steud., Syn. PI. Glum. 1:164 (1854). A. exarata microphylla (Steud.) Hitchc., Amer. Journ. Bot. 2:303 (I9I5)A. palustris Huds., Fl. Angl. 27 (1762). A. sandwicense Hillebr., Fl. Haw. Is. 515 (1888). [Fig. 11] A. rockii Hack, in Fedde, Repert. Sp. Nov. 10:167 (1911). A. scabra Willd., Sp. PI. 1:370 (1797). A. semiverticillata (Forsk.) C. Christ, in Dansk. Bot. Arkiv 4(3) : 1 2 (1922). [Fig. 12] A. kauaiense Hillebr., Fl. Haw. Is. 516 (1888). A. verticillata Vill., Prosp. PI. Dauph. 16 (1779). A. stolonifera L., Sp. PI. 62 (1753). A. alba auctt. non L., Sp. PI. 63 (1753). A. tenuis Sibth., Fl. Oxon. 36 (1794). A. vulgaris With., Bot. Arr. Veg. Brit. ed. 3, 2 : 1 3 2 (1796).

FIG. 8. Agrostis alba L. (a) Scabrous branchlets of panicle, X 6;(b) leaf sheath, ligule, and blade, X 6; (c) habit, showing the decumbent culms and flat leaf blades, X 1/3; (d) floret, showing lemma, X 20; (e) floret, showing the short emarginate, palea, X 20; (f) and (g) grains (caryopses), X 20; (h) habit, upper part of culm, showing panicle, X 1/3; (i) spikelet, showing the glumes, each with scabrous keel, X 16. (From H. L . Mason, A Flora of the Marshes of California.)

FIG. 9. Agrostis exarata Trin. (a) Leaf sheath, Iigule, and blade, x 5; (b) spikelets in lower part of panicle, x 3; (c) habit, showing the leafy culms and young, close panicle, X 2/5; (d) upper part of culm, showing panicle, X 2/5; (e) floret, X 15; ( f ) spikelet, each glume with a scabrous keel, x 15. (From H. L . Mason, A Flora of the Marshes of California.)

FIG. IO. Agrostis fallax Hillebr. (From A. S. Hitchcock, The Grasses of Hawaii.)

rrp. ••

".il

FIG. i l . Agrostis sandwicense Hillebr. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

FIG. 12. Agrostis semiverticillata (Forsk.) C. Christ, (a) Habit, showing culms with decumbent base, short, horizontal leaf blades, and panicles, X 1/5; (b) young floret, showing the truncate lemma, toothed at apex, X 12; (c) branchlets of panicle, showing inflated base, x 4; (d) flowering spikelet, the glumes scabrous, X 20; (e) and ( f ) leaf sheath, dentate ligule, and scabrous blade, x 4. (From H. L. Mason, A Flora of the Marshes of California.)

6o

GRASSES OF

HAWAII

Alopecurus L., Sp. PI. 60 (1753). SPIKELETS solitary, pedicelled or subsessile, falling entire, FLORET I , bisexual, GLUMES equal, acute, unawned, keeled, 3-nerved, the margins united towards the base, or free, LEMMA I , equal to or shorter than the glumes, thin, keeled, 5-nerved, obtuse, the margins united towards the base, awned from below the middle or from about the middle, the awn slender, erect or geniculate, usually 2 or 3 times as long as the lemma. PALEA none, LODICULES wanting, STAMENS 3. OVARY glabrous; styles distinct; stigmas plumose, apically exserted. GRAIN ovate to lanceolateovate in outline, somewhat flattened, included in the dehiscent lemma but free from it. Perennial or annual grasses; leaf blade flat, usually flaccid, convolute in aestivation; ligule membranous, obtuse; inflorescence a dense, soft, spikelike panicle.

1.

Alopecurus pratensis L., Sp. PI. 60 (1753). [Fig. 13]

Ammophila Host, Gram. Austr. 4:24 (1809). SPIKELETS solitary, pedicelled, not falling entire, compressed; rachilla disarticulating above the glumes and produced as a short bristle beyond the palea, FLORET I , bisexual, GLUMES subequal, chartaceous, narrow, acutely acuminate, keeled, thinly 5-nerved. LEMMA subequal to, or slightly longer than the glumes and almost similar to them in shape and texture, acute or subacute, with or without a small tooth near the apex; callus bearded with rather long silky hairs, PALEA equal in length to the lemma, thin, 2-keeled and bidentate, deeply sulcate between the keels. LODICULES 2. STAMENS 3. OVARY glabrous; styles distinct; stigmas plumose, laterally exserted or enclosed, GRAIN obovoid, included in, but free from the lemma and palea. Tall stoloniferous perennials of rather coarse texture, erect, with hard scaly creeping rhizomes and deep roots; leaf blades long, linear, tough, involute; ligule hyaline, long and large; inflorescence a narrow and dense spikelike panicle.

1.

Ammophila arenaría (L.) Link, Hort. Berol. 1:105 ( I ^27).

Apera Adans., Fam. PI. 2:495 (1763). SPIKELETS I-flowered, disarticulating above the glumes, the rachilla prolonged back of the palea as a naked bristle, GLUMES subequal,

6i

FIG. 13. Alopecurus pratensis L . Plant, x 1/2; glumes and floret, x 10. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

62

GRASSES

OF

HAWAII

acuminate, L E M M A firm, subindurate at maturity, acute, bearing a long delicate straight awn just below the tip. PALEA nearly as long as the lemma, strongly 2-nerved. Annuals with flat blades and loose or narrow panicles. i.

Apera interrupta (L.)

Beauv., Ess. Agrost. 31, 151 (1812).

Agrostis interrupta L., Syst. Nat. ed. 10, 2:872 (1759). Calamagrostis Adans., Fam. PL 2:31 (1763). solitary, shortly pedicelled, not falling entire, FLOWER I . persistent, equal or subequal, linear-lanceolate, long-acuminate, acute, keeled, membranous; lower glume 1-nerved; upper glume 3nerved. L E M M A I , shorter than the glumes, lanceolate, subacute, more or less shortly bifid, 3-5-nerved, awned from near the base, membranous, glabrous; awn longer or shorter than the glumes, kneed or straight. CALLUS small, long hairy, with the hairs usually much exceeding the lemma, PALEA as long as or shorter than the lemma, 2-nerved, hyaline. LODICULES 2, hyaline, STAMENS 3. O V A R Y glabrous; styles free, plumose above, laterally exserted. GRAIN enclosed by the hardly changed lemma and palea, free; embryo small. SPIKELETS

GLUMES

Usually robust perennials mostly with creeping rhizomes; culm striate, glabrous; leaf sheath ribbed, glabrous; ligule scarious; blade linear, flat, ribbed, glabrous; inflorescence a spikelike panicle. 1.

Calamagrostis canadensis var. scabra (Presl) Journ. Bot. 21:135 (1934).

2.

C. expansa (Munro)

Hitchc.,

Hitchc.,

M e m . Bernice P. Bishop

Amer. Mus.

8:149 (1922)- [ F i s - H ] Deyeuxia expansa Munro in Hillebr., Fl. Haw. Is. 519 (1888). 3.

C. hillebrandii (Munro)

Hitchc., M e m . Bernice P. Bishop Mus.

8:147 (1922). [Fig. 15] Deyeuxia hillebrandii Munro in Hillebr., FL Haw. Is. 519 (1888). 4.

C. langsdorfi (Link)

Trin., Gram. Unifl. 225 (1824).

Dichelachne Endl., Prodr. FL Ins. Norf. 20 (1833). SPIKELETS solitary, pedicelled, narrow-lanceolate, acuminate, not falling entire; rachilla disarticulating above the glumes, shortly produced or not at all produced beyond the flower, FLOWER I . GLUMES subequal,

FIG. 14. Calamagrostis expansa (Munro) Hitchc. (From A. S. Hitchcock, The Grasses of Hawaii.)

64

GRASSES OF H A W A I I

FIG. 15. Calamagrostis hillebrandii (Munro) Hitchc. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

FESTUCOIDEAE -

AGROSTIDEAE

65

narrow, membranous, acuminate, keeled, scabrous on the keels, LEMMA much shorter than the glumes, rounded on the back or more or less keeled, entire or bifid, faintly nerved, the teeth when present, erect, scarcely acute; AWN arising from below the apex of the lemma or from the sinus of the lobes, straight or flexuose and more or less spirally twisted; callus short, blunt, hairy, PALEA narrow, hyaline, slightly shorter than the lemma, 2-keeled. LODICULES 2. STAMENS 3 or 2. OVARY glabrous; styles short, distinct, stigmas plumose, included or laterally exserted. GRAIN narrow, pale, shining, subterete; embryo small. Annuals, tall, slender, caespitose grasses. Leaf blades narrow, flat or setaceous-convolute; ligule white, hyaline; inflorescence a dense cylindrical spikelike panicle, or loose and almost secund. 1. 2.

Dichelachne montana Endl., Prod. Fl. Ins. Norf. 20 (1833). D. sciurea (R. Br.) Hook, f., Fl. New Zealand 294 (1853). D. micrantha (Cav.) Domin., Bibl. Bot. 85:353 (1915).

Gastridium Beauv., Ess. Agrost. 21 (1812). SPIKELETS solitary, unequally pedicelled, not falling entire; pedicel linear, thickening above, GLUMES persistent, unequal, slightly saccate near the base, membranous; lower glume longer than the upper, lanceolate, long-acuminate, with membranous margins, 1-nerved, somewhat keeled, scabrid on the keel; upper glume similar to the lower, LEMMA 1, elliptic, rounded on the back, concave on the face, awned from the back, pilose; awn capillary, 5 times as long as the body of the lemma, scabrid. PALEA shorter than lemma, elliptic 2-lobed, 2-nerved, pilose on the back, LODICULES 2, hyaline, OVARY subglobose; styles free, plumose above. Annuals; culm smooth, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous; ligule membranous; blade narrowlinear, flat, ribbed, scabrid, glabrous; inflorescence a number of racemes whorled on the central rachis and the whole forming a very compact spikelike panicle.

1.

Gastridium phleoides (Nees et Meyen) I 9S4 : 375

C.E.Hubb.,

K e w Bull.

(i954)Lachnagrostis phleoides Nees et Meyen, Nov. Act. Acad. Caes. Leop. Carol. 19 (Suppl. 1)1146 (1843).

66 2.

GRASSES

OF

HAWAII

G. ventricosum (Gouan) Schintz et Thell., Vierteljohrs. Nat. Ges. Zurich 58:39 (1913). Agrostis ventricosa Gouan, Hort. Reo. Monsp. 39 (1762). G. australe Beauv., Ess. Agrost. 21, 164 (1812). G. lendigerum (L.) Desv., Obs. Angers 48 (1818). Milum lendigerum L., Sp. PI. 91 (1762).

Lycurus H.B.K.,

Nov. Gen. et Sp. 1:141 (1815).

One species in Hawaii described as follows: Lycurus phleoides H.B.K.,

Nov. Gen. et Sp. 1:142 (1815). [Fig. 16]

SPIKELETS usually in pairs, occasionally in threes; to 5 m m long; awned; upper spikelet perfect, lower spikelet male or neuter; falling with the pedicel, GLUMES about 1/2 as long as lemmas, awned, awns unequal, more or less ciliate on the margins; the first glume oblong, thin, 2-nerved, about 1.5 mm long, with 2 or rarely 3 unequal scabrous awns, L E M M A S 3.5-4 mm long, lanceolate, acuminate; thicker than the glumes; 3-nerved; more or less pubescent; terminating in a scabrous awn as long as the lemma, PALEA lanceolate; 2-toothed; nearly as long as its lemma; thinly pubescent. A tufted, slender perennial grass, to 60 cm in height. Culms erect or ascending, sometimes decumbent with a slightly thickened or bulbous base, freely branching, rather weak. Leaf blades mostly less than 10 cm long, about 1 - 3 m m wide, upper leaves shorter, radical leaves numerous and shorter, flat or folded, rough on the margins and on the upper surface toward the apex. Leaf sheaths much shorter than the internodes, loose. Ligule membranous. Panicles, dense, narrow, cylindrical, 3-6 cm long; often with an axillary panicle at each node, with peduncles sometimes as long as that of the terminal panicle.

Muhlenbergia Schreb., Gen. 44 (1789). SPIKELETS 1-flowered (occasionally 2-flowered), the rachilla disarticulating above the glumes, GLUMES usually shorter than the lemma, sometimes as long, obtuse to acuminate or awned, keeled or convex on the back, the first sometimes small, rarely obsolete, L E M M A firm-membranaceous, 3-nerved (the nerves sometimes obscure or rarely an obscure additional pair), with a very short callus, rarely long-pilose, usually minutely pilose, the apex acute, awned from the tip or just below it,

67

FIG. 16. Lycurus phleoides H . B . K . Plant, x 1/2; glumes and floret, x 10. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

68

GRASSES OF H A W A I I

from between very short lobes, sometimes only mucronate, the awn straight or flexuous. Sometimes only moderately tall or rarely robust grasses, tufted or rhizomatous, the culms simple or much branched, the inflorescence a narrow sometimes spikelike or open panicle. 1.

Muhlenbergia microsperma (DC.) Kunth, Rev. Gram. 1:64 (1829). [Fig. 17] Trichochloa microsperma DC., Cat. Hort. Monsp. 151 (1813). Agrostis microsperma Lag., Nov. Gen. et Sp. 2 (1816). M. fasciculata Trin., Gram. Unifl. 192 (1824).

2.

M. porteri Scribn. in Beal, Grasses N. Amer. 2:259 (1896).

3.

M. racemosa (Michx.) B.S. Presl, Cat. PI. N.Y. 67 (1888).

4.

M. repens (Presl) Hitchc. in Jepson, Fl. Calif. 1 : 1 1 1 (1912).

Phleum L., Sp. PI. 59 ( I 7 5 3 ) . SPIKELETS solitary, shortly pedicelled, laterally compressed, not falling entire; rachilla disarticulating above the glumes, shortly produced. FLORET 1, bisexual, GLUMES equal, membranous, oblong or oblong-cuneate in outline, abruptly truncate, strongly complicately keeled, the keel produced into a mucro or short awn, 3-5-nerved, usually ciliate along the keel, LEMMA much shorter than the glumes, hyaline, ovoid or elliptical, keeled, truncate or denticulate, 3-5-nerved, unawned. PALEA nearly as long as the lemma, narrow, hyaline, LODICULES minute. STAMENS 3. OVARY ovoid, glabrous; styles distinct, rather long; stigmas plumose, apically exserted. GRAIN ovoid, free from, but included in, the unaltered lemma and palea. Annuals or perennials, erect; leaf blade long, linear, flat; convolute in the bud; ligule white, membranous; inflorescence a dense cylindrical spikelike panicle.

1.

Phleum pratense L., Sp. PI. 59 (1753). [Fig. 18]

2.

P. subulatum (Savi) 2:154(1899).

Aschers. et Graebn., Syn. Mitteleur. Fl.

Polypogon Desf., Fl. Atlant. 1:66 (1798). SPIKELETS solitary, subsessile, falling entire, GLUMES equal or subequal, lanceolate, longer than the lemma, keeled, entire or shortly 2-lobed,

FESTUCOIDEAE -

AGROSTIDEAE

69

FIG. 17. Muhlenbergia microsperma (DC.) Kunth. A, Plant, x 1/2; B, fascicle of cleistogamous spikelets, x 1 . 5 ; C, glumes of terminal spikelet, x 1 0 ; D, prophyllum, X 5 ; i , terminal spikelet, X 5. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

FIG. 18. Phleum pratense L. Plant, x 1/2. (U.S.D.A. photograph.)

FESTUCOIDEAE -

AGROSTIDEAE

71

ciliate on the keel, membranous, each with a single awn, pubescent or glabrous; awns capillary, 2 - 1 2 times as long as the body of the glume. LEMMA 1, lanceolate or elliptic-oblong, truncate or emarginate, 5-nerved, with the lateral nerves shortly excurrent or evanescent, with the hyaline tip awned, glabrous; awn as long as to 6 times as long as the body of the lemma, sometimes awn reduced to a mucro or absent, often deciduous. PALEA slightly or much shorter than the lemma, lanceolate acute, 2-keeled. LODICULES 2, delicate, OVARY glabrous; styles free, very short, loosely plumose above, laterally exserted. GRAIN oblong in outline, enclosed by the unaltered lemma and palea; embryo small. Loosely tufted annuals or perennials, culm ribbed, glabrous; leaf sheath ribbed, rough; ligule membranous; blade flat, linear, closely ribbed, with a distinct midrib, scabrid, glabrous; inflorescence a dense terminal solitary spikelike panicle; spikelets often hidden by the numerous long awns. 1.

Polypogon interruptus H.B.K., Nov. Gen. et Sp. 1 : 1 3 4 (1815). Agrostis littoralis With., Bot. Arr. Veg. Brit, ed 3, 2:129 (1796). P. lutosus (Poir.) Hitchc., U. S. Dept. Agr. Bull. 772, 138 (1920); missapplied to P. interruptus H.B.K. (see Agropogon littoralis [Smith] C.E. Hubb.).

2.

P. monspeliensis (L.) Desf., Fl. Atlant. 1:67 (1798). [Fig. 19] Alopecurus monspelions (Sphalm) L., Sp. PI. 61 (1753). A. monspeliensis L., Sp. PI. 61 (1753). Phleum crinitum Schreb., Beschr. Graes. 1 : 1 5 1 (1769).

72

FIG. 19. Polypogon monspeliensis ( L . ) D e s f . 1, G r o u p of spikelets, x 3 ; 2, spikelet, x 17 J , L o w e r g l u m e ; 4, u p p e r g l u m e ; 5, l e m m a ; 6, p a l e a ; all x 12. ( F r o m D . M e r e d i t h , T h e G r a s s e s and Pastures of S o u t h A f r i c a . )

FESTUCOIDEAE -

AVENEAE

73

AVENEAE Dumort., Obs. Gram. Belg. 82, 120 (1823) as Avenaceae. Spikelets all alike, 2-7-flowered, more or less laterally compressed, with all the florets bisexual or the uppermost often sterile, rarely one or both of the two lowest florets male or neuter; rachilla disarticulating above the glumes and nearly always between the florets as well; callus often bearded. Glumes persistent, mostly equal, usually as long as the lowest lemma and often as long as the spikelet and enclosing the florets, membranous to chartaceous, frequently with shining margins; lemmas membranous to cartilaginous in texture, 5- to several-nerved, hyaline or scarious and shining on the margins, entire or 2-3-lobed at the tip, usually awned from the back or from between the lobes, rarely awnless; awn consisting usually of a twisted column and a bristle, kneed, rarely straight, scabrid; palea shorter, as long as the lemma, or slightly longer, hyaline, 2-keeled. Lodicules 2. Stamens 3, 2, or 1. Styles 2 or 1. Grain often tightly clasped by the lemma and palea; hilum linear, long; embryo up to 1/3 the length of the grain; starch grains compound. Annual or perennial herbs. Leaf blades narrow, with festucoid anatomy; silica bodies oblong or elliptic; no 2-celled microhairs; ligules well developed, membranous. Spikelets pedicelled and arranged in open or contracted panicles. Chromosomes large; basic number 4, 5, 7. Genera: Air a Deschampsia Anthoxanthum Arrhenatherum Avena

Holcus Sphenopholis Trisetum

KEY TO THE GENERA OF AVENEAE

i.

Lower floret male or neuter 2.

Lower pair of florets sterile or both male or the lower male and the other sterile; bisexual floret not awned; lodicules absent Anthoxanthum P- 75 2. Lowest floret male, the other bisexual, both awned Arrhenatherum P- 77 1. Lowest florets bisexual, the succeeding similar or reduced

GRASSES OF H A W A I I

74 3.

Rachilla disarticulating below the glumes

3. 4.

.

Sphenopholis P- 87

Rachilla disarticulating above the glumes Spikelets 1- to several-flowered; rachilla produced 5. Ovary hairy above or all over; awns mostly twisted at the base; spikelets nodding, over 8 mm long . . Avena P- 77 5. Ovary glabrous; awns not twisted at the base; spikelets less than 8 mm long 6. Lemmas finely erose-dentate or lobed, awned dorsally; spikelets 2- rarely 3-flowered Deschampsia p. 80 6.

Lemmas cleft or 2-toothed or 2-lobed with lobes sometimes prolonged into an awn; rachilla of spikelet pubescent or ciliate Trisetum p. 87 4. Spikelets strictly 2-flowered; rachilla not produced 7. Spikelets falling entire at maturity, strongly laterally compressed ; lower lemma awnless; upper lemma awned, usually male Holcus p. 84 7. Spikelets breaking up at maturity leaving the glumes persistent Air a P- 74 Aira L., Sp. PI. 63 (1753). Aspris Adans., Fam. PI. 2:496 (1763). SPIKELETS solitary, pedicelled, not falling entire; pedicel terete, glabrous, FLOWERS 2. GLUMES persistent, equal or subequal, ovate or elliptic, acute, 1-nerved, delicately membranous, LEMMAS 2, shorter than the glumes, lanceolate or ovate-lanceolate, rounded on the back, acute, awned from below the middle or awnless, the upper lemma sometimes acuminate, often 2-toothed or -lobed, faintly or obscurely 5-nerved, awned and the lower awnless; both lemmas subtending a palea and a bisexual flower; awn kneed, twisted below, capillary, shorter than to almost twice as long as the body of the lemma, PALEAS slightly shorter than their lemmas, narrow, 2-toothed, membranous or subhyaline. LODICULES 2, delicate, STAMENS 3. OVARY glabrous; styles free, short,

FESTUCOIDEAE -

AVENEAE

75

plumose above, laterally exserted near the base of the spikelet. GRAIN more or less adhering to its lemma and palea; embryo suborbicular, 1/5-1/6 as long as the grain. Small annuals; culm smooth, glabrous; leaf sheath shorter than the internode of the culm; ligule hyaline; blade almost filiform; inflorescence a lax panicle; primary branches of the inflorescence often in pairs on the main rachis. 1.

Aira carophyllea L., Sp. PI. 66 (1753). [Fig. 20] Aspris carophyllea (L.) Nash, in Britton and Brown, Illus. Fl. ed. 2, 1:214 (1913).

There has been much taxonomic and nomenclatural confusion among the genera Aira, Aspris, and Deschampsia. After the selection of Aira praecox L . as the lectotype of the genus Aira (Intern. Rules Bot. Nomencl. ed. 3, 139 [1935]), the generic name Aira is accepted for those species without prolongation of the rachilla and with the lemmas terminating in two slender teeth, and Aspris becomes a synonym for Aira. Species characterized by a prolonged rachilla and truncate lemmas are now placed in Deschampsia. See notes on Deschampsia for further information. Anthoxanthum L., Sp. PI. 28 (1753). solitary, pedicelled, not falling entire, FLOWERS 1-3. persistent, unequal, strongly or evidently keeled, with evident lateral veins, membranous or subhyaline; lower glume about 3/4 as long as the upper, lanceolate in profile, acute; upper glume as long as or longer than the lemmas, lanceolate or lanceolate-elliptic in profile, acute, sometimes shortly mucronate. L E M M A S 3, dissimilar; 2 lowest lemmas very similar, oblong-linear in profile, bifid or deeply 2-lobed, with an awn from the base or from the back or from the sinus, membranous, villous or pilose, each usually subtending a palea and one or both sometimes also subtending a male flower; awns usually unequal, that of the upper lemma as long as the body of the lemma, that of the lower lemmas much shorter, PALEAS (when present) linear, almost as long as their respective lemmas, 2-keeled, hyaline, U P P E R M O S T L E M M A 1/3-1/2 as long as the lower lemmas, ovate to oblong in profile, obtuse, awnless, delicately 7-1-nerved, 2-keeled, membranous, subtending a palea and a bisexual flower, PALEA as long as its lemma, linear, hyaline. L O D I C U L E S none, STAMENS 3 in the male flower, 2 in the bisexual flower. SPIKELETS

GLUMES

76

GRASSES OF H A W A I I

FESTUCOIDEAE - AVENEAE

77

OVARY ellipsoid; styles distinct, plumose above, exserted from the top of the spikelet. GRAIN ovoid; embryo 1/4 the length of the grain. Annuals or perennials, sometimes stoloniferous; culm ribbed, glabrous; leaves sometimes all basal and lanceolate; leaf sheath longer or shorter than the internode of the culm, ribbed, glabrous; ligule membranous; blade flat, narrow-linear, closely ribbed or ribs distinct, with or without a distinct midrib, rough or smooth, glabrous; inflorescence a cylindrical spikelike panicle or a more or less compact but evident panicle. 1.

Anthoxanthum odoratum L., Sp. PL 28 (1753). [Fig. 21]

Arrhenatherum Beauv., Ess. Agrost. 55 (1812). SPIKELETS solitary, pedicelled, laterally compressed, not falling entire, the rachilla very short, glabrous, disarticulating above the glumes but not between the lemmas, FLORETS 2, the lower male, the upper bisexual. GLUMES persistent, unequal, the lower much shorter than the upper, hyaline or scarious, acute or acuminate, rounded on the back, 5-7nerved. LEMMAS 2, the lower usually subtending a male, the upper a bisexual floret, slightly longer than the glumes, chartaceous or subherbaceous, acuminate, 5-7-nerved, the lower bearing a long twisted awn near the base, the upper usually awnless; awn geniculate, twisted below the knee; callus glabrous, PALEA shorter than lemma, narrow, 2-keeled, entire or minutely 2-toothed, the keels ciliate. LODICULES 2, lanceolate triangular, large, STAMENS 2. OVARY obovoid, poorly developed in the lower floret, densely villous toward the apex; styles free, the stigmas plumose, laterally exserted. GRAIN oblong, pubescent or villous at the summit. Tall slender perennials; leaf blade linear, elongated, flaccid; ligule hyaline; inflorescence a lax elongated symmetrical or secund panicle. 1.

Arrhenatherum elatius (L.) J.S. et C.B. Presl, Fl. Cech. 17 (1819); Mert et Koch, Deutsch. FL 1:546 (1823). [Fig. 22] Avena elatior L., Sp. PL 79 (1753). Arrhenatherum, avenaceum (Scop.) Beauv., Ess. Agrost. 152, 153 (1812).

Avena L., Sp. PL 79 (1753). SPIKELETS solitary, pedicelled, not falling entire, FLOWERS 1-2, rarely 3, bisexual, rarely with 1 - 2 additional male flowers, GLUMES persistent,

7»

GRASSES OF H A W A I I

I FIG. 2 1 . Anthoxanthum odoratum L . Plant, X 1/2; spikelet, sterile lemmas, and fertile floret, X 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FESTUCOIDEAE -

AVENEAE

79

FIG. 22. Arrhenatherum elatius (L.) J . S. et C. B. Presl. Plant, X 1/2; spikelet and upper floret, X 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

8o

GRASSES OF H A W A I I

equal or subequal, longer than the lemmas, lanceolate or lanceolatelinear, acute, sometimes acuminate, rounded on the back, 7-11-nerved, scarious, glabrous, LEMMAS 3-5, unequal and dissimilar, UPPERMOST LEMMA hyaline, empty or subtending a rudimentary male flower; next lemma (when ,4-5 lemmas present) lanceolate, almost convolute, obtuse, awnless, subtending a flower; normal lower lemmas lanceolate, rounded on the back, acuminate, subacute, awned from the middle of the back, stiffly membranous, each subtending a palea and a bisexual flower; awn slightly longer than to almost twice as long as the body of the lemma, kneed and twisted below, PALEAS almost as long as their respective lemmas, 2-toothed or bifid, 2-keeled, ciliate on the keels. LODICULES 2, rather large, hyaline, OVARY densely villous from the base; styles free, extremely short or lacking; stigmas laterally exserted. GRAIN oblong in outline, grooved in front, hairy, tightly embraced by the hardened lemma and palea; embryo small. Annuals; culm smooth, glabrous; leaf sheath ribbed, glabrous; ligule hyaline; blade linear, flat, strongly ribbed, scabrid; inflorescence a lax panicle, sometimes a subraceme. 1.

Avena barbata Pott ex Link in Schräder, Journ. für Bot. 2 : 3 1 5 ( I 799)-

2.

A. byzantina C. Koch in Linnaea 2 1 : 3 9 2 (1848).

3.

A. fatua L., Sp. PI. 80 (1753). [Fig. 23]

4.

A. sativa L., Sp. PI. 79 (1753).

Deschampsia Beauv., Ess. Agrost. 91 (1812). SPIKELETS solitary, pedicelled, not falling entire; pedicel long, capillary. FLOWERS 2, very rarely some spikelets with only one flower, rarely a third terminal male flower present, GLUMES persistent, as long as the lemmas, lanceolate, acute, keeled, shining, membranous, glabrous. LEMMAS 2, oblong-lanceolate, toothed at the apex, awned from the base, subhyaline or membranous, each usually subtending a bisexual flower, rarely a third lemma present which is empty or subtends a male flower; awn slightly longer than the body of the lemma, slender, twisted at the base, PALEA narrow, 2-nerved, hyaline, STAMENS 3. OVARY ellipsoid; styles free, plumose above, GRAIN oblong in outline, free within the palea. Perennials or annuals; culm smooth, glabrous; leaf sheath ribbed, glabrous; ligule hyaline; blade narrow-linear, with a prominent midrib

FIG. 23. Avena jatua L . (U.S.D.A. photograph.)

1 , Plant, x 1/2; 2, inflorescence, x 1/2; 3, spikelet, X 2.

82

G R A S S E S OF

HAWAII

and obscure lateral veins, scabrid, glabrous ; inflorescence a lax panicle ; primary branches of the inflorescence rather distant and whorled on the main axis. i.

Deschampsia australis Nees ex Steud., Syn. PI. Glum. 1:220 (1854), in Hillebr., Fl. Haw. Is. 520 (1888).

The Hawaiian species of Deschampsia have had an interesting but confusing history with regard to nomenclature. The author will attempt to outline the nomenclatural revisions that have been made with regard to D. australis and will let the reader draw his own conclusions. Hillebrand (1888) recognized 3 species of Deschampsia: (a) D. australis Nees ex Steud., Syn. PI. Glum. 1:220 (1854). Air a australis Nees (mpt. sub. Deschampsia) in Steud., Syn. PI. Glum. 1:220 (1854) non A. australis Raoul. (b) D. pollens Hillebr., Fl. Haw. Is. 520 (1888). D. pallens var. beta Hillebr., Fl. Haw. Is. 520 (1888). (c) D. nubigena Hillebr., Fl. Haw. Is. 521 (1888). Hitchcock (1922) examined the above named species in Hillebrand and concluded, 'The three species given by Hillebrand (D. pallens, D. australis, D. nubigena) seem to be all forms of A. nubigina,' choosing the species name nubigena, since the names australis and pallens were preoccupied under Aira. Thus the species became Air a nubigena (Hillebr.) Hitchc., Mem. Bernice P. Bishop Mus. 8:145 (1922). Skottsberg (1926), after studying his own material and that of Hillebrand, concluded that it would be best to restore Hillebrand's species which then became: (a) Aira pallida (Hillebr.) Skottsb., Acta Hort. Gothob. 2:204 (1926). A. pallida var. tenuissima Skottsb., Acta Hort. Gothob. 2:205 (1926). Deschampsia pallens var. beta Hillebr., Fl. Haw. Is. 520 (1888). A. hawaiiensis (Hillebr.) Skottsb., Acta Hort. Gothob. 2:205 (1926). (b) A. hawaiiensis f. depauperata Skottsb., Acta Hort. Gothob. 2:207 (1926).

FESTUCOIDEAE -

AVENEAE

83

A. hawaiiensis f. haleakalensis Skottsb., Acta Hort. Gothob. 2:207 (1926). (c) A. nubigena (Hillebr.) Skottsb., Acta Hort. Gothob. 2:208 (1926). In 1944, Skottsberg published further descriptions of forms of Deschampsia australis and proposed the following tentative arrangements: (a) D. australis Nees ms. ex Hillebr., Fl. Haw. Is. 520 (1888); sensu strictu {Aira australis Nees) ms. sub. (Deschampsia ex Steud.), Syn. PI. Glum. 1:220 (1854). Aira nubigena Hitchc., Mem. Bernice P. Bishop Mus. 8:145 (1922) pro parte. A. hawaiiensis Skottsb., Acta Hort. Gothob. 2:207 (1926). (b) D. australis f. haleakalensis Skottsb., Acta Hort. Gothob. 15:279 (1944). (c) D. australis subsp. nubigena (Hillebr.) Skottsb., Acta Hort. Gothob. 15:279 (1944) sensu strictu D. nubigena Hillebr., Fl. Haw. Is. 520(1888); A. nubigena Skottsb., Acta Hort. Gothob. 2:208 (1926); A. nubigena (Hillebr.) Hitchc., Mem. Bernice P. Bishop Mus. 8 (3):i45 (1922) pro parte. (d) D. australis subsp. nubigena var. gracilis Skottsb., Acta Hort. Gothob. 15:280 (1944), as a new variety. Rock (Indig. Trees Haw., 87 [1913]) mentions a variety, pumila, from Waialeale, undescribed, which is placed in the variety gracilis by Skottsberg. Under the variety gracilis, Skottsberg describes 3 forms: f. major, f. media, and f. minor. (e) Deschampsia australis subsp. nubigena var. tenuissima Skottsb., Acta Hort. Gothob. 2:205 (1926), D. pallens Hillebr. var. ß, Fl. Haw. Is. 520 (1888). Skottsberg's (1944) last comment on var. tenuissima was: 'Possibly better described as a form of var. gracilis.' St. John (Bull. Torrey Bot. Club 72:23 [1945]) made the following new combinations, apparently without being aware of Skottsberg's (1944) suggestions: (a) 'Deschampsia hawaiiensis (Skottsb.) St. John, comb. nov. Aira hawaiiensis (Hillebrand herb.) Skottsb., Medd. Göteborgs Bot. Trädgard 2:205-207 f. ib, c. (1926), D. australis Hillebrand, Fl. Haw. Is. 520-521. 1888; not Raoul, 1846.'

84

GRASSES OF HAWAII

(b) 'Deschampsia hawaiiensis (Skottsb.) St. John, f. depauperata (Skottsb.) St. John, comb. nov. Air a hawaiiensis (Hillebrand herb.) Skottsb. f. (depauperata) Skottsb., Medd. Göteborgs Bot. Trädgard 2:207-208.

(c)

1926.'

'Deschampsia hawaiiensis (Skottsb.) St. John, comb. nov. (Air a hawaiiensis) (Hillebrand herb.) Skottsb. f. haleakalensis Skottsb. Medd. Göteborgs Bot. Trädgard 2:207. 1926.'

2.

D. caespitosa (L.) Beauv., Ess. Agrost. 9 1 , 1 4 9 , 1 6 0 ( 1 8 1 2 ) . [Fig. 2 4 ]

3.

D. elongata (Hook.)

4.

D. flexuosa (L.) Trin., Bull. Acad. Sei. Petersb. Aira flexuosa L., Sp. PI. 65 (1753).

Holcus L., Sp. PI.

Munro ex Benth., PI. Hartw.

342 (1857). 1:66

(1836).

1047 (1753).

SPIKELETS solitary, pedicelled, falling entire; pedicel terete, pilose. FLOWERS 1-2. GLUMES unequal; lower glume lanceolate, keeled, ciliate on the keels and margins, 1-nerved, membranous, awned, pubescent; awn, when present, 1/2 as long as the body of the glume; upper glume larger than the lower, elliptic-lanceolate or ovate, keeled, with 2 prominent lateral nerves, ciliate on the keel and margins, pubescent, awned or shortly mucronate; awn (when present) as long as the body of the glume, LEMMAS 2, shorter than the glumes, unequal; lower lemma ovate, convex on the back, obtuse or acute, obscurely 3-5-nerved, awnless, subtending a palea and a bisexual flower, PALEA smaller than the lemma, narrow-lanceolate, hyaline or membranous, LODICULES 2, delicate. STAMENS 3. OVARY glabrous; styles distinct, plumose above, laterally exserted. GRAIN enclosed by the lemma and palea, often adhering to the palea; embryo small, UPPER LEMMA shorter than to about as long as the lower, lanceolate, awned, glabrous, subtending a palea and a bisexual or male flower or sometimes subtending only a palea; awn as long as to twice as long as the body of the lemma, PALEA smaller than its lemma. Annuals; culm ribbed, glabrous or pilose above; leaf sheath about as long as the internode of the culm, ribbed, pubescent or pilose; ligule membranous; blade flat, linear, without a distinct midrib, smooth, pubescent or pilose; inflorescence a contracted panicle, sometimes almost spikelike, oblong or interrupted; primary branches of the inflorescense sometimes fascicled on the main rachis. 1.

Holcus lanatus L., Sp. PI. 1 0 4 8 ( 1 7 5 3 ) . [Fig. 2 5 ] Ginannia lanata (L.) F.T. Hubb., Rhodora 18:234 ( I 9 I Q -

FESTUCOIDEAE - AVENEAE

85

FIG. 24. Deschampsia caespitosa (L.) Beauv. Plant, x 1/2; glumes and floret, x 10. (From Hitchcock and Chase, Manual of Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

86

GRASSES

OF

HAWAII

FIG. 25. Holcus lanatus L . Plant, x 1/2; spikelet, florets, and mature fertile floret, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FESTUCOIDEAE -

AVENEAE

87

Sphenopholis Scribn., Rhodora 8:142 (1906). SPIKELETS 2- or 3-flowered, the pedicel disarticulating below the glumes, the rachilla produced beyond the upper floret as a slender bristle. GLUMES unlike in shape, the first narrow, usually acute, i-nerved, the second broadly obovate, 3-5-nerved, the nerves sometimes obscure, mostly somewhat coriaceous, the margins scarious. L E M M A S firm, scarcely nerved, awnless or rarely with an awn from just below the apex, the first a little shorter or a little longer than the second glume, PALEA hyaline, exposed.

Slender perennials (rarely annual) with usually flat blades and narrow shining panicles. 1.

Sphenopholis obtusata (Michx.) [Fig. 26]

Scribn., Rhodora 8:144 (1906).

Aira obtusata Michx., Fl. Bor. Amer. 1:62 (1803). Eatonia obtusata A . Gray, Man. Bot. ed. 2, 558 (1856). Trisetum Pers., Syn. PI. 1:97 (1805). solitary, pedicelled, not falling entire, FLOWERS 2 to several. persistent, equal or more or less unequal, acute, keeled, hyaline; lower glume 1-3-nerved; upper glume 3-nerved. L E M M A S 2-6, equalling or exceeding the glumes, acutely 2-toothed, with broad margins, faintly or obscurely 3-5-nerved, membranous, hyaline at the tip, awned and sometimes with fine short bristles from the teeth, subtending bisexual or male flowers or the upper without flowers; awn from the back near the tip, fine, straight or kneed and twisted below, PALEA shorter than the lemmas, 2-toothed, hyaline, STAMENS 3. O V A R Y glabrous; styles free, very short, plumose above, laterally exserted. G R A I N oblong in outline, embraced by the usually slightly hardened back of the lemma; embryo small. SPIKELETS

GLUMES

Perennials, rarely annuals; leaf blades usually flaccid; ligule hyaline, sometimes minutely ciliate and hairy from the back; inflorescence usually a contracted often spikelike panicle, more rarely opening and lax. 1.

Trisetum flavescens (L.J Beauv., Ess. Agrost. 88, 153 (1812). Avena flavescens L., Sp. PI. 80 (1753).

2.

T. glomeratum (Kunth) Trin. in Steud., Syn. PI. G l u m . 1:229 (1854). [Fig. 27] Koeleria glomerata Kunth, Rev. Gram. 2:611, pi. 219 (1834).

3.

T. inaequale Whitney, Occas. Papers Bernice P. Bishop Mus. 1 3 1 6 - 1 3 1 7 (1937).

88

GRASSES

OF

HAWAII

FIG. 26. Sphenopholis obtusata (Michx.) Scribn. A, Plant, x 1/2; B, spikelet, x 10; C, glumes, X 1 0 ; D, lower floret, x 10; E, upper floret, x 10. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

FESTUCOIDEAE -

AVENEAE

89

FIG. 27. Trisetum glomeratum (Kunth) Trin. (From O. Degener, Flora Hawaiiensis.)

90

GRASSES

BRACHYPODIEAE Harz,

OF

HAWAII

Linnaea 43:15 (1880).

Spikelets all alike, very shortly pedicelled, several- to many-flowered, terete; upper florets often imperfect; rachilla disarticulating above the glumes and between the florets. Glumes unequal persistent, the lower 1-5-nerved, the upper 3-7-nerved; lemmas 7-9-nerved, lanceolate, acuminate, awned from the tip, rounded on the back, scarious on the margins; palea as long as the lemmas, hyaline, 2-nerved and 2-keeled, pectinately ciliate on the keels; awn terminal, straight, scabrid. Ovary elongate, with a terminal villous or hairy appendage; styles 2, borne laterally below the appendage; stigmas plumose. Lodicules 2, ciliate. Stamens 3. Grains hairy at the top, tightly enclosed by the lemma and palea; hilum elongate, linear; embryo small; starch grains simple. Slender annual or perennial herbs. Leaf blades narrow, acuminate, with festucoid anatomy; silica bodies variable; microhairs 1-celled. Inflorescence terminal, spikelike, with the spikelets seated on very short pedicels on the opposite sides of and with the sides of the lemmas touching the tough axis. Chromosomes small; basic numbers 5, 7, 9. Genus:

Brachypodium

Brachypodium Beauv., Ess. Agrost. 100 (1812). SPIKELETS solitary, sessile or subsessile, not falling entire, FLOWERS 8 - 1 7 . GLUMES persistent, equal or subequal, lanceolate, or oblonglanceolate, acute, 3-7-nerved, stiffly membranous, glabrous, L E M M A S 5 - 1 7 , exserted from the glumes, similar, more or less equal or decreasing in size upwards, lanceolate, 7-nerved, awned, ciliate, scabrid, membranous, subtending a palea and a bisexual flower; the few upper glumes sometimes reduced to a little more than awns, PALEAS thinner in texture than their lemmas, sometimes equalling them, oblong or oblong-linear, almost truncate at the apex, 2-keeled and ciliate on the keels, L O D I C U L E S 2, lanceolate, usually ciliolate. S T A M E N S 3, rarely 2. O V A R Y with a villous appendage at the top; styles laterally exserted. G R A I N linear or linearoblong in outline, more or less adhering to the palea; embryo small. Annuals or perennials; culm ribbed, glabrous; leaf sheath ribbed, glabrous or pilose; ligule membranous; blade flat, linear, with evident veins and a more or less distinct midrib, ciliate, glabrous or pilose.

1.

Brachypodium distachyon (L.) Beauv., Ess. Agrost. 101 (1812).

F E S T U C O I D E A E - BROMEAE

BROMEAE Dumort., Obs. Gram. Belg. 82:115 (1823)

91 as

Bromaceae.

Spikelets all alike, several- to many-flowered, terete or laterally compressed, bisexual; upper florets often imperfect; rachilla disarticulating above the glumes and between the florets. Glumes unequal, persistent, the lower 1-3-nerved, the upper usually 3-5(7)-nerved, glabrous or hairy, often scarious or shining on the margins; lemmas rounded or keeled on the dorsal surface, 5-9-nerved, 2-cleft or lobed, awned from between the teeth or lobes, awn straight or recurved, scabrid, sometimes twisted at the base; lobes rarely aristate; paleas usually shorter than the lemmas, 2-nerved, 2-keeled; keels ciliate or scabrid. Ovary oblong, with a terminal appendage which is hairy or villous; styles 2, inserted laterally below the appendage, short; stigmas plumose. Lodicules 2, obcuneate or lanceolate. Stamens 3 or 2. Grain linear-oblong, usually concave-convex in section, adherent to the palea; hilum linear, filiform, the length of the grain; embryo small, basal; starch grains simple. Annual or perennial grasses. Leaf blades usually narrow, glabrous or hairy, with festucoid anatomy; leaf sheaths tubular; microhairs 1 -celled; silica bodies oblong; ligules membranous. Spikelets pedicelled, arranged in dense or loose panicles. Chromosomes large; basic number 7. Genus:

Bromus

Bromus L., Sp. PI. 76 (1753). solitary, pedicelled, not falling entire, FLOWERS 2-1 I. persistent, equal or subequal, similar, lanceolate, more rarely ovate or linear-lanceolate, acute or subacute, sometimes acuminate, keeled, scabrid on the keels, stiffly membranous, sometimes villous; lower 1-7-nerved; upper 3-9-nerved. LEMMAS 2-12, usually exserted from the glumes and of the same texture, similar, decreasing in size upwards, lanceolate, 2-toothed or 2-lobed, awned or mucronate from the sinus, 5-13-nerved, keeled, scabrid on the keel and main nerves, glabrescent or sometimes villous, subtending a palea and a bisexual flower, or the uppermost lemma or few upper lemmas empty, more rarely with rudimentary flowers, PALEA subequalling their lemmas, thinner in texture, linear or lanceolate, entire or bifid, 2-keeled, ciliate on the keels, sometimes pilose, LODICULES 2, oblong or lanceolate, entire or lobed. SPIKELETS

GLUMES

92

GRASSES OF

HAWAII

3. O V A R Y obovoid, often with a large villous 3- or 2-lobed terminal appendage, plumose above, laterally exserted or in the cleistogamic species permanently enclosed, G R A I N linear to linear-oblong in outline, usually adhering to the lemma and palea or at least to the latter; embryo small. Annuals or perennials; culms smooth, striate, or ribbed, glabrous; leaf sheath as long as or shorter than the internode of the culm, ribbed, somewhat scabrid, pubescent or villous, with distinct veins and a prominent midrib, scabrid, pilose, villous, or glabrous; inflorescence a lax or contracted panicle, more rarely a raceme or subraceme; primary branches in pairs or threes or clustered on the main axis, glabrous or pilose, sometimes scabrid; main axis scabrid. STAMENS

1.

Bromus breviaristatus Buckl.,

Proc. Acad. Nat. Sci. Phila.

2.

B. carinatus Hook, et Am., Bot. Beechey's Voy. 403 (1840).

3.

B. commutatus Schrad., Fl. Germ. 353 (1806).

4.

B. diandrus Roth, Bot. Abh. 44 (1787). B. rigidus var. gussonei (Pari.) Coss. et Dur., Expl. Sci. Alger. 2:159 (1855).

1862:98 (1862).

B. maximus var. gussonei (Pari.) Pari., Fl. Ital. 1:407 (1848). B. villosus var. gussonei (Pari.) Aschers. et Graebn., Syn. Mitteleur. Fl. 2:595 (1901). 5.

B. erectus Huds., Fl. Angl. 39 (1762).

6.

B. hordeaceus L., Sp. PI. 77 (1753).

7.

B. inermis Leyss., Fl. Hal. 16 (1761).

8.

B. marginatus Nees in Steud., Syn. PI. Glum. 1:322 (1854).

9.

B. molliformis Lloyd, Fl. Loire-Inf. 315 (1844).

10. B. mollis L., Sp. PI. ed. 2, 112 (1762). 11. B. polyanthus Scribn. in Shear, U. S. Dept. Agr. Div. Agrost. Bull. 23,56(1900). 12. B. racemosus L., Sp. PI. ed. 2, 1:114 13. B. rigidus Roth, Mag. Bot. Roem. and Ust. 10:21 (1790). [Fig. 28] B. maximus Desf., Fl. Atlant. 1:95 (1798). B. villosus Forsk., Fl. Aegypt.-Arab. 23 (1775). 14. B. rubens L., Cent. PI. 1:5 (1755); Amoen. Acad. 4:265 (1759).

FESTUCOIDEAE - BROMEAE

93

FIG. 28. Bromus rigidus Roth. A, Plant, X 1/2; B, spikelet, x 10; C, glumes, X 2. (From an original drawing in the files of the Dept. of Agronomy and Soil Scier ce, Univ. of Hawaii.)

GRASSES OF

94

HAWAII

15. B. squarrosus L., Sp. PI. 76 (1753). 16. B. squarrosus var. villosus (Gmel.)

Reich., Agrost. Germ. 32

(1834)17. B. sterilis L., Sp. PI. 77 (1753). 18. B. tectorum L., Sp. PI. 77 (1753). 19. B. unioloides (Willd.)

H.B.K.,

Nov. Gen. et Sp. 1:151 (1815).

B. catharticus Vahl, Sym. Bot. 2:22 (1791). Bor (i960) adds 'nomen dubium,' and refers the reader to C . E. Hubbard, D e Flora Lusitana Commentarii fasc. 9:7 (1956) for reasons to reject the epithet catharticus and why it should be called B. unioloides H . B . K . 20. B. vulgaris (Hook.) 23. 43 (^oo)-

Shear, U . S. Dept. Agr. Div. Agrost. Bull.

FESTUCOIDEAE - FESTUCEAE

95

FESTUCEAE Dumort., Obs. Gram. Belg. 82, 85 (1823). Spikelets all alike, bisexual, rarely dioecious, or dissimilar and then the latter consisting of empty glumes and lemmas, more or less laterally compressed, 2- to many-flowered, with the uppermost floret or florets reduced; rachilla disarticulating above the upper glume and between the florets, rarely below the glumes and then the spikelets falling entire. Glumes usually persistent, awned or awnless, the lower more or less equal to or shorter than the upper, rarely absent altogether, rarely both minute, the lower 1-nerved, the upper 3-nerved. Lemmas herbaceous, becoming firmer, 5-13-nerved, awned or awnless; awn straight or curved; paleas 2-keeled. Lodicules 2, rarely o. Stamens 3, rarely 1. Grain usually tightly enclosed by the lemma and palea; hilum punctiform and basal, or linear and then as long as the grain; starch grains compound. Annual or perennial grasses. Leaf blades setaceous, filiform or linear, with festucoid anatomy; silica bodies oblong or circular; 2-celled microhairs absent; ligule membranous. Spikelets pedicelled in loose or contracted panicles, rarely in racemes or sessile in panicled or solitary spikes. Chromosomes large; basic number 7 (mostly) or 5. Genera:

Beckmannia Briza Cynosurus

Dactylis Festuca Lamarckia

Lolium Poa Vulpia

KEY T O T H E G E N E R A O F F E S T U C E A E

i.

Spikelets of two kinds, fertile and sterile, the latter surrounding the former, or collected at the base of a dense panicle 2.

Fertile spikelet 2-3-flowered; sterile spikelets consisting of numerous awned scales Cynosurus P- 97

2.

Fertile spikelets 1 -flowered; sterile spikelets consisting of obtuse glumes Lamarckia p. 101 Spikelets all of one kind

3.

Spikelets disarticulating above the glumes, not subcircular in outline, more than 1-flowered

1.

96

GRASSES OF HAWAII

4.

Inflorescence a spike or occasionally the spikes panicled Lolium p. 103 Inflorescence a true panicle

4. 5.

Lemmas firmly keeled on the back 6.

Spikelets broadly ovate, pendulous; lemmas inflated, awnless, cordate at the base, closely imbricate, boatshaped, many-nerved, horizontally spreading Briza P- 97 Spikelets and lemmas not as above

6. 7.

Spikelets borne in dense one-sided clusters on the branches of a panicle, lemmas tipped with a rather stout awn up to 1.5 mm long; tufted perennials Dactylis P- 99 7. Spikelets borne in loose or contracted panicles; lemmas not awned Poa P- 105 Lemmas rounded on the back

5. 8.

Florets with 1 stamen; panicle rather dense, erect, of long-awned spikelets Vulpia p. 109

8.

3.

Florets with 3 stamens; lemmas awned or awnless, chartaceous to cartilagenous; perennials . Festuca P- 99 Spikelets 1 -flowered, disarticulating below the glumes, subcircular in outline, sessile, imbricate, secund in 2 rows along a continuous rachis; the spikes panicled Beckmannia p. 96

Beckmannia Host, Gram. Austr. 3:5

(1805).

SPIKELETS 1- or 2-flowered, laterally compressed, subcircular, nearly sessile and closely imbricate, in 2 rows along one side of a slender continuous rachis, disarticulating below the glumes, GLUMES falling entire; nerves equal, inflated, obovate, 3-nerved, rounded above but the apex apiculate; LEMMA 5-nerved, acuminate, about as long as the glumes, PALEA nearly as long as the lemma.

FESTUCOIDEAE -

FESTUCEAE

97

Erect rather stout annuals with flat blades with numerous short appressed or ascending spikes in a narrow more or less interrupted panicle. i.

Beckmannia syzigachne (Steud.) Fern., Rhodora 30:27 (1928). [Fig. 29] B. erucaeformis var. uniflora Scribn. ex A. Gray, Man. Bot. ed. 6, 628 (1890).

Briza L., Sp. PI. 70 (1753). solitary, pedicelled, not falling entire; pedicels capillary. 4-17. GLUMES persistent, sometimes almost at right angles to the rachilla, subequal, broadly ovate, very concave, 3- to 7-nerved, membranous, glabrous, LEMMAS 4-20, exserted from the glumes, broadly ovate, very concave, subacute, clasping the rachilla at the base, 7-9-nerved, rarely 5-nerved, more or less papery, finely pilose, each subtending a palea and a bisexual flower or the upper few empty, PALEAS suborbicular, 2-keeled, finely ciliate on the keels and with the keels often winged, subhyaline. LODICULES 2, obliquely ovate, STAMENS 3. OVARY glabrous; styles short, distinct, loosely plumose above, laterally exserted. GRAIN tightly embraced by the hardened back of the lemma and palea, usually adhering to the palea; embryo small. Annuals; culm striate, glabrous; leaf sheath ribbed, shorter than the internode of the culm, glabrous; ligule hyaline; blade flat, without a very distinct midrib; inflorescence a raceme or panicle. SPIKELET

FLOWERS

1.

Briza maxima L., Sp. PI. 70 (1753).

2.

B. minor L., Sp. PI. 70 (1753).

Cynosurus L., Sp. PI. 72 (1753). SPIKELETS sterile and fertile, in fascicles consisting of one fertile spikelet surrounded by the sterile spikelets. FERTILE SPIKELET sessile or subsessile. FLOWERS 1-6. GLUMES subequalling the lemmas, lanceolate, longacuminate, acute, 1-nerved, almost hyaline, scabrid on the keel, LEMMAS 2-7, similar, decreasing in size upwards, oblong or lanceolate, mucronate or awned from close below the tip, 5-nerved, membranous, each subtending a palea and a bisexual flower or sometimes the upper or uppermost lemma without a flower; awn about t.vice as long as the body of the lemma, scabrid. PALEAS subequalling the lemmas, and similar in structure, lanceolate, acuminate, acute, 2-toothed, 2-keeled.

98

GRASSES

OF

HAWAII

FIG. 29. Beckmannia syzigachne (Steud.) Fern, (a) Panicle, showing the ascending spikes, X 2/3; (b) habit, X 1/8; (c) floret, X 12; (d) spikelet, laterally compressed, X 8; (e) grain, x 12; ( f ) leaf sheath and ligule, x 4. (From H. L . Mason, A Flora of the Marshes of California.)

FESTUCOIDEAE - FESTUCEAE

99

2. STAMENS 3. OVARY glabrous; styles distinct, loosely plumose above, laterally exserted. GRAIN oblong in outline, more or less adhering to the lemma and palea; embryo small, STERILE SPIKELETS consisting of a number of linear-lanceolate long-acuminate empty lemmas on an elongated rachis; internodes of rachis very evident. Annuals; culm ribbed, glabrous; leaf sheath ribbed, shorter than the internode of the culm, glabrous; ligule hyaline; blade flat, conspicuously ribbed, scabrid, glabrous; inflorescence a more or less ovate spikelike raceme; primary branches sometimes all turned to one side giving the inflorescence a unilateral appearance. LODICULES

1.

Cynosurus cristatus L., Sp. PI. 72 (1753).

Dactylis L., Sp. PI. 71 (1753). SPIKELETS in dense secund shortly pedicelled fascicles, not falling entire, FLOWERS 3-7. GLUMES persistent, subequal or the lower shorter, keeled, mucronate, 1-3-nerved, rigid, membranous, LEMMAS 3-7, exceeding the glumes, 5-nerved, keeled, with the keel ciliolate and produced into a mucro or short awn, subtending a palea and a bisexual flower; uppermost lemma sometimes sterile, PALEAS slightly shorter than the lemmas, 2-keeled. STAMENS 3. OVARY glabrous; styles distinct, plumose above, GRAIN enclosed by the lemma and palea. A tall perennial; leaf blade flat; ligule scarious; inflorescence a crowded panicle or false spike.

1.

Dactylis glomerata L., Sp. PI. 71 (1753). [Fig. 30]

Festuca L., Sp. PI. 73 (1753). solitary, pedicelled, not falling entire, FLOWERS 2-9. persistent, unequal or subequal, lanceolate or ovate-lanceolate, acute or obtuse, 1-nerved or the upper 3-5-nerved, often scabrid on the keel, membranous, glabrous or pubescent, LEMMAS 2-10, exserted from the glumes, of the same texture as the glumes, similar, decreasing in size upwards, lanceolate, sometimes awned, 5-7-nerved, often scabrid on the keel, usually glabrous, subtending a palea and a bisexual flower or the uppermost usually subtending a rudimentary palea and sometimes a rudimentary flower; awn, when present, 1/4-1/2 as long as the body of the lemma, PALEAS subequalling the lemmas, lanceolate, 2-keeled, almost hyaline, LODICULES 2, unequally 2-lobed, hyaline. STAMENS 3. OVARY glabrous or the top minutely hairy; styles distinct, very short, terminal or subterminal, plumose, laterally exserted. GRAIN SPIKELETS

GLUMES

FIG. 30. Dactylis glomerata L . Plant, X 1/2; spikelet and floret, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FESTUCOIDEAE -

FESTUCEAE

IOI

oblong in outline, dorsally convex, grooved or concave in front, free or more or less adhering to the palea or to the lemma and palea, tightly enclosed by the slightly hardened lemma and palea. Annuals or perennials; culm smooth or striate, glabrous; leaf sheath shorter than the internode of the culm, glabrous; ligule scarious; blade linear, usually prominently ribbed, glabrous, sometimes scabrous. 1. Festuca arundinacea Schreb., Spic. Fl. Lips. 57 (1771). 2. F. drymeja Mert. et Koch in Roehl, Deutschl. Fl. ed. 3, 12:670 (1823). 3. F. elatior L„ Sp. PI. 75 (1753). 4. F.hawaiiensis//iVc/ic.,Mem. BerniceP. BishopMus.8:ii5(1922). [Fig- 3 1 ] 5. F. idahoensis Elmer., Bot. Gaz. 36:53 (1903). 6. F. kashmiriana Stapf in Hook., f., Fl. Brit. India 7:351 (1896). F. duriuscula Duthie, Grasses N.W. India 41 (1883), non L . (1753). 7. F. ovina L., Sp. PI. 73 (1753). 8. F. pratensis Huds., Fl. Angl. 37 (1762). 9. F. rubra L., Sp. PI. 74 (1753). F. dumetorum L., Sp. PI. ed. 2, 109 (1762). 10. F. tenuifolia Sibth., Fl. Oxon. 44 (1794). 11. F. viridula Vasey, U. S. Dept. Agr. Div. Bot. Bull. 13(2), pi. 93 (I893)Lamarckia Moench, Meth. PL 201 (1794). SPIKELETS in groups of 4, with i spikelet fertile and surrounded by 3 sterile spikelets. FERTILE SPIKELET pedicelled, falling entire, FLOWERS 1-2. GLUMES subequal, long-acuminate, acute, keeled, almost hyaline. LEMMAS 2, sometimes 1, lanceolate or ovate-oblong, bifid, awned from the middle or below the tip, membranous, each subtending a palea and a bisexual flower or the upper lemma empty or subtending a male flower; awn 3 times as long as the body of the lemma, PALEA subequalling the lemma and of the same texture, lanceolate, subacuminate, 2-toothed, 2-keeled. LODICULES 2, minute, STAMENS 3. STYLES distinct, barbellate. GRAIN elliptic-oblong in outline, enclosed in the unaltered lemma and palea. STERILE SPIKELET pedicelled. GLUMES lanceolate, acute, membranous. LEMMAS 12-13, obovate, keeled, membranous, empty, INTERNODES OF RACHILLA evident between the lemmas.

102

GRASSES OF H A W A I I

FIG. 31. Festuca hawaiiensis Hitchc. (From A. S. Hitchcock, The Grasses of Hawaii.)

FESTUCOIDEAE -

FESTUCEAE

IO3

A low-growing tufted annual; culm striate; leaf sheath longer than the internode of the culm, ribbed, glabrous; ligule hyaline; blade flat, finely ribbed, with a distinct midrib; inflorescence a raceme (usually unilateral) composed of the groups of spikelets. 1.

Lamarckia aurea (L.) Moench, Meth. PI. 201 (1794).

Lolium L., Sp. PI. 83 (1753). SPIKELETS solitary, sessile, not falling entire, FLOWERS 4-1 I . GLUMES of terminal spikelets equal and similar; lower glume absent in the lateral spikelets, very rarely present in some inflorescences and then very much smaller than the upper glume; upper glume persistent, facing the rachis, lanceolate or linear-lanceolate, obtuse or subacute, 7-9-nerved, glabrous, LEMMAS 6-12, usually not exserted from the glumes, sometimes decreasing in size upwards, lanceolate or elliptic, rounded on the back, awnless or awned or mucronate, 5-nerved, coriaceous, glabrous, subtending a palea and a bisexual flower, sometimes the uppermost or the two upper lemmas much reduced and empty; awn less than half as long as the body of the lemma, PALEAS equalling or subequalling the lemmas, somewhat thinner in texture, lanceolate, lanceolate-elliptic or lanceolate-oblong, 2-keeled, ciliate on the keels or the keels more or less crested, LODICULES 2, lanceolate, with a lateral tooth, STAMENS 3. OVARY truncate, glabrous; styles distinct, plumose above, laterally exserted. GRAIN elliptic-oblong to linear-oblong in outline, tightly enclosed by the lemma and palea and adhering to both; embryo short. Annuals or perennials; culm striate or ribbed, glabrous; leaf sheath longer than the internode of the culm, ribbed, glabrous; ligule membranous; blade flat, linear, closely ribbed, often scabrid; inflorescence a simple spike; the central axis with concavities for the reception of the spikelets, scabrid.

1.

2. 3. 4. 5.

Lolium multiflorum Lamk., Fl. Franc. 3:621 (1778). [Fig. 32] L. italicum A. Br., Flora 17:243 (1834). L. westernwoldicum Breakw., Grasses and Fodder PI. N.S.W. 171 (!923)L. perenne L., Sp. PI. 83 (1753). L. rigidum Gaud., Agrost. Helv. 1:334 (1811). L. subulatum Vis., Fl. Dalm. 1:90 (1842). L. temulentum L., Sp. PI. 83 (1753).

. (U.S.D.A. photograph.)

FESTUCOIDEAE -

FESTUCEAE

Poa L., Sp. PI. 67 (1753). SPIKELETS solitary, pedicelled, not falling entire; pedicel swollen at the apex, FLOWERS 2-6. GLUMES unequal or subequal, similar, ovate or oblong, 1-3-nerved, keeled, membranous, glabrous, LEMMAS 2-6, exserted from the glumes, similar, decreasing in size upwards, of the same texture as the glumes, lanceolate or oblong, obtuse or subacute, often scabrid on the keel, scantily pilose, more rarely glabrous, sometimes with long wool from the base, all subtending a palea and a bisexual flower or only the lemmas or lowermost lemma with a bisexual flower; the uppermost lemma or two upper lemmas subtending a female flower. PALEAS smaller than or subequalling the lemmas, 2-keeled, thinner in texture than the lemmas, glabrous or pilose, LODICULES 2, more or less 2-lobed. STAMENS 3. OVARY glabrous; styles distinct, plumose above, laterally exserted. GRAIN ovate, oblong, or linear in outline, free or adhering to the palea; embryo small. Annuals or perennials; culm striate, glabrous; leaf sheath longer than the internode of the culm, ribbed; ligule membranous or hyaline; blade long-linear, prominently or finely ribbed, smooth or sometimes scabrous on the margins, glabrous; inflorescence a lax or contracted panicle, primary branches solitary or usually in pairs or clustered on the central axis, pubescent.

1. Poa ampla Merr., Rhodora 4:145 (1902). 2. P. annua L., Sp. PI. 68 (1753). [Fig. 33] Aira pumila Pursh., Fl. Amer. Sept. 1:76 (1814). 3. P. arachnifera Torr. in Marcy, Expl. Red. Riv. 301 (1853). 4. P. bulbosa L., Sp. PI. 70 (1753). 5. P. compressa L., Sp. PI. 69 (1753). 6. P. flabellata (Lamk.) Rasp, in Saig. and Rasp., Ann. Sci. Obs. 2:86 (1829). 7. P. mannii Munro in Hillebr., Fl. Haw. Is. 526 (1888). [Fig. 34] 8. P. nemoralis L., Sp. PI. 69 (1753). 9. P. nervosa (Hook.) Vasey, U. S. Dept. Agr. Div. Bot. Bull. 13 (2), pi. 81 (1893). 10. P. pratensis L., Sp. Pi. 67 (1753). 11. P. sandvicensis (Reich.) Hitchc., Mem. Bernice P. Bishop Mus. 8:121 (1922). [Fig. 35]

io6

GRASSES OF H A W A I I

FESTUCOIDEAE - FESTUCEAE

FIG. 34. Poa mantiii Munro. (From A. S. Hitchcock, The Grasses of Hawaii.)

107

io8

GRASSES OF H A W A I I

FIG. 35. Poa sandvicensis (Reich.) Hitchc. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

FESTUCOIDEAE - FESTUCEAE

Festuca sandvicensis Reich., Sitzungsb. Akad. Wiss. Math. Naturw. (Wien) 76:726 (1878). P. longeradiata Hillebr., Fl. Haw. Is. 526 (1888). 12. P. scabrella (Thurb.) Benth. ex Vasey, Grasses U. S. 42 (1883). 13. P. siphonoglossa Hack, in Fedde, Repert. Sp. Nov. 11:24. (I912)[Fig- 36] 14. P. trivialis L., Sp. PI. 67 (1753). Vulpia Gmel., Fl. Bad. 1:8 (1805). SPIKELETS solitary, pedicelled, not falling entire; pedicel 3-angled, scabrid on the margins, FLOWERS 4-7. GLUMES persistent, similar, unequal; lower glume shorter than the upper, very minute or obsolete or subulate-lanceolate, 1-nerved; upper glume subulate to subulatelanceolate, 3-nerved. LEMMAS 5-8, much exserted from the glumes, lanceolate, passing into an awn, rounded on the back, faintly 5-nerved, glabrous or shortly pilose, subtending a palea and a bisexual flower; the upper lemma without flowers; awn slightly longer to twice as long as the body of the lemma, scabrid. PALEAS lanceolate to linear-lanceolate, entire or minutely 2-toothed, 2-keeled; hyaline, scabrid on the keels. LODICULES 2, unequally lobed, hyaline, STAMENS 1-3. OVARY glabrous or minutely hispid on the top; stigmas sessile, plumose, permanently enclosed between the lemma and the palea, or shortly exserted at the base, GRAIN linear in outline, more or less adhering to the palea, embryo small. Annuals; culm smooth, glabrous; leaf sheath ribbed, glabrous; ligule membranous; blade linear, very narrow, usually convolute or involute, at least when dry; inflorescence a spikelike panicle.

1.

Vulpia bromoides Dum., Obs. Gram. Belg. 101 (1823). Festuca bromoides L., Sp. PI. 75 (1753). F. dertonensis (All.) Aschers. et Graebn., Syn. Mitteleur. Fl. 2:558(1901).

2.

V. megalura (Nutt.) Rydb., Bull. Torrey Bot. Club 36:538 (1909). F. megalura Nutt., Journ. Acad. Phila. 2:1, 188 (1848).

3.

V. myuros (L.) Gmel., Fl. Bad. 1:8 (1806). F. myuros L., Sp. PI. 74 (1753).

4.

V. octoflora (Walt.) Rydb., Bull. Torrey Bot. Club 36:538 (1909). F. octoflora Walt., Fl. Carol. 81 (1788).

no

GRASSES

OF

HAWAII

FIG. 36. Poa siphonoglossa Hack. Fertile plant on left, sterile plant on right. (From A. S. Hitchcock, The Grasses of Hawaii.)

FESTUCOIDEAE -

GLYCERIEAE

III

GLYCERIEAE Endl., Fl. Poson. 1 1 7 (1830) as Glycerinae. Spikelets few- to many-flowered, subterete or slightly compressed; rachilla disarticulating above the glumes and between the florets. Glumes persistent, unequal, 1 -nerved, hyaline, usually scarious; lemmas obtuse, membranous, rounded on the back, 7-9-nerved, hyaline at the tip; nerves prominent; palea as long as the lemma or shorter, hyaline, 2-keeled, smooth or scaberulous on the keels. Lodicules 2, minute. Stamens 3; anthers large or small. Ovary glabrous; styles 2, short, distinct; stigmas plumose. Lodicules connate, fleshy, swollen. Grain long, slender; hilum linear, as long as the grain; embryo small; starch grains compound. Perennial or annual, usually aquatic herbs. Leaf blades usually narrow, linear, with festucoid anatomy and transverse nervules; silica bodies with wavy outlines; microhairs 1-celled; ligules membranous or scarious. Margins of the leaf sheaths fused. Spikelets arranged in effuse panicles or racemes. Chromosomes small; basic number 4 or 5. Genus:

Glyceria

Glyceria R. Br., Prodr. Fl. Nov. Holl. 179 (1810). SPIKELETS solitary, pedicelled, cylindrical or slightly compressed, not falling entire; rachilla disarticulating above the glumes and between the lemmas, FLORETS few to many (6-20), bisexual, GLUMES persistent, unequal, herbaceous or hyaline, oblong, obtuse or subacute, faintly nerved at the base, mostly 1-nerved, LEMMAS 6-20, broad, convex on the back, firm, exserted from the glumes, awnless, 3-9-nerved, the nerves parallel and not quite reaching the hyaline apex, the upper, lemmas often sterile; callus obscure, PALEA as long as or longer than the lemma, 2-keeled. LODICULES 2, fused, STAMENS 3. OVARY glabrous; styles distinct, rather long; stigmas plumose, laterally exserted. GRAIN surrounded by the hardened lemma and palea, oblong in outline; hilum basal and punctiform.

Usually tall aquatic or marsh perennials, with creeping and rooting bases or with creeping rhizomes; culms with closed or partly closed sheaths, flat blades; ligule hyaline, lacerated; inflorescence an open or contracted panicle.

112

G R A S S E S OF H A W A I I

1.

Glyceria fluitans (L.) R. Br., Prodr. Fl. Nov. Holl. 179 (1810). Festuca fluitans L., Sp. PI. 75 (1753). Panicularia fluitans (L.) Kuntze, Rev. Gen. PI. 1:782 (1891).

2.

G. grandis S. Wats ex A. Gray, Man. Bot. ed. 6, 667 (1890). [Fig- 37] Poa aquatica var. americana Torr., Fl. Northern and Middle Sect. U.S. 1:108 (1823).

FIG. 37. Glyceria grandis S. Wats, (a) Floret, showing palea, x 16; (b) spikelets, solitary on tips of branchlets, x 8; (c) habit, lower part showing the conspicuous joints of culm and the long, lax leaf blades, x 1/5; (d) habit, upper pait of culm showing panicle, x 1/5; (e) leaf sheath, blade, and luptured ligule, X 4; ( f ) floiet, showing lemma, the strong nerves papillose, x 16; (g) young upper leaf, the sheath and ligule enclosing culm, x 4. (From H. L. Mason, A Flora of the Marshes of California.)

ii4

GRASSES

ISACHNEAE Benth., parte.

OF

HAWAII

Journ. Linn. Soc. (Bot.) 19:30, 92 (1881) pro

Spikelets alike, bisexual, 1-, 2-, very rarely 3-flowered, small to very small, awnless; rachilla disarticulating above the glumes and usually between the florets. Glumes persistent, eventually deciduous, usually equal, membranous, 3-9-nerved, rarely nerveless, shorter than or rarely slightly longer than the spikelet; lemma membranous or chartaceous, 5-7-nerved; palea as long as the lemma or absent; upper lemma similar to the lower or often smaller, firmer or of the same texture, 5-7-nerved. Lodicules 2, cuneate, truncate. Stamens 2 or 3. Grain free between the lemma and palea, fusiform, ovoid or ellipsoid, plano-convex or elliptic in section; hilum small and basal, or linear and elongated; embryo up to 1/3 the length of the grain; starch grains simple. Annual or perennial grasses. Leaf blades usually broad, with festucoid anatomy; silica bodies dumbbell-shaped, cross-shaped or roughly rectangular with concave sides; microhairs simple or 2- or more-celled; ligules a row of hairs or absent. First foliage leaf of the seedling vertical. Inflorescence of panicles or spikelike panicles or spikelike racemes. Chromosomes small; basic number 10. Genus:

Isachne

Isachne R. Br., Prodr. Fl. Nov. Holl. 196 (1810). SPIKELETS small, ovoid to subglobose, continuous with the pedicel, articulate above the sterile lemmas, FLOWERS 2, bisexual, or rarely the upper female or the lower male, GLUMES membranous; about equal and as long as the fruits or at maturity exceeded by them, L E M M A S coriaceous and similar in form and texture to their paleas. Both FLORETS (or FRUITS) plano-convex, obtuse, equal in size or the upper shorter, the pair usually remaining attached by the minute rachilla joint between them, GRAIN enclosed by the lemma and palea.

Perennial or rarely annual grasses with simple or usually branching stems, flat strongly nerved blades, and paniculate inflorescence with slender branches. 1.

2. 3.

Isachne distichophylla Munro, Journ. Bot. Brit, and For. 7:178 (1869) nom. nud.; Munro ex Hillebr., Fl. Haw. Is. 504 (1888). [Fig- 38] I. distichophylla var. beta Hillebr., Fl. Haw. Is. 504 (1888). I. myosotis Nees, Journ. Bot. K e w Misc. 2:98 (1850). I. pallens Hillebr., Fl. Haw. Is. 504 (1888).

"5

FIG. 38. Isachne distichophylla Munro. Plant, x 1/3; leaf and sheath, x 2; part of panicle, X 4; floret, glumes and spikelet, x 10. (From O . Degener, Flora Hawaiiensis.)

n6

GRASSES OF H A W A I I

LEPTUREAE Holmb., Bot. Not. 1926:80 (1926). Spikelets 1-flowered, bisexual, sessile, solitary, alternate, appressed to or sunken in the hollows of the joints on opposite sides of the readily disarticulating rachis of slender cylindric spikes; rachilla disarticulating above the glumes and between the florets, produced beyond topmost floret and carrying a rudimentary floret. Lower glume absent; upper glume coriaceous, thick, tailed or not, eventually becoming coriaceous; lemma with its back to the rachis, hyaline or membranous, awnless, 3-nerved; palea of the same texture, 2-nerved. Lodicules 2. Stamens 3. Styles 2, thick, distinct; stigmas plumose. Grain oblong, dorsally compressed; embryo longer than 1/2 the length of the grain; hilum punctiform, basal; starch grains simple. Low annual or perennial herbs spreading widely by means of stolons; leaves narrow, with saddle-shaped or transversely dumbbell-shaped silica bodies; pyriform 2-celled microhairs sometimes present. Ligules very short, ciliate. Inflorescence a terminal spike. Chromosomes small; basic number 9. Genus:

Lepturus

Lepturus R. Br., Prodr. Fl. Nov. Holl. 207 (1810). SPIKELETS solitary, sessile, not falling entire, FLOWER I . GLUMES persistent, of terminal spikelets both developed, similar, equal; lower glume of lateral spikelets absent; upper glume longer than the lemma, facing the main axis, lanceolate, acute, 5-nerved, coriaceous, glabrous. LEMMA i, lanceolate, acute, 3-nerved, membranous, subtending a palea and a bisexual flower, PALEA equalling the lemma, lanceolate, 2-nerved, hyaline, LODICULES 2. STAMENS 3. OVARY truncate, glabrous; styles distinct, plumose above, laterally exserted. GRAIN narrow-oblong in outline, enclosed with the lemma and the palea between the glume and the axis of the inflorescence. Annuals or perennials; culm smooth, glabrous; leaf sheaths longer or shorter than the internode of the culm, glabrous; ligule membranous; blade narrow, linear, ribbed, without a distinct midrib, glabrous; inflorescence a simple cylindrical spike; main axis with concavities for the reception of the spikelets, very closely adpressed to the axis.

1.

Lepturus cinereus Burcham, Contr. U. S. Natl. Herb. 30:424 (1948).

FESTUCOIDEAE -

LEPTUREAE

117

L. repens var. cinereus (Burcham) Fosberg, Occas. Papers Bernice P. Bishop Mus. 21:292 (1955). 2.

L. repens (G. Forst.) R. Br., Prodr. Fl. Nov. Holl. 207 (1810). Rottboellia repens G. Forst., Fl. Ins. Austr. Prodr. 9 (1786).

II8

GRASSES

OF

HAWAII

MELICEAE Reichb., Consp. Reg. Veg. 53 (1828) as Melicaceae. Spikelets 1- to several-flowered, laterally compressed, the lower florets bisexual, the uppermost reduced and represented by a bundle of empty rudimentary lemmas; rachilla disarticulating above the glumes, with long internodes, crowned by rudimentary florets. Glumes equal or unequal, the lower 1-3-nerved, the upper 5-nerved, membranous; lemmas 5-9-nerved, longer than the glumes, compressed, hairy or glabrous, awnless; the lowest stipitate; palea hyaline, oblong, 2-keeled, ciliolate on the keels. Lodicules 2, fused laterally. Ovary glabrous; styles 2, free; stigmas plumose. Grain oblong, fusiform or terete; hilum linear, elongate. Starch grains compound. Erect perennial herbs. Leaf blades narrow, with festucoid anatomy; silica bodies simple; no microhairs. Margins of the leaf sheaths fused. Spikelets arranged in open or spiciform or racemiform panicles, jointed or not on the pedicels. Ligules membranous or absent. First foliage leaf of the seedling erect. Chromosomes large; basic number 9. Genus:

Melica

Melica L., Sp. PI. 66 (1753). SPIKELETS 2- to several-flowered (rarely with 1 perfect floret), the rachilla disarticulating above the glumes and between the fertile florets (in some species spikelets falling entire), prolonged beyond the perfect florets and bearing 2 or 3 gradually smaller empty lemmas, each enclosing the one above, GLUMES somewhat unequal, thin, often papery, scarious-margined, obtuse or acute, sometimes nearly as long as the lower floret, 3- to 5-nerved, the nerves usually prominent, L E M M A S convex, several nerved, membranous or rather firm, scarious margined, sometimes conspicuously so, awnless or sometimes awned from between the teeth of the bifid apex, the callus not bearded.

Rather tall perennials, the base of the culm often swollen into a corm, with enclosed sheaths, usually flat blades, narrow or sometimes open, usually simple panicles of relatively large spikelets. 1.

Melica californica Scribn., Proc. Acad. Nat. Sci. Phila. 1885:46 (1885).

FESTUCOIDEAE -

PHALARIDEAE

II9

PHALARIDEAE Kunth, Rev. Gram. 1 : 1 2 (1829). Spikelets all similar or sometimes the lower reduced, bisexual, strongly compressed, 3-flowered, with the two lower florets sterile and reduced to the lemmas or one or both absent; rachilla disarticulating above the glumes but not between the florets. Glumes persistent, equal and as long as the spikelet; lower two lemmas small, sometimes reduced to minute scales; fertile lemma becoming indurated, awnless, 5-nerved; palea as long as the lemma, 2-nerved, 2-keeIed. Lodicules 2, hyaline. Ovary glabrous; styles 2; stigmas 2, plumose. Stamens 3. Grain closely invested by the indurated lemma and palea; hilum oblong, short; embryo small; starch grains compound. Annual or perennial grasses. Leaf blades narrow with festucoid anatomy and oblong silica bodies, 2-celled microhairs absent; ligules membranous. First foliage leaf narrow, erect. Panicles contracted, sometimes spikelike. Chromosomes large; basic number 6 or 7. Genus:

Phalaris

Phalaris L., Sp. PL 54 (1753). SPIKELETS solitary, shortly pedicelled, not falling entire, FLOWER usually 1, sometimes a second flower present, GLUMES persistent, completely enclosing the lemmas, boat-shaped, almost plano-convex in outline, subobtuse, broadly winged on the keel, scabrid, membranous. LEMMAS 3, dissimilar; the two lower ones varying from more than 1/2 as long as the uppermost lemma to minute scales, empty, rarely one subtending a male flower, UPPERMOST LEMMA ovate-elliptic or ovatelanceolate in outline, obtuse or acute, chartaceous, 5-nerved, softly pilose, subtending a palea and a bisexual flower, PALEA oblong-linear in outline, ciliate on the keel, almost hyaline, LODICULES 2, hyaline. STAMENS 3. OVARY ellipsoid; styles distinct, plumose above, exserted from the top of the spikelet. GRAIN ovate in outline, enclosed by the lemma and palea; embryo 1/4 as long as the grain. Annuals or perennials; culm ribbed or smooth, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous; ligule hyaline; blade flat, closely ribbed, with the midrib distinct or scarcely so, glabrous ; inflorescence a dense oblong or, more rarely, a spikelike panicle, sometimes subcapitate or interrupted and lobed; primary branches of

120

G R A S S E S OF H A W A I I

the inflorescence sometimes bearing a definite cluster of 7 spikelets with the lowermost long-pedicelled and the upper three usually fertile. 1.

Phalaris arundinacea L., Sp. PL 55 (1753). [Fig. 39]

2.

P. californica Hook, et Am., Bot. Beechey's Voy. 161 (1841).

3.

P. caroliniana Walt., Fl. Carol. 74 (1788).

4.

P. coerulescens Desf., Fl. Atlant. 1:56 (1798).

5.

P. commutata Roem. et Schult., Syst. Veg. 2:403 (1817).

6.

P. minor Retz., Obs. Bot. 3:8 (1783).

7.

P. paradoxa L., Sp. PI. ed. 2, 2:1665 (1763).

8.

P. tuberosa L., Mant. PI. 2:557 (1771). P. tuberosa var. stenoptera (Hack.) Hitchc., Journ. Washington Acad. Sci. 24:292 (1934). P. stenoptera Hack, in Fedde, Repert. Sp. Nov. 5:333 (1908).

FIG. 39. Phalaris arundinacea L . (a) Floret, showing fertile and sterile lemmas, X 8; (b) spikelet, showing the strongly keeled glumes, fertile lemma, and palea, X 8; (c) panicle, interrupted below, X 2/5; (d) leaf sheath, ligule, blade, and node, X 4/5; (e) habit, showing creeping rhizome, X 1/8; ( f ) upper part of culm, showing panicle, X 1/8. (From H. L . Mason, A Flora of the Marshes of California.)

122

GRASSES

OF

HAWAII

STTPEAE Dumort., Obs. Gram. Belg. 83, 139 (1824). Spikelets all alike, bisexual, 1-flowered; rachilla disarticulating above the glumes, not prolonged beyond the floret. Glumes persistent, of a much more delicate texture than the lemma and palea, longer than the lemma, 1-3-nerved; lemma terete becoming slightly or strongly indurated at maturity, with convolute or involute margins, 5-7-nerved with nerves close together at the apex, often with a bearded, sharp or obtuse callus, awned from the entire or minutely 2-lobed tip or rarely the lobes produced into stout scabrid bristles; awn simple, continuous with or articulated to the tip of the lemma, bent or geniculate, twisted below, plumose or naked above the knee; palea of similar texture to the lemma, 2-keeled, 2-nerved. Lodicules 3. Stamens 3. Grain tightly embraced by the lemma and palea; hilum linear, almost the length of the grain or shorter; embryo short. Annual or perennial herbs often inhabiting very dry localities. Leaf blades narrow, often rolled, with the chlorenchyma evenly distributed between the vascular bundles; silica bodies simple, variable; 2-celled microhairs present, elongated; ligule membranous to scarious. First foliage leaf of the seedling narrow and erect. Inflorescence of spikelets arranged in open or contracted panicles. Chromosomes small; basic number 7 - 1 3 and 17. Genera:

Oryzopsis Stipa KEY TO THE GENERA OF STIPEAE

i.

1.

Lemma narrowly linear, long, slender, awn kneed; usually twisted below; callus long and sharp-pointed; lodicules 3, large Stipa p. 123 Lemmas elliptic, plump; awn eventually deciduous, not twisted but curved, callus short, obtuse, never long and pointed Oryzopsis p. 122

Oryzopsis Michx.,

Fl. Bor. Amer. 1:51 (1803).

SPIKELETS 1-flowered, disarticulating above the glumes, GLUMES about equal, obtuse to acuminate, L E M M A indurate, usually about as long as the glumes, broad, oval or oblong, nearly terete, usually pubescent, with a short, blunt, oblique callus, and an early deciduous, sometimes bent and twisted awn. PALEA enclosed by the edges of the lemma.

FESTUCOIDEAE - STIPEAE

123

Mostly slender perennials, with flat or often involute blades and terminal narrow or open panicles. 1. 2. 3.

Oryzopsis coerulescens (Desf.) Hack., Denkschr. Akad. Wiss. (Wien) 502:8, 75 (1885). O. hymenoides (Roem. et Schult.) Ricker in Piper, Contr. U. S. Natl. Herb. 1 1 : 1 0 9 (1906). O. miliacea (L.) Benth. et Hook, ex Aschers. et Schweinf., Mem. Inst. Egypte 2:169 (1887).

Stipa L., Sp. PI. 78 (1753). SPIKELET solitary, pedicelled, not falling entire; pedicel swollen at the apex, minutely pubescent, FLOWER I . GLUMES usually persistent, equal or subequal, as long as or much shorter than the lemma, lanceolate or linear in outline, acute, sometimes long-acuminate, 3-nerved, membranous, puberulous or glabrous, LEMMA I , lanceolate, usually convolute, acuminate or minutely 2-lobed, 3-7-nerved, awned, membranous, subtending a palea and the bisexual flower; awn 2 - 1 6 times as long as the body of the lemma, terminal, simple, kneed, twisted below, plumose or naked above the knee, sometimes disarticulating from the lemma. CALLUS usually pungent and bearded, PALEA enclosed in the lemma, and either equalling it or much shorter, LODICULES 2 or 3, equal and rather large or much reduced, STAMENS 3 ; anthers often barbellate at the apex. OVARY glabrous; styles free, plumose above, GRAIN narrow, subterete, tightly embraced by the hardened lemma and palea.

Perennials; culm striate, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous or pilose; ligule very narrow, membranous, sometimes minutely ciliate; blade flat, closely veined, with a distinct midrib, glabrous or pilose; inflorescence a lax panicle or a contracted raceme; primary branches usually in pairs or threes or whorled and distant on the main rachis, bearing spikelets on the upper half; spikelets sometimes direct on the main rachis. 1. 2. 3. 4. 5.

Stipa cernua Stebbins et Love, Madroño 6:137 (1941). S. comata Trin. et Rupr., Mém. Acad. Sci. Pétersb. sér. 6, 5 1 : 1 5 (1842). S. leucotricha Trin. et Rupr., Mém. Acad. Sci. Pétersb. sér. 6, 51:54(1842). S. papposa Nees, Agrost. Bras. 377 (1829). S. viridula Trin. Mém. Acad. Sci. Pétersb. sér. 6, 21:39 (1836).

124

GRASSES OF

TRITICEAE Dumort.,

HAWAII

O b s . G r a m . Belg. 82 (1823).

Hordeeae Spenner, Fl. Friburg. 1 : 1 5 5

as

Hordeaceae.

Spikelets 1 - to many-flowered, solitary or in groups of 2 - 6 (usually 3), mostly bisexual and sessile and alike, or the lateral spikelets of a cluster sometimes shortly pedicelled and male or sterile and m u c h reduced, alternating on the opposite sides of a continuous or jointed rachis; rachilla disarticulating above the glumes and between the florets or tough in cultivated cereals. G l u m e s coriaceous, strongly nerved, often reduced to awl-shaped empty structures, those of fertile spikelets awnless or a w n e d ; lemmas 5- to many-nerved, awnless or a w n e d ; awn short or long, straight or recurved, not twisted or kneed. L o d i c u l e s 2. Stamens 3. O v a r y obovate with a hairy appendage; styles 2, distinct, often very short; grain free or adhering to the lemma and/or palea; hilum long, linear; e m b r y o small; starch grains simple. A n n u a l or perennial herbs. L e a f sheaths usually with small auricles at the m o u t h ; blades narrow with festucoid anatomy; silica bodies oblong, rounded or elliptic; 2-celled microhairs absent. Inflorescence spikelike with a continuous or disarticulating rachis (spike axis). C h r o m o s o m e s large; basic number 7. Genera:

Agropyron

Secale

Elymus

Triticum

Hordeum KEY TO THE GENERA OF TRITICEAE i.

Spikelets normally solitary at each node of the spike axis 2.

Spikelets 2-flowered; glumes very narrow, subulate, 1 - n e r v e d ; lemmas stiffly hairy on the keels Secale p. 129

2.

Spikelets usually more than 2-flowered; glumes broader, usually more than 1 - n e r v e d ; lemmas not stiffly hairy on the keels or midrib 3.

N e r v e s of the lemmas not confluent at the tip; glumes and lemmas keeled on the back; annuals

.

.

Triticum P- 131

3.

N e r v e s of the lemmas confluent at the tip; glumes not keeled (except A. pectiniforme); perennials . . . Agropyron P- 125

FESTUCOIDEAE -

i.

TRITICEAE

125

Spikelets more than one at each node of the spike axis (solitary in part of the spike in some species of Elymus and in hybrids between Agropyron and Elymus) 4.

Spikelets 1-flowered, usually with a rudiment of the second floret Hordeum p. 126

4.

Spikelets 2-7-flowered

Elymus p. 126

Agropyron Gaertn., Nov. Comm. Acad. Sci. Petrop. 14:539 (1770). SPIKELETS solitary, sessile, not falling entire, FLOWERS 4-10. GLUMES with their edges turned to the axis of the inflorescence, similar, equal, lanceolate, keeled, 5- to many-nerved, coriaceous, glabrous; keel not median, L E M M A S 5 - 1 1 , exserted from the glumes, lanceolate, obtuse, sometimes 3-toothed or mucronate, sometimes acuminate or produced into an awn, 5- (rarely 7-9-)veined, coriaceous, glabrous, subtending a palea and a bisexual flower; uppermost lemma with a rudimentary palea. PALEAS subequalling the lemmas, elliptic, obovate, or lanceolate, shortly acuminate, truncate, 2-keeled, with the keels acute or somewhat winged or shortly ciliate. LODICULES 2, oblique or unequally lobed, densely ciliate. STAMENS 3. O V A R Y obovoid, with a large villous top; styles almost absent; stigmas delicately plumose, G R A I N linear-oblong in outline, tightly enclosed by the lemma and palea and adhering to them or at least to the palea. Perennials, often with creeping rhizomes, culm smooth, glabrous; leaf sheath longer than the internodes of the culm, glabrous; ligules scarious; blade rigid, convolute, ribbed, smooth, glabrous. Inflorescence a spike; internodes of axis concavo-convex; spikelets distichous.

1.

Agropyron cristatum (L.) (1770).

Gaertn., Nov. Comm. Petrop. 14:540

2.

A. dasystachyum (Hook.)

Scribti., Bull. Torrey Bot. Club 10:78

(1883). 3.

A. desertorum (Fisch.)

Schult.,

4.

A. elongatum (Host) Beauv., Ess. Agrost. 102, 146, 180 (1812).

5.

A. inerme (Scribn. 36:539 (1909).

et Smith)

Mantissa 2:412 (1824).

Rydb.,

Bull. T o r r e y Bot. Club

126

GRASSES OF H A W A I I

6. A. intermedium (Host) Beauv., Ess. Agrost. 102, 146, 180 (1812). 7. A. repens (L.) Beauv., Ess. Agrost. 102, 146, 180 (1812). [Fig. 40] 8. A. smithii Rydb., Mem. New York Bot. Gard. 1:64 (1900). 9. A. spicatum (Putsch.) Scribn. et Smith, U. S. Dept. Agr. Div. Agrost. Bull. 4:33 (1897). 10. A. trachycaulum (Link) Malte, Canada Natl. Mus. Ann. Rept. (1930); Bull. 68:42 (1932). A. pauciflorum (Schwein) Hitchc. in Silveus, Tex. Grasses 158 ( I 933)A. tenerum Vasey, Bot. Gaz. 10:258 (1885). 1 1 . A. trichophorum (Link) Richt., PI. Eur. 1 : 1 2 4 (1890). Elymus L., Sp. PI. 83 (1753). SP 1 RELETS 2- to 6-flowered, sessile in pairs (rarely 3 or more, or solitary) at each node of a usually continuous rachis, the rachilla distorted at base, bringing the florets more or less dorsiventral to the rachis; rachilla disarticulating above the glumes and between the florets. GLUMES equal, somewhat asymmetric, usually rigid, sometimes indurate below, narrow to subulate, 1- to several-nerved, acute to aristate. LEMMAS rounded on the back or nearly terete, obscurely 5-nerved, acute or usually awned from the tip. PALEA shorter than the glume, 2-keeled. STYLES very short, distinct, STIGMAS plumose. Usually tall, erect perennials, often with creeping rhizomes; inflorescence a slender or bristly spike.

1.

Elymus canadensis L., Sp. PI. 83 (1753).

2.

E. condensatus J . et C. Presl, Rel. Haenk. 1:265 (1830).

3.

E. glaucus Buckl., Proc. Acad. Nat. Sci. Phila. 1862:99 (1862).

4.

E. triticoides Buckl., Proc. Acad. Nat. Sci. Phila. 1862:99 (1862). [Fig- 4 i ] E. virginicus L., Sp. PI. 84 (1753).

5.

Hordeum L., Sp. PI. 84 (1753). SPIKELETS in groups of 3's at the end of a short pedicel, one sessile, two pedicelled, usually all awned, rarely one of the pedicelled spikelets absent or represented by the two glumes only, SESSILE SPIKELET i-flow-

FIG. 40. Agropyron repens (L.) Beauv. 1 , Plant, x 1/2; 2, spike, X Z; 3, spikelet, x 5. (U.S.D.A. photograph.)

128

GRASSES

OF

HAWAII

FIG. 41. Elymus triticoides Buckl. (a) Node, showing group of spikelets, X 4; (b) floret, the lemma removed to show the lodicules, X 4; (c) leaf sheath and ligule, X 4; (d) floret, X 4; (e) habit, X 1/8; ( f ) inflorescence, X 2/5. (From H. L . Mason, A Flora of the Marshes of California.)

FESTUCOIDEAE -

TRITICEAE

129

ered; rachilla produced as a bristle, GLUMES linear-lanceolate, long-ciliate, or reduced to 2 scabrid awns, L E M M A I , lanceolate, acuminate, awned, membranous, glabrous, subtending a palea and a bisexual flower; awn 2 - 4 times as long as the body of the lemma, scabrid. PALEA about as long as the lemma, lanceolate, acuminate, acute or truncate, sometimes 2-toothed, membranous, L O D I C U L E S 2, lanceolate, ciliate or ciliolate, hyaline, STAMENS 3. O V A R Y villous above; styles distinct, plumose above, laterally exserted. G R A I N tightly enclosed and usually adhering to the lemma and palea, oblong or narrowly oblong in outline; embryo short. PEDICELLED SPIKELETS both similar (very rarely one absent or represented by 2 glumes), 1 -flowered or both reduced to 2 glumes and a sterile lemma. G L U M E S linear-lanceolate, awned, ciliate or represented by 2 scabrid awns, L E M M A S lanceolate, acuminate, keeled, usually awned, scabrid on the keel, membranous, sometimes pilose within, empty or subtending a palea and a male or bisexual flower; awn, when present as long as or twice as long as the body of the lemma, PALEA, when present, subequalling its lemma, lanceolate, keeled, scabrid on the keels; membranous, sometimes pilose, O V A R Y , when present, often rudimentary. Annuals or perennials; culm ribbed, glabrous; leaf sheath longer or shorter than the internode of the culm, ribbed, glabrous; ligule membranous ; blade flat, linear, closely ribbed, smooth or scabrid, sometimes pubescent; inflorescence a dense spikelike raceme. 1.

Hordeum brachyantherum Nevski, Act. Inst. Bot. Acad. Sci. U.R.S.S. ser. 1, fasc. 2:61 (1936).

2.

H. bulbosum L., Cent. PI. 2:8 (1756). According to Chase and Niles (1952), '[Diagnosis and description; base, root and c u l m = i / . bulbosum L . s p i k e = H . murimim L.] fide C. E. Hubbard.'

3.

H. murinum L., Sp. PI. 85 (1753).

4.

H. nodosum L., Sp. PI. 126 (1762). According to Chase and Niles (1962) ' L ' s nodosum is a mixture, fides Hubbard; description suggests "H. bulbosum" L., also a mixture. United States species so called is H. brachyantherum.'

5.

H. vulgare L., Sp. PI. 84 (1753). [Fig. 42]

Secale L., Sp. PI. 84 (1753). SPIKELETS FLOWERS

2.

solitary, sessile, falling entire with an internode of the axis. equal or slightly unequal, linear-subulate, keeled,

GLUMES

130

GRASSES

OF

HAWAII

FIG. 42. Hordeum vulgare L. Plant, x 1/2; group of spikelets and floret, x 3; (a) spike of beardless barley, X 1/2. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FESTUCOIDEAE -

TRITICEAE

stiff, with membranous margins, rigidly ciliate on the keels, LEMMAS 2, more or less exserted from the glumes, oblong or lanceolate, passing into a straight awn, 5-nerved, keeled, firmly membranous, subtending a palea and a bisexual flower; awn slightly shorter than the lemma, scabrid. PALEAS as long as or almost as long as the lemmas, lanceolate, 2-toothed, 2-keeled, membranous, rigidly ciliate on the keels, LODICULES 2, hyaline, ciliate. STAMENS 3. OVARY obovoid, densely tomentose above; styles distinct, plumose above, laterally exserted. GRAIN oblong in outline, enclosed by the lemma and palea, free or almost so; embryo 1/3 the length of the grain. Annuals or perennials; culm ribbed, glabrous; leaf sheath ribbed, glabrous; ligule short, scarious; blade linear, ribbed, minutely pubescent ; inflorescence a dense cylindric spike; internodes of axis flattened, densely pilose on the margins. 1.

Secale cereale L., Sp. PI. 84 (1753). [Fig. 43]

Triticum L., Sp. PI. 85 (1753). SPIKELETS solitary, sessile, slightly laterally compressed or turgid, when flattened with the flattened side towards the rachis, not falling, or rarely falling entire, the rachilla disarticulating above the glumes and between the lemmas, FLORETS 2-5, bisexual, or the uppermost sterile. GLUMES rigid, equal or subequal, more or less keeled, 3- to severalnerved, the apex abruptly mucronate or toothed, or terminating in 1 or several awns, LEMMAS 2-5, exserted from the glumes, rigid, broad, keeled, more or less asymmetrical, many-nerved, abruptly pointed or awned; callus none, PALEA shorter than lemma or nearly as long, 2keeled, the keels ciliate, entire or splitting into two longitudinal halves when mature, LODICULES 2, rather thick, STAMENS 3. OVARY pubescent or villous at the apex; styles 2, short; stigmas plumose, GRAIN ovoid or oblong in outline, subterete, grooved in front, enclosed by the lemma and palea, either entirely free, or adhering to both lemma and palea, pubescent at the apex; embryo rather small.

Short or tall annual grasses; leaf blade flat, somewhat flaccid, with more or less conspicuous auricles at the base; ligule hyaline; inflorescence a spike, the rachis more or less zigzag, each internode flattened and curved, and presenting the concave side to the spikelet. 1.

Triticum aestivum L., Sp. PI. 85 (1753). [Fig. 44] T. vulgare Vill., Hist. PI. Dauph. 2:153 (1787). T. sativum Lamk., Fl. Franc. 3:625 (1778).

132

GRASSES

OF

HAWAII

F i g . 43. Secale cereale L . Plant, X 1/2; spikelet, x 3; floret, X 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FESTUCOIDEAE -

TRITICEAE

*33

FIG. 44. Triticum aestivum L . Plant with awned spike (bearded wheat) and a nearly awnless spike (beardless wheat), x 1/2; spikelet and floret, X 3. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

J

34

GRASSES OF

HAWAII

CHLORroEAE Agardh, Aphor. Bot. 150 (1823). Cynodoneae Dumort., Obs. Gram. Belg. 83, 140 (1823). Spikelets 1- to several-flowered, slightly compressed laterally, with one floret bisexual and with or without imperfect florets above or below it; rachilla disarticulating above the persistent glumes; glumes herbaceous, pale, translucent, 1-3-nerved; lemmas membranous to chartaceous, entire, emarginate or 2-lobed, awned or awnless, 1 - 3 nerved with the lateral nerves near the margins and often ciliate; palea often obovate, 2-keeled, narrowly winged on the keels. Lodicules 2, rarely absent. Stamens 3. Ovary glabrous. Styles 2; stigmas plumose. Grain enclosed between the scarcely changed lemma and palea; hilum in a small depression at the base of the grain; embryo about 1/2 as long as the grain; starch grains compound. Annual or perennial herbs; leaf blades narrow, with panicoid anatomy; silica bodies saddle-shaped or transversely dumbbell-shaped; microhairs two-celled, swollen or round; ligule a ciliate rim or membranous. Spikelets sessile or subsessile, secund in one or two rows on the continuous rachis of solitary, digitate or scattered spikes or spikelike racemes. First foliage leaf of the seedling broad, flat, horizontal. Chromosomes small; basic number 9, 10, rarely 7. Genera:

Bouteloua

Enteropogon

Buchloe

Schedonnardus

Chloris

Spartina

Cynodon KEY TO THE GENERA OF CHLORIDEAE i.

Male and female flowers in separate inflorescences

1.

Flowers bisexual or sometimes reduced

.

Buchloe P- 137

2.

Inflorescence terminal, spicate, not of digitate or racemose spikes; Enteropogon lemmas not obovate p. 146

2.

Inflorescence of digitate spikes or spikes racemose, very rarely solitary, but if so the lemmas obovate 3.

Maritime grasses with erect spikes

Spartina p. 149

ERAGROSTOIDEAE -

3.

CHLORIDEAE

i3S

Not maritime 4.

Spikelets racemose, distant falling entire

4.

Spikes not falling entire

.

Bouteloua P- 135

5.

Spikelets with one or more fertile florets and one or more imperfect florets above them; lemmas usually widened upwards Chloris p. 138

5.

Spikelets with one fertile floret and no imperfect florets 6.

Spikelets disarticulating above glumes

6.

Spikelets falling entire .

Cynodon p. 142

Schedonnardus p. 149

Bouteloua Lagasca, Var. Cienc. 2:4, 134 (1805). SPIKELETS 1-flowered, with the rudiments of one or more florets above, sessile in two rows along one side of the rachis. GLUMES 1-nerved; acuminate or awn-tipped, the first shorter and narrower, LEMMAS as long as the second glume or a little longer; 3-nerved, the nerves extending into the short awns or mucros, the internerves usually extending into lobes or teeth, PALEA sometimes 2-awned; rudiment various, usually 3-awned, the awns usually longer than those of the fertile lemma, a secondary rudimentary floret sometimes present. Perennial or sometimes annual, low or rather tall grasses, with 2 to several or many spikes racemose on a common axis, or sometimes solitary, the spikelets few to many in each spike, rarely solitary, pectinate or more loosely arranged and appressed, the rachis of the spike usually naked at the tip. The sterile florets forming the rudiment are variable in all the species and commonly in individual specimens. The general pattern of the rudiment is fairly constant for each species, the variability being in the reduction or increase in number and size of the sterile florets, the reduction from 3 awns to 1, and in the amount of pubescence.

1.

Bouteloua chondriosioides (H.B.K.) Amer. Acad. 18:179 ( i 8 83)-

Benth. ex Wats., Proc.

2.

B. curtipendula (Michx.) Torr. in Emory, Notes Mil. Reconn. 154 (1848). [Fig. 45] Chloris curtipendula Michx., Fl. Bor. Amer. 1:59 (1803).

FIG. 45. Bouteloua curtipettdula(Michx.) Torr. Plant, X 1/2; spikelet and florets, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

ERAGROSTOIDEAE -

CHLORIDEAE

137

3.

B. eriopoda (Ton.) (1856).

Ton., U. S. Expl. Miss. Pacif. Rept. 4:155

4.

B. filiformis (Fourn.) Griffiths, Contr. U. S. Natl. Herb. 14:413 (1912).

5.

B. gracilis (H.B.K.) Lag. ex Steud., Nom. Bot. ed. 2 , 1 : 2 i g (1840). B. oligostachya (Nutt.) Torr. ex A. Gray, Man. Bot. ed. 2, 553 (1856).

6.

B. heterostega (Trin.) Griffiths, Contr. U. S. Natl. Herb. 14:414 (1912).

7.

B. hirsuta Lag., Var. Cienc. 4:141 (1805).

8.

B. parryi (Fourn.) (1912).

9.

B. rothrockii Vasey, U. S. Dept. Agr. Spec. Rept. 63:33 (1883) nom. nud., Contr. U. S. Natl. Herb. 1:268 (1893).

Griffiths, Contr. U. S. Natl. Herb. 14:381

Buchloe Engelm., Trans. Acad. Sci. St. Louis 432 (1859). STAMINATE SPIKELETS 2-flowered, sessile and closely imbricate, in two rows on one side of a slender rachis, forming a short spike, GLUMES somewhat unequal, rather broad, 1 -nerved, LEMMAS longer than the glumes, 3-nerved, rather obtuse, whitish, PALEA as long as its lemma. PISTILLATE SPIKELETS mostly 4 or 5 in a short spike or head, this falling entire, usually 2 heads to the inflorescence, the common pedicel short and included in the somewhat inflated sheaths of the upper leaves, the thickened indurate rachis and broad outer (second) glumes forming a rigid white obliquely globular structure crowned by the green-toothed summit of the glumes, FIRST GLUME (inside) narrow, thin, mucronate, well developed to obsolete in a single head, SECOND GLUME firm, thick and rigid, rounded on the back, obscurely nerved, expanded in the middle, with inflexed margins, enveloping the floret; abruptly contracting above, the summit with 3 green rigid acuminate lobes, LEMMA firmmembranous, 3-nerved, dorsally compressed, broad below, narrowed into a 3-lobed green summit, the middle lobe much larger, PALEA broad, obtuse, about as long as the body of the lemma, enveloping the grain.

A stoloniferous perennial with short curly blades, the staminate flowers in 2 or 3 short spikes on slender erect culms, the pistillate in sessile heads partly hidden among the leaves. Plants dioecious or monoecious.

GRASSES OF

i.

HAWAII

Buchloe dactyloides (Nutt.) Engelm., Trans. Acad. Sci. St. Louis 1:432 (1859). [Fig. 46]

Chloris Sw., Prodr. Veg. Ind. Occ. 25 (1788). SPIKELETS solitary, sessile, not falling entire, FLOWERS 1-3. GLUMES persistent, unequal, linear, membranous; lower glume about 1/2 as long as the upper; upper as long as lemmas, 2-lobed, mucronate from the sinus to the keel passing into a short acumen, LEMMAS 2-4, dissimilar. LOWER or LOWERMOST LEMMA obovate-acuminate or lanceolate, folded, awned below the tip, with a tuft of long cilia on the upper portion or scantily ciliate, membranous or chartaceous, subtending a palea and a bisexual flower; awn slightly shorter than to 4 times as long as the body of the lemma, PALEA as long as its lemma, linear to obovate-Iinear, 2-keeled, hyaline, SECOND LEMMA spathulate or obovate, truncate, awned, membranous or chartaceous, glabrous, empty or subtending a palea and either a male or a bisexual flower; awn shorter than or 3-5 times as long as the body of the lemma, THIRD and FOURTH LEMMAS (when present) empty; the upper often reduced to little more than an awn. LODICULES 2, minute, STAMENS 3 . OVARY glabrous; styles distinct, plumose above, GRAIN narrow-ovate in outline; pericarp very thinly and delicately hyaline, loose, swelling slightly in water; embryo rather large. Perennials; culm ribbed, glabrous; leaf sheath shorter than the internodes of the culm, ribbed, glabrous; ligule membranous or a row of cilia; blade flat, ribbed, with a prominent midrib, smooth or scabrid, glabrous or rarely with long hairs from bulbous bases; inflorescence of few to many linear spikes arranged digitately at the end of the culm.

1.

Chloris barbata (L.) Sw., Fl. Ind. Occ. 1:200 (1797). [Fig. 47] C. inflata Link, Enum. PI. Hort. Berol. 1:105 (1821). C. paraguayensis Steud., Syn. PI. Glum. 1:204 (1854). Andropogon barbatus L., Mant. PI. Alt. 302 (1771), non L. (1759).

2.

C. berroi Arech., Anal. Mus. Nac. Montevideo 1:388 (1896).

3.

C. ciliata Sw., Prodr. Veg. Ind. Occ. 25 (1788).

4.

C. cucullata Bisch., Ann. Sci. Nat. 111. Bot. 19:357 (1853).

5.

C. distichophylla Lag., Gen. et Sp. Nov. 4 (1816).

6.

C. divaricata R. Br., Prodr. Fl. Nov. Holl. 1 :i86 (1810). [Fig. 48]

139

FIG. 46. Buchloe dactyloides (Nutt.) Engelm. Pistillate and staminate plants, X 1/2; pistillate spike and floret, x 5; staminate spikelet, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A photograph.)

140

GRASSES

OF

HAWAII

FIG. 47. Chloris barbata (L.) Sw. Plant, X 1 jz; glumes and spikelet, x 6. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

ERAGROSTOIDEAE - CHLORIDEAE

141

FIG. 48. Chloris divaricata R. Br. Plant, X 1; (a) spikelet, x 10; (b) second glume, X 10; (c) first glume, x 10; (d) lemma, x 10; (e) seed, X 10. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

142

GRASSES OF H A W A I I

7. C. gayana Kunth, Rev. Gram. 1189 (1829) nomen; 293 (1830), pi. 58; ex Stapf in Dyer, Fl. Cap. 7:642 (1900). [Fig. 49] 8. C. polydactyla (L.) Sw., Prodr. Veg. Ind. Occ. 26 (1788). Andropogon polydactylon L., Sp. PI. ed. 2, 1483 (1763). A. barbatum L., Amoen. Acad. 5:412 (1759). C. barbata (L.) Nash, Bull. Torrey Bot. Club 25:443 (1898). 9. C. radiata (L.) Sw., Prodr. Veg. Ind. Occ. 26 (1788). [Fig. 50] Agrostis radiata L., PI. Jam. Pugill. 7 (1759); Syst. Nat. ed. 10, 2:873 (*759)10. C. truncata R. Br., Prodr. Fl. Nov. Holl. 186 (1810).

11. C. virgata Sw., Fl. Ind. Occ. 1:203 (I797)- [Fig- 51] C. elegens H.B.K., Nov. Gen. et Sp. 1:66 (1816). Cynodon Rich, in Pers., Syn. PI. 1:85 (1805). Capriola Adans., Fam. PI. 2:31 (1763). SPIKELETS solitary, awnless, sessile in two rows along one side of a slender continuous rachis and appressed to it, not falling entire; rachilla sometimes produced, FLOWER I. GLUMES equal or unequal, lanceolate or boat-shaped, acute, keeled, sometimes scabrid on the keel; upper glume usually deciduous; lower glume subpersistent. LEMMA I , very rarely a second reduced lemma present, ovate, ovate-lanceolate, or elliptic-lanceolate, obtuse, keeled, ciliate on the keel, membranous, subtending a palea and a bisexual flower, PALEA as long as or somewhat shorter than the lemma, oblong-linear, obtuse or subacute, membranous. LODICULES 2, minute, glabrous, STAMENS 3. OVARY glabrous; styles distinct, plumose above, GRAIN oblong in outline; embryo about 1/3 as long as the grain.

Creeping perennials, rooting at the nodes; culm smooth, glabrous; ligule ciliate; blade linear, closely ribbed, with a distinct midrib; inflorescence of 2-7 narrow spikes arranged digitately at the end of the culm, very rarely only 1 spike present; rachis keeled; spikelets unilateral in two rows on the rachis. 1.

Cynodon arcuatus J. S. Presl ex C. B. Presl, Rel. Haenk. 1:290 (1830).

2.

C. barberi Rang, et Tad., Journ. Bombay Nat. Hist. Soc. 24:846 (1916).

ERAGROSTOIDEAE

-

CHLORIDEAE

r

43

FIG. 49. Chloris gayana Kunth. 1, Plant and inflorescence, x 1/2. 2, Spikelet; 3, spikelet with glumes removed; 4, lower glume; 5, upper glume; 6, lower lemma; 7, palea of lowest lemma; all X 6. 8, Ligule, X 1. (From N . L . Bor, T h e Grasses of Burma, Ceylon, India and Pakistan.)

144

ERAGROSTOIDEAE -

CHLORIDEAE

145

FIG. SI. Chloris virgata SW. A, Plant, x 1/2; B, glumes, x 6; C, florets, x 6; D, spikelets, x 6. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

146

GRASSES OF H A W A I I

3.

C. dactylon (L.) Pers., Syn. PI. 1:85 (1805). [Fig. 52] Capriola dactylon (L.) O. Ktze., Rev. Gen. PI. 2:764 (1891). Panicum dactylon L., Sp. PI. 58 (1753). Digitaria stolonifera Schrad., Fl. Germ. 1 : 1 6 5 (1806). D. dactylon (L.) Scop., Fl. Cam. ed. 2, 1:52 (1772).

4.

C. dactylon var. maritimus (H.B.K.) Hack, in Fries, Arkiv Bot. 8:40 (1909). C. maritimus H.B.K., Nov. Gen. et Sp. 1 : 1 7 0 (1816). Capriola dactylon var. maritimus (H.B.K.) Hitchc., U. S. Dept. Agr. Bull. 722, 179 (1920). Cynodon dactylon Hitchc., U. S. Dept. Agr. Misc. Publ. 200, 484 (1935), non Cynodon dactylon Pers., Syn. PI. 1 ¡85 (1805).

5.

C. magennisii Hurcombe, Journ. So. Afr. Bot. 13:109, 1 1 5 (1947). (A natural hybrid between C. dactylon and C. transvaalensis.)

6.

C. plectostachyus (K. Schum.) Pilger, Bot. Jahrb. Engler 40:82 (1907). [Fig. 53]

7.

C. transvaalensis Burtt.-Davy,

Kew Bull. 1921:281 (1921).

Enteropogon Nees in Lindl., Introd. Nat. Syst. ed. 2, 448 (1863). SPIKELETS solitary, sessile, not falling entire, FLOWER I . GLUMES persistent, unequal; lower glume about 1/2 as long as the upper, lanceolate, acuminate, acute, produced into a short awn, 1-nerved, membranous; upper lanceolate, 1-nerved, produced into an awn, membranous; awn about 1 ¡2 as long as the body of the glume, LEMMAS 3 ; lowest lanceolate, 2-lobed, awned from the sinus, 3-nerved, membranous, subtending a palea and the bisexual flower; awn about twice as long as the body of the lemma, RACHILLA produced beyond the lemma, tufted at the apex. PALEA lanceolate, shortly 2-lobed, 2-keeled, concave between the keels, minutely ciliolate on the keels above, LODICULES none, OVARY oblong in outline, glabrous; styles distinct, plumose above, laterally exserted. SECOND LEMMA similar to the lowest but smaller and subtending a palea only; uppermost lemma reduced to an awn.

Perennials; culm smooth, striate, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous; ligule a fringe of long hairs; blade long, narrow-linear, folded, ribbed, glabrous; inflorescence a solitary long narrow spike at the end of the culm. 1.

Enteropogon machrostachya (Höchst.) Munro ex Benth., Journ. Linn. Soc. 19:101 (1881).

ERAGROSTOIDEAE -

Schedonnardus Steud., Gram. 146 (1855).

CHLORIDEAE

149

Flora 33 (1850) 228, 229, nomen; Syn. PI.

SPIKELETS i-flowered, sessile and somewhat distant in two rows on one side of a slender, continuous 3-angled rachis, appressed to its slightly concave sides, the rachilla disarticulating above the glumes, not prolonged. GLUMES narrow, stiff, somewhat unequal, acuminate, 1-nerved. L E M M A S narrow, acuminate, a little longer than the glumes, 3-nerved. L o w tufted perennial, with stiff, slender, divergent spikes rather remote along a common axis.

1.

Schedonnardus paniculatus (Nutt.) Trel. in Branner and Coville, Rept. Geol. Surv. Ark. 1888, 4:236 (1891). [Fig. 54]

Spartina Schreber, Gen. 43 (1789). solitary, much flattened laterally, subsessile, falling entire. 1. GLUMES unequal, dissimilar, 1-5-nerved, keeled; lower glume linear, acute or obtuse, ciliate on the keel, membranous; upper glume shorter than the lemma, narrow-linear, membranous or subcoriaceous, pubescent, L E M M A oblong, 1-nerved, thinly membranous, subtending a palea and a bisexual flower, PALEA similar to its lemma in shape, 2-nerved, delicate in texture, LODICULES none, STAMENS 3. O V A R Y glabrous; styles very long, connate at the base, plumose above, G R A I N linear or oblong in outline, included in the lemma and palea. SPIKELETS

FLOWER

Erect, often stout, maritime grasses, usually tall and coarse, with tufted or creeping bases (firm scaly rhizomes); culm glabrous; leaf sheath longer than the internode of the culm, ribbed, glabrous; ligule a minute line of hairs; blade scarcely ribbed, folded, awl-shaped, rigid; inflorescence of 2-3 spikes from the apex of the culm, rarely of a single spike; rachis prominently keeled on the face, flattened on the back; spikelets 2-ranked on the rachis. 1.

Spartina x townsendii H. et J. Groves, Rept. Bot. Exch. Club Brit. Isles 1880:37 (1881).

Spartina x townsendii is the male sterile primary hybrid between 5 . maritima (S. striata) X S. alterniflora with 2n = c. 60-63 chromosomes. T h e plant introduced into Hawaii is the fertile seed-yielding derivative of S. x townsendii with double the number of chromosomes, 2n = c. 122, and which at this time is without a name.

FIG. 54. Schedonnardus paniculatus (NuttjTrel. Plant, x 1/2; spikelet and floret, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U.S.D.A. photograph.)

ERAGROSTOIDEAE - ERAGROSTEAE

ERAGROSTEAE Stapf in Dyer, Fl. Cap. 7:316 (1898). Spikelets 2- to many-flowered, bisexual, usually strongly laterally compressed, rarely terete; rachilla disarticulating above the glumes and the spikelets breaking up from above downwards, or somewhat tough and the glumes and lemmas falling from below upwards in succession. Glumes sometimes persistent, membranous to coriaceous, usually shorter than the lemmas; lemmas exserted from the glumes, membranous to chartaceous, not becoming firmer at maturity, 1-3-nerved, emarginate or 2-4-lobed at the tip, awnless, mucronate or with a straight awn from the tip or sinus, rarely with the three nerves running out as awns; paleas hyaline, 2-nerved, 2-keeled, occasionally winged on the keels which are scabrid or rarely ciliate, usually caducous, rarely persistent. Lodicules 2. Stamens 2 or 3. Grain sometimes with a free pericarp, loosely or tightly enclosed between the lemma and palea; hilum punctiform, oblong or ellipsoid; embryo about 1/2 the length of the grain; starch grains compound. Annual or perennial herbs. Leaf blades narrow, often setaceous, with panicoid anatomy; microhairs 2-celled, swollen, club-shaped; silica bodies saddle-shaped; ligule a ciliate rim, rarely membranous. Spikelets pedicelled in open or contracted panicles, or sessile or subsessile in secund spikes or spikelike racemes. Chromosomes small; basic number 9, 10. Genera:

Dactyloctenium Eleusine Eragrostis Leptochloa

Trichoneura Tridens Triraphis

K E Y T O T H E G E N E R A OF E R A G R O S T E A E

i.

Lemmas usually entire at the apex, obtuse, acute or acuminate, or if 2-toothed or awned from the tip then glabrous near the margins and along the side nerves; cleistogamous spikelets not developed in the axils of the leaf sheaths 2.

Spikelets in open, contracted or spikelike panicles, rarely in solitary secund spikes 3.

Lemmas rounded on the back, 3-5-nerved; grain hollowed out on the adaxial surface, rugose on the other

152

GRASSES

3.

OF

HAWAII

Lemmas more or less acutely keeled, 3-nerved; grain not rugose or hollowed out on the adaxial surface Eragrostis P- 155

2.

Spikelets sessile or very short-pedicelled, loosely to densely imbricate in digitate or racemosely arranged spikes or spikelike racemes, very rarely in solitary spikes (Eleusine sp.) 4.

4.

1.

Axis and branches of the inflorescence ending in a spikelet Eleusine P- i53 Axis of spikes terminating in a sharp point; upper glume mucronate or awned; spikes digitate, rarely racemose Dactyloctenium P- 152

Lemmas usually emarginate or 2-4-lobed or toothed at the apex, rarely entire and then hairy along the nerves, frequently mucronate or awned; cleistogamous spikelets sometimes developed in the sheaths; racemes densely spiculate, conspicuously secund; spikelets oblong, laterally compressed; lemmas keeled 5.

Spikelets in slender, more or less secund, racemosely arranged panicles 6.

Glumes usually exceeding the florets; lateral nerves of the lemmas long ciliate Trichoneura p. 169

6.

Glumes shorter than the florets; lateral nerves not longciliate Leptochloa p. 169 Spikelets in loose or contracted panicles

5. 7.

Lemmas awnless

7.

Lemmas 3-awned

Tridens p. 171 Triraphis p. 174

Dactyloctenium Willd., Enum. Hort. Berol. 1029 (1809). SPIKELETS solitary, sessile, not falling entire, FLOWERS 2-4. GLUMES dissimilar subequal, membranous, glabrous; lower glume persistent, boat-shaped, sometimes shortly acuminate, keeled, ciliate on the keel; upper glume deciduous, oblong, with an awn from the back of the tip;

ERAGROSTOIDEAE -

ERAGROSTEAE

iS3

times as long as the body of the awn erect or somewhat spreading, glume, scabrid. LEMMAS 3-5, similar, decreasing in size upwards, exserted from the glumes, ovate, much compressed, shortly mucronate, sometimes long-acuminate, keeled, membranous; the lower lemmas each subtending a palea and a bisexual flower; the uppermost empty. PALEAS shorter than their respective lemmas, ovate-lanceolate or oblonglanceolate, 2-keeled, deeply concave between the keels, ciliate on the keels, membranous, LODICULES 2, minute, hyaline, STAMENS 3. OVARY glabrous; styles distinct, plumose above, subterminally exserted. GRAIN subglobose, rugose or punctate; pericarp very delicate, irregularly breaking away; embryo scarcely equalling 1/2 the length of the grain. Annuals sometimes stoloniferous and rooting at the nodes; culm simple or branched, striate, glabrous; leaf sheath striate, glabrous; ligule membranous; blade flat, linear, distinctly ribbed, with a distinct midrib, ciliate with tubercule-based hairs, blade glabrous or pilose with tubercule-based hairs; inflorescence of 2-4 spikes, arranged digitately or subdigitately at the end of the culm; spikelets crowded and unilateral on the rachis. 1.

Dactyloctenium aegyptium (L.) Beauv., Ess. Agrost. 15 (1812).

[Fig- 55] Cynosurus aegyptius L., Sp. PI. 72 (1753). Eleusine aegyptia (L.) Desf., Fl. Atlant. 1:85 (1798). Eleusine Gaertn., Fruct. & Sem. 1:7 (1788). SPIKELETS solitary, sessible, not falling entire, FLOWERS 3-6. GLUMES persistent, unequal, keeled, scabrid on the keels, 3-5-nerved, herbaceous, with membranous margins, glabrous; lower glume 1/2 as long as the upper, narrowly winged on keel, LEMMAS 3-6, exserted from the glumes, similar, resembling the upper glume, 3-nerved near the base, each subtending a palea and a bisexual flower, PALEAS slightly shorter than their respective lemmas, notched at the apex, 2-keeled, with keels winged, LODICULES 2, minute, cuneate. STAMENS 3. OVARY glabrous; styles slender from a broadened base, distinct, plumose above, laterally exserted. GRAIN globose or broadly oblong in outline; pericarp loose, delicate, breaking up irregularly or almost circumscissle; seed finely striate; embryo suborbicular.

Annuals; culm simple or branched, somewhat compressed, ribbed, glabrous; leaf sheath longer or shorter than the internode of the culm,

GRASSES OF

HAWAII

FIG. 55. Dactyloctenium aegyptium ( L . ) B e a u v . Plant, x i . i, l i g u l e ; 2, s p i k e l e t ; 3, l e m m a f r o m t h e side; 4, p a l e a ; all x 6. ( F r o m N . L . Bor, T h e G r a s s e s of B u r m a , C e y l o n , India and Pakistan.)

ERAGROSTOIDEAE -

ERAGROSTEAE

!55

compressed, strongly ribbed, glabrous or with few scattered hairs; ligule membranous; blade flat or longitudinally folded, linear, closely ribbed, with a distinct midrib, smooth, glabrous or with a few scattered long hairs; inflorescence of 3 - 1 2 spikes arranged digitately or subdigitately at the end of the culm; spikelets crowded and unilateral on the flattened rachis of the spikes. 1.

Eleusine coracana (L.)

2.

E. indica (L.)

Gaertn., Fruct. & Sem. 1:8 (1788).

Gaertn., Fruct. & Sem. 1:8 (1788). [Fig. 56]

Cynosurus indicus L., Sp. PI. 72 (1753). E. gracilis Salisb., Prodr. Stirp. 19 (1796). Cynodon indicus (Gaertn.) Rasp., Ann. Sci. Nat. Bot. 5:303 (1825). E. polydactyla Steud., Syn. PL Glum. 1:211 (1854). Eragrostis Beauv.,

Ess. Agrost. 70 (1812), emend. Reichb. (1828).

Eragrostis Host, Ic. Gram. Austr. 4:14 (1809) sine descr. Megastachya

Beauv., Ess. Agrost. 74, 167 (1812) partim.

solitary, pedicelled, rarely sessile, not or rarely falling entire. 2 to many, GLUMES persistent or deciduous, equal or unequal, 1-nerved or the upper sometimes 3-nerved, keeled, usually membranous. L E M M A S 2 to many, more or less imbricate, ovate or lanceolate or almost orbicular, acute or obtuse, entire, muticous, 3-nerved, with the side nerves short or almost precurrent, membranous to chartaceous, glabrous or very rarely minutely pubescent; all except the uppermost subtending a palea and a bisexual flower, PALEAS subequal to the lemmas, 2-keeled, membranous, deciduous or persistent on the rachilla. L O D I C U L E S 2, more or less fleshy, S T A M E N S 3, rarely 2. O V A R Y glabrous; styles distinct, plumose above, laterally exserted. GRAIN enclosed by the scarcely altered lemma and palea and deciduous with them or more commonly falling with the deciduous lemma leaving the more or less persistent palea behind, oblong, ovate or round in outline; embryo often 1/2 as long as the grain. SPIKELETS

FLOWERS

Annuals or perennials, sometimes forming cushion-like growths; culm smooth or striate, glabrous or villous; leaf sheath striate, shorter than the internode of the culm, glabrous or sometimes pilose from bulbous-based hairs; ligule a row of hairs, very rarely membranous; blade linear, rolled, or setaceous, finely ribbed, often scabrid; inflorescence a lax or contracted panicle, more rarely a raceme or composed of

GRASSES OF HAWAII

FIG. 56. Eleusine indica (L.) Gaertn. (From D. Meredith, The Grasses and Pastures of South Africa.)

ERAGROSTOIDEAE -

ERAGROSTEAE

157

spikes arranged on the main axis; primary branches solitary or more or less whorled on the main axis, sometimes distant; spikelets sometimes secund and 2-ranked on the rachis. 1. Eragrostis atropioides Hillebr., Fl. Haw. Is. 531 (1888). [Fig. 57] 2. E. brownei (Kunth) Nees in Hook, et Arn., Bot. Beechey's Voy. 253 (1838); et Fl. Afr. Austr. 396 (1841). Poa brownei Kunth, Rev. Gram. 1 : 1 1 2 (1829). 3. E. chloromelas Steud., Syn. PI. Glum. 1:271 (1854). 4. E. cilianensis (All.) Vignolo-Lutati, Malpighia 18:386 (1904). [Fig. 58] E. major (L.) Host, Ic. Gram. Austr. 4:14, pi. 24 (1809). E. megastachya (Koel.) Link, Hort. Berol. 1:187 (1827). Poa cilianensis All., Fl. Pedem. 2:246 (1785). See Bor (i960) for a discussion of the nomenclature of E. cilianensis and E. poaeoides. 5. E. curvula (Schrad.) Nees, Fl. Afr. Austr. 397 (1841). 6. E. deflexa Hitchc., Mem. Bernice P. Bishop Mus. 8:131 (1922). [Fig- 59] 7. E. elongata (Willd.) Jacq., Ecol. Gram. pi. 3 (1813). 8. E. falcata (Gaud.) Gaud, ex Steud., Nom. Bot. ed. 2, 1 ¡563 (1840). Poa falcata Gaud, in Freyc., Voy. Uran. Bot. 408 (1830). 9. E. fosbergii Whitney, Occas. Papers Bernice P. Bishop Mus. 13:75 (1937). [Fig. 60] 10. E. grandis Hillebr., Fl. Haw. Is. 528 (1888). [Fig. 61] 1 1 . E. grandis var. oligantha Hillebr., Fl. Haw. Is. 528 (1888). 12. E. grandis var. polyantha Hillebr., Fl. Haw. Is. 528 (1888). 13. E. hosakai Degener, Fl. Hawaiiensis 4: descr. with pi. Feb. 10 (1940). [Fig. 62] 14. E. intermedia Hitchc., Journ. Washington Acad. Sci. 23:450 (I933)15. E. lehmannia Nees, Fl. Afr. Austr. 402 (1841). 16. E.leptophyllaHitchc., Mem. BerniceP. Bishop Mus. 8:133 (1922). [Fig- 63]

GRASSES OF H A W A I I

FIG. 57.

Eragrastis atropioides Hillebr. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

ERAGROSTOIDEAE -

ERAGROSTEAE

iS9

FIG. 58. Eragrostis cilianensis (All.) Vignolo-Lutati. Plant, x 1 ¡2; spikelet, X 3 ; floret, x 5; grain, x 20. (From O. Degener, Flora Hawaiiensis.)

i6o

GRASSES OF H A W A I I

ERAGROSTOIDEAE -

ERAGROSTEAE

FIG. 60. Eragrostis fosbergit Whitney. (From O. Degener, Flora Hawaiiensis.)

L6L

IÖ2

FIG. 6I. Eragrostis grandis Hillebr. Plant, X 1/6; panicle, X 1/6; (From O. Degener, Flora Hawaiiensis.)

spikelet, X 3.

ERAGROSTOIDEAE -

ERAGROSTEAE

163

FIG. 62. Eragrostis hosakai Degener. Plant, x 1/2; spikelet, front and side view, x 4; floret, x 10. (From O. Degener, Flora Hawaiiensis.)

164

GRASSES OF H A W A I I

ERAGROSTOIDEAE -

ERAGROSTEAE

i6 5

17. E. leptostachya (R. Br.) Steud., Syn. PI. Glum. 1:279 ( I ^54)18. E. mauiensis Hitchc., Mem. Bernice P. Bishop Mus. 8:129 ( x 9 2 2 )[Fig. 64] 19. E. m e x i c a n a (Hörnern.) Link, Hort. Berol. 1:190 (1827). 20. E. montícola (Gaud.) Hillebr., Fl. Haw. Is. 531 (1888) [Fig. 65] Poa montícola Gaud, in Freyc., Voy. Uran. Bot. 408 (1830). 21. E. n e o m e x i c a n a Vasey, Contr. U. S. Natl. Herb. 2:542 (1894). 22. E. niihauensis Whitney, Occas. Papers Bernice P. Bishop Mus. 12:6 (1936). [Fig. 66] 23. E. paupera Jedw., Bot. Archiv 5:214 (1924). 24. E. pectinacea (Michx.) Nees, Fl. Afr. Austr. 406 (1841). E. caroliniana (Spreng.) Scribn., Torrey Bot. Club Mem. 5:49 (1894). Poa caroliniana Spreng., Mant. Fl. Hal. 33 (1807). 25. E. pilosa (L.) Beauv., Ess. Agrost. 71, 162, 175 (1812). Poa senegalensis Desv., Opuse. 100 (1831). 26. E. plana Nees, Fl. Afr. Austr. 390 (1841). 27. E. poaeoides Beauv., Ess. Agrost. 162 (1812). 28. E. superba Peyr., Sitzungsb. Akad. Wiss. Math. Naturw. (Wien) 38:584(1860). 29. E. tef (Zucc.) Trotter, Bull. Soc. Bot. Ital. 62 (1918). E. abyssinica (Jacq.) Link, Hort. Berol. 1:192 (1827). Poa tef Zucc., Diss, l'ist. Pianta Panizz. Abiss. (1774). P. abyssinica Jacq., Misc. Austr. 2:364 (1781). 30. E. tenella (L.) Beauv. ex Roem. et Schult., Syst. Veg. 2:576 (1817). E. plumosa (Retz.) Link, Hort. Berol. 1:192 (1827). Poa amabilis L., Sp. PI. 68 (1753). P. tenella L., Sp. PI. 69 (1753). E. amabilis (L.) Wight et Arn. ex Hook, et Arn., Bot. Beechey's Voy. 251 (1838). See Bor (i960) for discussion of nomenclature of E. tenella. 31. E. trichodes (Nutt.)

Wood, Class-book Bot. ed. 1861, 796 (1861).

32. E. unioloides (Retz.) Nees ex Steud., Syn. PI. Glum. 1 ¡264 (1854).

L66

GRASSES

OF

HAWAII

FIG. 64. Eragrostis mauiensis Hitchc. (From A. S. Hitchcock, The Grasses of Hawaii.)

ERAGROSTOIDEAE - ERAGROSTEAE

IÓ7

L68

GRASSES

OF

HAWAII

FIG. 66. Eragrostis niihauensis Whitney, (a) Plant; (b) spikelet. (From Whitney and Hosaka, 1936. Occas. Papers Bernice P. Bishop Mus. 12, No. 5. Reprinted with permission of T h e Bernice P. Bishop Museum.)'

ERAGROSTOIDEAE -

ERAGROSTEAE

169

33. E. variabilis (Gaud.) Steud., Nom. Bot. ed. 2, 1:564 (1840). [Fig. 67] Poa variabilis Gaud, in Freyc., Voy. Uran. Bot. 408 (1830). E. wahowensis Trin., Mém. Acad. Sci. Pétersb. sér. 6, 11412 (1830). E. equitans Trin., Mém. Acad. Sci. Pétersb. sér. 6, 1:413 (1830). E. hawaiiensis Hillebr., Fl. Haw. Is. 530 (1888). E. phleoides Hillebr., Fl. Haw. Is. 530 (1888). E. variabilis var. beta ciliata Hillebr., Fl. Haw. Is. 529 (1888). E. thyrosoidea Hillebr., Fl. Haw. Is. 529 (1888). 34. E. whitneyi Fosb., Occas. Papers Bernice P. Bishop Mus. 15 (3) : 39 (1939)- [Pacific equatorial and leeward Hawaiian Islands.] 35. E. whitneyi Fosb. var. caumii Fosb., Occas. Papers Bernice P. Bishop Mus. 15 (3) ¡41 (1939). [Only westernmost leeward Hawaiian Islands.] Leptochloa Beauv., Ess. Agrost. 71 (1812). SPIKELETS solitary, 1-6-flowered, sessile or very shortly pedicellate, glabrous, not falling entire; rachilla disarticulating above the glumes and between the lemmas, GLUMES persistent, subequal or unequal, membranous, 1-nerved, LEMMA or LEMMAS i, or 5-6, firmly membranous, broadly ovate to ovate elliptical in outline, entire, obtuse or mucronate, 3-nerved, dorsally convex, the two lateral nerves submarginal and usually hairy, PALEA nearly as long as the lemma, hyaline, 2-keeled, the keels ciliate. LODICULES 2, cuneate or oblong, fleshy. STAMENS 3 ; anthers short, OVARY glabrous ; styles distinct, the stigmas plumose, GRAIN subglobose, oblong-obovoid, or triangular, closely enclosed by the lemma and palea, the pericarp very thin and adnate. Annuals or perennials; leaf blade flat; ligule short, ciliate; inflorescence consisting of a number of spikelike racemes or spikes, racemosely or digitately arranged, the spikes or racemes secund.

Leptochloa virgata (L.) Beauv., Ess. Agrost. 71, 161, 166 (1812). Cynosurus virgatus L., Syst. Nat. ed. 10, 2:876 (1759).

1.

Trichoneura Anderss., Vet. Akad. Handl. Stockholm, 148 (1753). Crossotropis Stapf SPIKELETS

3-9.

GLUMES

solitary, shortly pedicelled, not falling entire, FLOWERS persistent, equal, similar, exceeding the lemmas, linear-

GRASSES

OF

HAWAII

FIG. 67. Eragrostis variabilis (Gaud.) Steud. The dense panicle from a specimen named by Hillebrand, E. phleoides. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

ERAGROSTOIDEAE - ERAGROSTEAE

lanceolate, long-acuminate, acute, i-nerved, keeled, scabrid on the keel, membranous, glabrous or glandular on the back, LEMMAS 3-9, similar, decreasing in size upwards, lanceolate or lanceolate-elliptic, notched or shortly 2-lobed, mucronate or shortly awned from the sinus, 3-nerved, keeled, membranous, long-ciliate, subtending a palea and a bisexual flower; awn about 4 times shorter than the body of the lemma, scabrid; the upper lemmas sometimes subtending a male flower and the uppermost lemma empty, PALEAS subequalling their respective lemmas, oblong-linear or lanceolate, 2-keeled, deeply concave on the back, sometimes ciliate on the keel above, LODICULES 2. STAMENS 3. OVARY obovoid, glabrous; styles distinct, plumose above, laterally exserted. GRAIN oblong in outline; embryo about 1/2 the length of the grain. A creeping grass; culms completely hidden by the leaf sheath; leaves crowded on the culms; leaf sheath ribbed, sparsely pilose; blade slightly longer than the sheath, standing at an angle of about 45 degrees to the culm, linear or linear-lanceolate, pungent, closely ribbed, without a distinct midrib, sparsely pilose; inflorescence a contracted panicle; main axis shortly pilose; rachis more or less flat on the back, keeled on the face, scabrid on the margins; spikelets 2-ranked. 1.

Trichoneura elegans Swollen, Amer. Journ. Bot. 19:439 (1932). [Fig. 68]

Tridens Roem. et Schult., Syst. Veg. 2:34 (1817). SPIKELETS several-flowered, the rachilla disarticulating above the glumes and between the florets, GLUMES membranous, often thin, nearly equal in length, the first sometimes narrower, 1-nerved or the second rarely 3- to 5-nerved, acute or acuminate, LEMMAS broad, rounded on the back, the apex from minutely emarginate or toothed to deeply and obtusely lobed; 3-nerved, the lateral nerves near the margins, the midnerve excurrent between the lobes as a minute point or as a short awn, the lateral nerves often excurrent as minute points, all the nerves pubescent below (subglabrous in one species) the lateral ones sometimes conspicuously so throughout, PALEA broad, the two nerves near the margin sometimes villous. Erect, tufted perennials, rarely rhizomatous or stoloniferous, the blades usually flat, the inflorescence an open to contracted capitate panicle. 1.

Tridens flavus (L.) Hitchc., Rhodora 8:210 (1906). [Fig. 69] Trioda flava Smyth, Trans. Kansas Acad. Sci. 25:95 (1913).

FIG. 68. Trichoneura elegans Swallen. Plant, X I ; spikelet and floret, x 10. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

FIG. 69. Tridens flavus (L.) Hitchc. 1, Plant, x 1/4; 2, panicle, X 1/2; 3, spikelet; 4, floret; all x 5. (U.S.D.A. photograph.)

GRASSES OF H A W A I I

i74

Triraphis R. Br., Prodr. Fl. Nov. Holl. 85 (1810). solitary or sometimes in pairs, pedicelled, not falling entire. persistent, subequal, lanceolate or linear, acute or sometimes mucronate, 1 -nerved, membranous to almost hyaline. LEMMAS 5 - 1 2 , much exserted from the glumes, lanceolate or linearoblong, 2-lobed, with the lobes sometimes bristle-like, 3-awned, 3nerved, keeled, membranous, villous, subtending a palea and a bisexual flower, or the uppermost empty; awns scabrid, with the central awn times as long as the body of the lemma; lateral awns usually shorter, PALEAS somewhat shorter than their lemmas, linear or linearoblong, hyaline, LODICULES 2, minute, delicate, OVARY glabrous; styles distinct, plumose above, laterally exserted. GRAIN tightly embraced by the scarcely changed lemma and palea, linear in outline; embryo short. Perennials; culm striate, glabrous; leaf sheath shorter than the internode of the culm, striate or ribbed, glabrous or pilose with tuberculebased hairs; ligule a row of long cilia; blade folded, ribbed, with a distinct midrib, smooth, glabrous; inflorescence a somewhat contracted panicle or, more rarely, compact and spikelike. SPIKELETS

FLOWERS 5 - 1 1 . GLUMES

1.

Triraphis mollis R. Br., Prodr. Fl. Nov. Holl. 85 (1810).

ERAGROSTOIDEAE -

PAPPOPHOREAE

r

7S

PAPPOPHOREAE Kunth, Rev. Gram. 1:82 (1829). Spikelets all alike, 2-3- to many-flowered, with the lower florets bisexual and the upper male or sterile and often much reduced; rachilla disarticulating above the glumes and continuous between the florets. Glumes similar, usually as long as the florets, 3-5-7-nerved, membranous; lemmas similar in texture to the glumes, very broad, rounded on the back, 9- to many-nerved, with the nerves continued out as antrorsely scabrid to ciliate awns; uppermost lemma reduced to a bundle of awns; palea elliptic-obtuse or truncate when flattened, 2-nerved, 2-keeled. Lodicules 2, minute, cuneate, fleshy. Ovary glabrous; styles distinct; stigmas plumose. Grain oblong, more or less compressed; hilum basal, punctiform; embryo long. Starch grains compound. Perennial or annual herbs. Leaf blades usually narrow, often convolute, with mixed festucoid and panicoid anatomy; silica bodies saddleshaped; microhairs with a glandular head upon a long slender stalk. Spikelets arranged in a somewhat narrow bristly panicle. Ligule a ridge of hairs. Chromosomes small; basic number 10. Genus:

Enneapogon

Enneapogon Desv. ex Beauv., Ess. Agrost. 81 (1812). SPIKELETS solitary, sessile or pedicelled, or in pairs with both spikelets pedicelled or 1 subsessile and the other shortly pedicelled, not falling entire, GLUMES persistent, similar, equal or subequal, lanceolate, distinctly 3-5- or sub-7-nerved, sometimes ciliate on the nerves, membranous, frequently pilose; upper glume as long as the lemmas. LEMMAS 3, dissimilar, 7-9-awned, 9-nerved, membranous; awns I J times as long as the body of the lemma, plumose, ciliate, or scaberulous. LOWEST LEMMA oblong or obovate-elliptic, pilose, subtending a palea and bisexual flower, PALEA linear or lanceolate, sometimes ciliate on the keels, hyaline, MIDDLE LEMMA similar to the lowest lemma, but glabrous, subtending a palea and a male flower or subtending a palea only, UPPERMOST LEMMA reduced to a tuft of minute awns, LODICULES 2, minute, fleshy, STAMENS 3. OVARY glabrous; styles distinct, loosely plumose, laterally exserted. GRAIN oblong in outline; embryo large.

176

GRASSES OF H A W A I I

Perennials; culm striate, pubescent or glabrous; leaf sheath ribbed, glabrous or pubescent; ligule ciliate; blade setaceous, rolled or flat, with a distinct midrib, smooth or scabrid, glabrous or pilose; inflorescence a spikelike raceme or panicle or a much contracted panicle; main rachis pubescent or pilose. 1.

Enneapogon brachystachyus (Jaub. et Spach) Stapf in Dyer, Fl. Cap. 7:654 (1900). [Fig. 70] Pappophorum brachystachyum Jaub. et Spach, Ann. Sci. Nat. Bot. ser. 3, 14:365 (1850); et in 111. PI. Or. 4:34 (1851).

FIG. 70. Enneapogon brachystachyus (Jaub. et Spach.) Stapf. (From D. Meredith, The Grasses and Pastures of South Africa.)

178

GRASSES

OF

HAWAII

SPOROBOLEAE Stapf in Dyer, Fl. Cap. 7:315 (1898). Spikelets all alike, bisexual, terete, gaping, 1-flowered, small; rachilla disarticulating above the glumes, very rarely produced beyond the floret. Glumes membranous, nerveless or 1-nerved, usually shorter than the lemmas, equal or unequal, truncate, acute or acuminate, more or less persistent; lemmas 1-3-nerved, membranous, not becoming indurated, usually shining, mostly greyish; palea hyaline, 2-nerved, often split by the developing grain. Lodicules 2, broadly cuneate, glabrous. Stamens 2 or 3. Ovary glabrous; styles 2, free; stigmas plumose. Grain enclosed in a delicate pericarp which swells when wetted and often ejects the seed; hilum small, punctiform, basal; embryo 1/2 as long or as long as the grain; starch grains compound. Annual or perennial herbs. Leaf blades narrow with panicoid anatomy; silica bodies saddle-shaped; microhairs 1-celled occasionally 2-celled, swollen; ligule a line of hairs which are sometimes connate at the base. First leaf blade of the seedling spreading. Inflorescence an open or contracted panicle of usually very small spikelets. Chromosomes small; basic number 9, 10 (12). Genus:

Sporobolus

Sporobolus R. Br., Prodr. Fl. Nov. Holl. 169 (1810). SPIKELETS solitary, sessile, or pedicelled, not falling entire; pedicel usually capillary, swollen at apex, F L O W E R I . GLUMES equal or unequal, persistent or caducous, 1-nerved or nerveless, emucronate, seldom keeled; lower often very small or almost obsolete; upper much shorter than, to as long as, the lemma, lanceolate, sometimes acuminate, acute or obtuse, membranous, L E M M A I , elliptic, obtuse, hyaline or similar in texture to the upper glume, glabrous, PALEA equalling the lemma or a little shorter, 2-nerved, often caniculate between the nerves, obtuse, truncate, emarginate or minutely bilobed, sometimes splitting at the base, L O D I C U L E S delicately hyaline, truncate, S T A M E N S 2-3. O V A R Y often globose; styles free, plumose above, G R A I N either falling entire or often the pericarp remains attached to the rachilla and the seed alone falls through the bursting of the pericarp; pericarp delicate, loose, more or less swelling in water; embryo 1/2 or less than the length of the seed. Annuals or perennials, sometimes with a creeping rootstock, sometimes stoloniferous; culms smooth, glabrous; leaves sometimes all basal; leaf

ERAGROSTOIDEAE -

SPOROBOLEAE

179

sheath ribbed, sometimes ciliate on the upper portion, glabrous, rarely pilose; ligule ciliate, sometimes reduced to a minute rim; blade usually narrow-linear or rolled or filiform or setaceous, more rarely flat, closely veined, with or without a midrib, sometimes ciliate with cartilaginous hairs, rarely scabrid on the margins, glabrous, more rarely pilose; inflorescence usually a lax panicle, sometimes a subraceme, more rarely contracted and spikelike or of spikelike racemes from the main rachis; primary branches of inflorescence erect or horizontal, usually in pairs or whorled on the main rachis, simple or bearing a single spikelet, or branched, sometimes appearing almost spikelike owing to the crowding of the spikelets, sometimes very short so that the spikelets appear as if fascicled on the main rachis. 1.

Sporobolus africanus (Poir.) Robyns et Tourn., Bull. Jard. Bot. Etat. Brux. 25:242 (1955). [Fig. 71] 5. capensis (Willd.) Kunth, Enum. PI. 1:212 (1833). Agrostis spicatus Thunb., Prodr. PI. Cap. 19 (1794). non A. spicata Vahl, Sym. Bot. 1:9 (1790). non A. capensis (L.) Lamk., Encycl. Meth. Bot. 1:58 (1783). A. africana Poir. in Lamk., Encycl. Meth. Bot. Suppl. 1:254 (1810). 2. S. airoides (Torr.) Torr., U. S. Expl. Miss. Pacif. Rept. 7:2i (1856). 3. S. cryptandrus (Torr.) A. Gray, Man. Bot. 576 (1848). 4. S. diander (Retz.) Beauv., Ess. Agrost. 26, 147, 178 (1812). Agrostis diandra Retz., Obs. Bot. 5:19 (1789). 5. S. elongatus R. Br., Prodr. Fl. Nov. Holl. 170 (1810). 6. S. fertilis (Steud.) W. D. Clayton, Kew Bull. 19:291 (1965). Agrostis fertilis Steud., Syn. PI. Glum. 1:170 (1854). S. indicus auctt. mult., non (L.) R. Br. 7. S. fimbriatus Nees, Fl. Afr. Austr. 156 (1841). 8. S. flexuosus (Thurb.) Rydb., Bull. Torrey Bot. Club 32:6oi (1905). 9. S. indicus (L.) R. Br. sensu strictu, Prodr. Fl. Nov. Holl. 170 (1810). [Fig. 72] S. poirettii (Roem. et Schult.) Hitchc., Bartonia 14:32 (1932). S. berteroanus (Trin.) Hitchc. et Chase, Contr. U. S. Natl. Herb. 18:370 (1917). Bor (i960, p. 630) indicates that this S. indicus (L.) R. Br. is a complex made up of several of the above-named species and may include S. elongatus R. Br. and others. 10. S. virginicus (L.J Kunth, Rev. Gram. 1:67 (1829). 11. S. wrightii Munro ex Scribn., Bull. Torrey Bot. Club 9:103 (1882).

i8o

GRASSES

OF

HAWAII

FIG. 71. Sporobolus africanus (Poir.) Robyns et Tourn. Plant, x 1/2; part of panicle, x 3 ; spikelet and grain, x 20. (From O. Degener, Flora Hawaiiensis.)

ERAGROSTOIDEAE

-

SPOROBOLEAE

181

FIG. 72. Sporobolus indicus (L.) R. Br. A, Plant, x 1/2; B, spikelet, x 20; C, glumes, x 20; D, floret, x 20. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

GRASSES

OF

HAWAII

ZOYSIEAE Miq., Fl. Ind. Bat. 3:365 (1857) emend. Stapf in Dyer, Fl. Cap. 7:315 (1898) partim. Spikelets all alike, bisexual, or rarely in groups with some of the spikelets male, 1-flowered; rachilla disarticulating below the floret, not produced beyond the floret; spikelets falling entire, either singly or in groups of 2-5. Glumes equal or the lower suppressed; the lower sometimes with a globose base and a cymbiform limb, keeled, with a cartilaginous wing, the upper coriaceous, rarely keeled and winged; lemma hyaline, much smaller than and enclosed by the glume or glumes; palea broad or narrow, hyaline. Lodicules absent or 2. Stamens 3 or 2; anthers small or long. Ovary glabrous; styles 2; stigmas plumose. Grain free between the lemma and palea; hilum punctiform. Annual or perennial herbs. Leaf blades flat or wiry, rigid, rarely flat, usually subulate or convolute, with panicoid anatomy; silica bodies saddle-shaped; 2-celled microhairs swollen. Spikelets arranged in narrow spikes or subspiciform racemes. Ligules absent or a ciliate rim. First foliage leaf of the seedling spreading. Chromosomes small; basic number 9, 10. Genera:

Hilaria

Tragus

Zoysia

KEY TO THE GENERA OF ZOYSIEAE

1.

Spikelets sessile in groups of three, the center spikelets bisexual, the two lateral male, the group falling entire . . Hilaria p. 182 Spikelets not arranged as above

1. 2.

Glumes of the spikelets with recurved spines or one armed and . Tragus one smooth; spikelets arranged in erect racemes . p. 183

2.

Glumes of the spikelets smooth, compressed, shining; spikelets arranged in rigid, erect spikes Zoysia p. 186

Hilaria H.B.K.,

Nov. Gen. et Sp. 1:116 (1816)

SPIKELETS sessile, in groups of three, the flowers falling from the axis entire, central spikelet (next to the axis) fertile, 1-flowered (occasionally 2-flowered), the two lateral spikelets staminate, 2-flowered (occasionally 3-flowered). GLUMES coriaceous, those of the three spikelets forming

ERAGROSTOIDEAE - ZOYSIEAE

a false involucre, in some species connate at the base, more or less asymmetric, usually bearing an awn on one side from about the middle (extension of the midnerve of the asymmetric glume), LEMMA and PALEA hyaline, about equal in length. Perennials, with stiff solid culms and narrow blades, the groups of spikelets appressed to the axis, in terminal spikes. 1.

Hilaria belangeri (Steud.) Nash, N. Amer. Fl.

17:135

(1912).

I

2.

H. cenchroides H.B.K., Nov. Gen. et Sp. 1:117 ( 8iS)-

3.

H. jamesii (Torr.) Benth., Journ. Linn. Soc. Bot. [Fig- 73]

19:62 (1881).

4.

H. mutica (Buckl.) Benth., Journ. Linn. Soc. Bot.

19:62 (1881).

Tragus Haller, Hist. Stirp. Helvet. nom. cons. Nazia Adans., Fam. PI.

2:203 (1768)

Scop., Intr.

73 (1777)

2 : 3 1 (1763).

SPIKELETS sessile, in clusters of 2 - 5 , all fertile, or often one or more more or less reduced; clusters of spikelets deciduous, SPIKELETS I flowered. GLUMES very dissimilar; lower facing the rachis, minute, hyaline, sometimes suppressed; upper exceeding the lemma, lanceolate, acute, rounded on the back, 3-5-nerved, or -ribbed, membranous, with the ribs or nerves hispid or spine-hooked, LEMMA I, lanceolate or lanceolate-oblong, acute, 1-nerved, glabrous, PALEA as long as lemma, 2-nerved. LODICULES 2, broad, cuneate. STAMENS 3. STYLES distinct, very slender, plumose above terminally exserted. GRAIN enclosed by the lemma and palea, oblong to elliptic in outline, hilum punctiform, basal; embryo about 1/3 the length of the grain. Annuals or perennials, sometimes with creeping stems rooting at the nodes; culm smooth, glabrous; leaf sheath shorter than the internodes of the culm, finely striate, glabrous; ligule a short ciliate rim; blades usually linear-lanceolate, acute, finely and closely nerved, without a distinct midrib, with small cartilaginous spines on margins, glabrous; inflorescence spikelike, terminal, solitary, apparently unbranched; rachis of inflorescence persistent, grooved on the internodes, pubescent.

1.

Tragus berteronianus Schult., Mantissa 2 : 2 0 5 T. occidentalis Nees, Agrost. Bras. 286 (1829).

(1824).

[Fig- 74]

FIG. 73- Hilaria jamesii (Torr.) Benth. Plant, x 1/2; single spike, x 1; A, group of spikelets, two views; B, fertile spikelet; C, staminate spikelet; and D, fertile floret; all x 5. (U.S.D.A. photograph.)

FIG. 74. Tragus berteronianus Schult. A, Plant, x 1 ; B, bur, x 10; C,spikelet, x 10; D, ligule, X 6. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

GRASSES OF H A W A I I

Zoysia Willd., in Neue Sehr. Ges. Naturf. Fr. Berlin 3:440 (1801) nom. cons. Osterdamia Neck., Elem. 3, 218 (1791). SPIKELETS i-flowered, laterally compressed, appressed flatwise against the slender rachis, glabrous, disarticulating below the glumes, FIRST GLUME wanting, SECOND GLUME coriaceous, mucronate, or short-awned, completely infolding the thin lemma and palea, the palea sometimes obsolete. Low perennials, with creeping rhizomes, short, pungently pointed blades, and terminal spikelike racemes, the spikelets on short appressed pedicels.

1.

Zoysia japónica Steud., Syn. PI. Glum. 1:414 (1855).

2.

Z. matrella (L.) Merr., Philipp. Journ. Sei. Bot. 7:230 (1912). [Fig- 75] Agrostis matrella L., Mant. PI. 2:185 (1771). Z. pungens Willd., Neue Sehr. Ges. Naturf. Fr. Berlin 3:44i (1801). Osterdamia matrella (L.) O. Ktze., Rev. Gen. PI. 2:781 (1891).

3.

Z. tenuifolia Willd. ex Trin., Mém. Acad. St. Pétersb. sér. 6, 4:96 (1836).

GRASSES OF

HAWAII

ANDROPOGONEAE Dumort., Obs. Gram. Belg. 84, 141 (1923) as Andropogineae. Spikelets usually in pairs, rarely in threes or solitary, one of each pair or three sessile, the other or both pedicelled; the sessile spikelet most often bisexual, 2-flowered, with the lower floret male or sterile or absent, the upper bisexual or female; the pedicelled spikelet 2-flowered with the lower male or sterile, and the upper male or sterile, very rarely bisexual, or both florets absent or the pedicelled spikelet reduced to the pedicel or the latter reduced or entirely absent. Spikelets deciduous from pedicels at maturity, the sessile often with the adjacent joint of the rachis and pedicel; rachilla not produced beyond the upper floret. Glumes more or less rigid and firmer than the lemma, the lower always as long as or longer than the florets. Lemmas hyaline or membranous, the upper sometimes reduced to the hyaline base of an awn or broader, entire, 2lobed or cleft with a stout geniculate awn from the tip, sinus or cleft. Paleas shorter than the lemma, frequently the lower or both absent. Ovary glabrous; styles 2; stigmas 2, plumose. Grain loosely enclosed by the glumes or lemmas; hilum subbasal, punctiform; embryo large; starch grains simple or compound. Annual or perennial grasses. Leaf blades linear, lanceolate or broad, narrowed, rounded, cordate or rarely sagittate at the base, rarely petioled, with panicoid anatomy; silica cells dumbbell-shaped, or with undulate margins; microhairs filiform, 2-celled. Spikelets panicled, racemose or spicate on tough or fragile rachides. Ligules membranous, a ciliate rim or absent. Chromosomes small; basic numbers 5, 9-15, 17, 19. Genera:

Andropogon

Erianthus

Ischaemum

Sehima

Bothriochloa

Eulalia

Narenga

Sorghastrum

Chrysopogon

Hackelochloa

Polytrias

Sorghum

Cymbopogon

Heteropogon

Saccharum

Spodiopogon

Dichanthium

Hyparrhenia

Schizachyrium

Themeda

Eremochloa

Imperata

Sclerostachya

KEY TO THE GENERA OF ANDROPOGONEAE i.

Spikelets in pairs (rarely threes, Polytrias), members of each pair similar, the sessile and the pedicelled bisexual; the pedicelled very rarely female, rarely the sessile spikelet male and the pedicelled

PANICOIDEAE - ANDROPOGONEAE

spikelet bisexual; joints of the panicle thin, linear or somewhat expanded at the top 2.

Spikelets arranged in more or less ample panicles or compound racemes arranged along a central axis; upper lemma awnless or awned from the entire tip or from between two terminal teeth 3.

Rachis of racemes tough or tardily breaking up; spikelets deciduous from their pedicels Lemmas unawned; spikelets other pedicelled

4. 5.

in pairs, one sessile, the

Panicles narrow, contracted, silvery, glumes very delicate; callus hairs twice as long as the glumes or more Imperata p. 208 Panicles broader or widely spreading; glumes indurated, brown or reddish-brown; callus hairs very short or absent

5.

6.

Spikelets oblong, one sessile the other pedicelled; rachis of racemes tardily breaking up . Narenga p. 210

6.

Spikelets in pairs, ovate or obovate, one short-, one long-pedicelled; rachis of the racemes tough Sclerostachya p. 214 Lemmas awned

4.

3.

7.

All spikelets pedicelled

7.

One spikelet pedicelled, the other sessile

Narenga p. 210

Rachis of the racemes breaking up 8.

Spikelets similar, unawned; culms solid; panicle ample Saccharum p. 212 Spikelets awned

8. 9.

Upper lemma cleft; lower glume rounded on the back or 2-keeled, with 5-9 prominent nerves Spodiopogon p. 220

GRASSES OF

9.

2.

HAWAII

Upper lemma not cleft or shortly 2-toothed; lower glume 2-keeled with infolded margins Erianthus p. 203

Spikelets arranged in a single or several racemes, the latter being digitate or arranged along a very short main axis; spikelets in pairs or threes; rarely pedicelled spikelets reduced to pedicel 10.

Spikelets in pairs; racemes 2 or more; upper glume not awned, or if awned, the awn not more than 3 m m long . Eulalia p. 203

10. Spikelets in threes, two sessile, one pedicelled 1.

Polytrias p. 212

Spikelets in each pair dissimilar, the sessile bisexual, the pedicelled male or sterile, or completely absent (or if more or less similar, then the joints of the raceme and the pedicel thick and swollen) 11. Joints of the rachis and pedicel of the pedicelled spikelet (if not much reduced) swollen, 3-angled, rounded or flattened 12.

Sessile spikelet with a male and a bisexual floret; upper lemma awned, rarely awnless

13.

Raceme solitary, terminal; lower glume of sessile spikelet not transversely grooved Sehima p. 214

13.

Racemes 2 to many; lower glume of sessile spikelet often with nodules on the margins or transversely furrowed Ischaemum p. 208

12.

Sessile spikelet with a bisexual floret only or occasionally with a male floret below; upper lemma unawned

14.

Sessile spikelet spherical; lower glume pitted all over; pedicelled spikelet reduced Hackelochloa p. 203

14.

Sessile spikelet not spherical in shape; lower glume with comblike spines on the margins of the lower glume narrowly or not winged at the apex; pedicelled spikelet much reduced Eremochloa p. 202

PANICOIDEAE -

IQI

ANDROPOGONEAE

11. Joints of the rachis and the pedicel narrow, seldom thickened upwards, occasionally with a translucent longitudinal groove; sessile spikelet usually awned 15.

Inflorescence a terminal raceme .

15.

Inflorescence a panicle

16.

.

.

.

Sehima p. 214

Spikelets in racemes which are not interrupted by spathes or solitary at the ends of the branches; the racemes collected into whorled panicles; joints and pedicels not furrowed

17.

Spikelets solitary at the ends of the branches Sorghastrum

17.

Spikelets in pairs or threes; pedicelled spikelets often

P- 215 more or less reduced 18.

Spikelets dorsally compressed, in panicles of many pairs or in threes of which one is bisexual

.

Sorghum p. 216

18. Spikelets compressed from the sides; lower glume smooth, not tuberculate Chrysopogon p. 197 16. Panicles of racemes which are interrupted by spathes, or the espatheate racemes digitate or in pairs or solitary and terminal, sometimes the joints and/or pedicels with a translucent median furrow 19. Margins of the lower glume of the sessile spikelets sharply infolded, 2-keeled; awn glabrous 20. Upper lemma of the sessile spikelet not cleft, often stipitate and passing into the awn 21. Joints and pedicel with a translucent longitudinal furrow; racemes ascending; glumes sometimes pitted Bothriochloa P- 195 22. Joints and pedicels without a translucent furrow; all spikelets, both sessile and pedicelled, in the racemes more or less alike; glumes herbaceous, greenish Dichanthium p. 201

192

GRASSES OF

HAWAII

20. Upper lemma of the sessile spikelet 2-lobed or 2-cleft, awned in the sinus ¿2. Aromatic grasses; racemes in pairs supported by spathes, often collected into huge panicles; one pair of spikelets in each raceme homogamous, male or neuter, all other pairs heterogamous Cymbopogon p. 199 22. Grasses not aromatic; racemes in pairs, or digitate or solitary 23.

Racemes solitary at the ends of the culms and their branches; joints and pedicles often somewhat stout; pedicelled spikelets male, neuter or suppressed Schizachyrium p. 213

23. Racemes in pairs or more rarely digitate, usually all pairs of spikelets heterogamous and all sessile spikelets bisexual and alike, or the lowest more or less reduced and sterile, but still resembling the upper fertile spikelets . Andropogon P- 193 19.

Margins of the lower glumes inturned and rounded at the sides, at the most keeled upwards; callus mostly sharp and long; awn usually hairy

24. Upper lemma 2-lobed or 2-cleft and awned from the sinus; the awn hairy . . . . Hyparrhenia p. 206 24. Upper lemma not 2-lobed or cleft, stipitate and passing into the awn 25.

Racemes solitary at the ends of the branches, surrounded at the base by an involucre of homogamous spikelets Themeda p. 220

25.

Racemes without an involucre

Heteropogon p. 204

PANICOIDEAE -

ANDROPOGONEAE

193

Andropogon L., Sp. PI. 1045 (1753). SPIKELETS in pairs, one sessile, the other pedicelled, those of each pair differing in sex and mostly also more or less in shape and size, or those of the lowest pair of the lowest raceme homogamous (male or imperfect). SESSILE SPIKELET falling with the adjacent internode of the rachis and the accompanying pedicel; callus short, mostly obtuse, shortly bearded. FLOWER 1, bisexual in the spikelet differing from the pedicelled spikelet. GLUMES equal to subequal, subcoriaceous to membranous; lower flat, concave, or channelled on the back, with sharply inflexed margins at least from the middle upwards, 2-keeled; upper more or less boat-shaped, keeled upwards, 1-3-nerved, sometimes awned. LEMMAS 2, usually ciliate or ciliolate; lower 2-nerved, without a flower or palea, hyaline; upper usually bifid or 2-lobed with an awn (very rarely without) from the sinus or sometimes entire and with the awn (if present) from the tip, upper lemma subtending the bisexual flower and sometimes a palea, awned w i t h the awn 2 - 8 times as long as the lemma, GRAIN narrowly lanceolate to oblong in outline, subterete to plane-convex; embryo about 1/2 as long as the grain, PEDICELLED SPIKELET 1-flowered, sometimes sterile or represented by the pedicel only, LOWER GLUME lanceolate, shortly awned or mucronate, usually several-nerved, often ciliate; upper thinner in texture, lanceolate, acute, sometimes produced into a short awn. LEMMAS (when present) 2, hyaline; lower empty; upper subtending a male flower, or empty, PEDICEL sometimes up to 4 times as long as the adjacent internode of the rachis, linear, linear-oblong, or spathulate, more rarely semiterete, cupshaped at the apex and generally projecting slightly on the inner face of the spikelet, usually long ciliate. Perennials; culm smooth, more rarely distinctly ribbed; leaf sheath ribbed, sometimes ciliate inside at the junction with the blade, glabrous; ligule ciliate (sometimes the cilia very minute); blade linear, flat or longitudinally folded, more rarely filiform, closely ribbed, with a distinct midrib, glabrous; inflorescence of digitate spikelike racemes terminating the culm or, when the culm is branched, the branches of tha culm; racemes usually in pairs, more rarely solitary or in threes, subtended by a spathe, but sometimes remote from the spathe, ciliate to densely villous; internodes of the rachis similar to the pedicel of the pedicelled spikelet.

1.

Andropogon bicornis L., Sp. PI. 1046 (1753).

2.

A. gerardi Vit., Summa PI. 6:16 (1792). [Fig. 76] A. furcatus M u h l . in Willd., Sp. PI. 4:919 (1806).

FIG. 76. Andropogon gerardi Vit. Plant, x 1/4; 2, flowering culm, x 1/2; 3 and 4, spikelets and floret, x 5. (Photograph from U . S . D . A . Yearbook, 1948. Grasses.)

PANICOIDEAE - ANDROPOGONEAE

!95

3.

A. glomeratus (Walt.) B.S.P. Presl, Cat. N.Y.

4.

A. hallii Hack., Sitzungsb. Akad. Wiss. Math. Naturw. (Wien)

67 (1888).

8 9 : 1 2 7 (1884).

5.

A. littoralis Nash, in Britton, Man. Fl. North. United States 69 (1901).

6.

A. virginicus L., Sp. PI. ed. [Fig- 77]

2, 1 4 8 2 ( 1 7 6 3 )

Bothriochloa Kuntze, Rev. Gen. PI.

in part (non L.,

1753).

2:762 (1891).

SPIKELETS in paL's, one sessile, the other pedicelled, differing in sex except sometimes in the lowermost pair which may be homogamous (male or neuter), similar in shape or the pedicelled reduced and smaller. SESSILE SPIKELET (of heterogamous pairs) 1-flowered, deciduous with the adjacent internode of the rachis and the pedicel, GLUMES equal, thinly chartaceous to membranous, lanceolate or elliptic-lanceolate, flat on the back, usually with a circular depression in the middle, ciliate, pilose on the back; upper lanceolate, shortly acuminate, distinctly keeled, glabrous, LEMMAS 2; lower hyaline, empty; upper reduced to an awned stipe, with the awn 3-7 times as long as the stipe, subtending the bisexual flower, PALEA none, or very minute, LODICULES 2, glabrous. STAMENS 3. OVARY ellipsoid; styles plumose above, exserted laterally. GRAIN oblong in outline; embryo about 1/2 the length of the grain. PEDICELLED SPIKELET usually sterile, more rarely 1-flowered, falling entire from the pedicel, LOWER GLUME lanceolate, subacute; upper about 1 ¡2 as long as the lower, hyaline, LEMMAS 2 or o, sometimes 1; the lower sometimes subtending a male flower. Tufted perennials; culms smooth or finely striate, glabrous or sometimes villous at the nodes; leaf sheaths shorter than the internodes of the culm, ribbed, glabrous, sometimes long-ciliate at the junction with blade; ligule membranous, sometimes ciliate; blade narrow-linear, usually with a distinct or prominent midrib, usually covered with bulbous-based hairs, usually scabrid to the touch; inflorescence of simple or branched spikelike racemes arranged digitately or subdigitately at the end of the culm or arranged along a main rachis; internodes of rachis similar to the pedicels of the pedicelled spikelet.

1.

Bothriochloa ambigua 5. T. Blake, Trans. Roy. Soc. South Austral. 6 7 : 4 3 ( 1 9 4 3 ) without descr.; S. T. Blake, Queensl. Univ. Dept. Biol. Papers 2(3) 129 (1944), spelled ambiguna.

196

GRASSES OF H A W A I I

FIG. 77. Andropogon virginicus L. A, Plant, x 1/6; B, racemes, x 1.5; C, spikelet with rachis joint and pedicel, x 5; D, ligule, x 5. (From an original drawing in the files of the Dept. of Agronomy and Soil Science.)

PANICOIDEAE -

ANDROPOGONEAE

197

2.

B. barbinodis (Lag.) Herter, Revist. Sudamer. Bot. 6:135 (1940). [Fig. 78] Andropogon barbinodis Lag., Nov. Gen. et Sp. 3 (1816).

3.

B. caucasia (Trin.) C.E. Hubb., K e w Bull. 1939:101 (1939). Andropogon caucasicus Trin., Mém. Acad. Sci. Pétersb. sér. 6, 2:286 (1832). A. intermedins var. caucasico (Trin.) Hack, in D C . , Monogr. Phan. 6, 486 (1889).

4.

B. insculpta (Hochst.) A. Camus, Ann. Soc. Linn. Lyon, n. s. 76 (i93o):i6s (1931).

5.

B. intermedia (R. Br.) A. Camus, Ann. Soc. Linn. Lyon, n. s. 76 (1930)1164 (1931). Andropogon intermedins R. Br., Prod. Fl. Nov. Holl. 202 (1810).

6.

B. ischaemum (L.) Keng, Contr. Biol. Lab. Sci. Soc. China, Bot. Ser. 10:201 (1936). Andropogon ischaemum L., Sp. PI. 1047 (1753).

7.

B. perforata (Trin. ex Fourn.) Herter, Revist. Sudamer. Bot. 6:135 (1940). Andropogon barbinodis var. perforatus (Trin. ex Fourn.) Gould., Madroño 14:23 (1957). A. perforatus Trin. ex Fourn., Mex. PI. 2:59 (1886).

8.

B. pertusa (L.) A. Camus, Ann. Soc. Linn. Lyon, n. s. 76 (1930): 164 (1931). Andropogon pertusus (L.) Willd., Sp. PI. 4:922 (1806).

9.

B. saccharoides (Sw.) Rydb., Brittonia 1:81 (1931). Andropogon saccharoides Sw., Prodr. Veg. Ind. Occ. 26 (1788). A. argenteus D C . , Cat. Hort. Monsp. 77 (1813).

Chrysopogon Trin., Fund. Agrost. 187 (1820), nom. cons. Rhaphis Lour., Fl. Cochinch. 552 (1790). Centrophorum Trin., Fund. Agrost. 106 (1820). Pollinia Spreng., PI. Pugili. 2:10 (1815) in part. SPIKELETS in threes, one sessile and two pedicelled, the trio falling entire, rarely in pairs (one sessile and the other pedicelled) and both falling with the adjacent internode of the rachis. SESSILE S P I K E L E T I-flowered, GLUMES subequal; lower coriaceous or chartaceous, involute,

198

FIG. 78. Bothriochloa barbinodis (Lag.) Herter. Plant, X 1/2; pair of spikelets, X 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

PANICOIDEAE -

ANDROPOGONEAE

199

with a rounded back, or complicate and more or less keeled upwards; upper boat-shaped, more or less keeled, sometimes awned; awn, when present, as long as or slightly longer than the glume, LEMMAS 2, hyaline; lower 2-nerved, empty; upper linear, entire, 2-toothed or 2-lobed, awned, subtending a palea and a bisexual flower, PALEA narrow, delicately hyaline, LODICULES 2, glabrous, STAMENS 3. OVARY ellipsoid; style plumose above, laterally exserted low down, GRAIN linear in outline; embryo 1/2 as long as the grain, PEDICELLED SPIKELET 1-flowered or sterile. LOWER GLUME lanceolate, awned or awnless, when awned the awn 1/2 as long as glume, chartaceous or membranous; upper somewhat thinner than the lower, mucronate. LEMMAS 2, hyaline; lower empty; upper long-ciliate, subtending a palea and a male flower or flower absent. PALEA linear, 1/3 as long as its lemma, hyaline. Mostly perennials; culm finely striate or smooth, glabrous; leaf sheath ribbed, glabrous; ligule membranous, sometimes ciliate; blade longitudinally folded, ribbed, sometimes scabrid on the keel and margins, glabrous or occasionally with long stiff bulbous-based hairs on the margins; inflorescence a panicle, consisting of a number of whorled capillary primary branches from the main rachis, with each branch bearing 3 spikelets at its apex. 1.

Chrysopogon aciculatus (Retz.) Trin., Fund. Agrost. 188 (1820).

[Fig. 79] Andropogon aciculatus Retz., Obs. Bot. 5:22 (1789). Rhaphis aciculatus (Retz.) Desv., Opusc. 69 (1831). R. trivialis Lour., Fl. Cochinch. 553 (1790). Chrysopogon subulatus Miq., Fl. Ind. Bot. 3:491 (1857). Andropogon subulatus Presl, Rel. Haenk. 1:341 (1828). 2.

Chrysopogon fulvus (Spreng.) Chiov., Fl. Somala. 327 (1929). Pollinia fulva (R. Br.) Benth., PI. Pugill. 2:10 (1815). C. montanus Trin. ex Spreng., Neue Entdeck. 2:93 (1821). C. monticola (Roem. et Schult.) Haines, Indian For. 40:495 (1914).

Cymbopogon Spreng., PI. Pugill. 2:14 (1815). SPIKELETS in pairs, one sessile, the other pedicelled, those of each pair usually differing in sex and more or less in shape; lowermost pair of spikelets of the lower or of both racemes homogamous (usually male). SESSILE SPIKELET of the heterogamous pair falling with the adjacent internode of the rachis and the accompanying pedicel of the pedicelled spikelet. FLOWER I . GLUMES equal or subequal, more or less chartaceous;

200

GRASSES OF H A W A I I

F i g . 79. Chrysopogon aciculatus (Retz.) Trin. Plant, X 1/2; leaf, x 2; male perfect florets, X 5; spikelet, X 5. (From O. Degener, Flora Hawaiiensis.)

florets,

PANICOIDEAE -

ANDROPOGONEAE

201

lower almost flat or slightly depressed, or narrowly grooved on the back, 2-keeled, with the margins sharply inflexed at least from the middle upwards; upper more or less boat-shaped, keeled upwards, usually 1-nerved, LEMMAS 2, ciliate or ciliolate; lower entire, 2-nerved, empty; upper bifid or 2-lobed, hyaline, awned, rarely firmer and almost stipelike below the insertion of the awn, subtending the bisexual flower; column of awn, if any, smooth, PALEA none, LODICULES 2, minute, glabrous. STAMENS 3. OVARY subglobose; styles plumose above, laterally exserted. GRAIN oblong in outline; embryo about 1/2 the length of the grain. PEDICELLED SPIKELET usually 1-flowered, GLUMES muticous; lower chartaceous to subchartaceous; upper thinner, LEMMAS 2; lower hyaline, 2-nerved, empty; upper sometimes subtending a male flower, PEDICEL linear, flat on the face, rounded on the back, usually obliquely cup-shaped at the apex, villous. Perennials, often robust; culms simple or branched, smooth, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous or very rarely pilose at the junction with the blade; ligule membranous or scarious; blade linear, tapering to a fine point, flat, longitudinally folded, convolute or almost filiform, with the midrib distinct and the lateral veins fine or distinct, sometimes scabrid to the touch, glabrous; inflorescence usually in pairs of spikelike racemes subtended by a common spathe, the whole either solitary and terminal on the culm or on the branches of the culm; internodes of the rachis similar to the pedicels of the pedicelled spikelet. 1.

Cymbopogon citratus (DC.) Stapf, Kew Bull. 1906:322 (1906). Andropogon citratus DC., Cat. Hort. Monsp. 78 (1813).

2.

C. nardus (L.) Rendle, Cat. Afr. PI. Welw. 2:155 (1899).

3.

C. refractus (R. Br.) A. Camus, Rev. Bot. Appl. 1:279 ( I 9 2 1 )-

4.

C. schoenanthus (L.) Spreng., PI. Pugill. 2:15 (1815).

Dichanthium Willemet in Usteri, Neue Ann. Bot. 12:11 (1796). SPIKELETS in pairs, one sessile and usually awned, the other pedicelled and awnless. SESSILE SPIKELET 1-flowered or those at the base of the raceme awnless, sterile or with a male flower, falling with the adjacent internode of the rachis and pedicel of the pedicelled spikelet. LOWER GLUME lanceolate or elliptic-oblong, almost truncate, membranous, pilose; upper somewhat thinner in texture, keeled, ciliate. LEMMAS 2 ; lower lanceolate, subhyaline, empty; upper reduced to an awned stipe,

202

GRASSES OF H A W A I I

with the awn 4-6 times as long as the stipe, subtending a bisexual flower. LODICULES 2, minute, glabrous, STAMENS 3. OVARY ellipsoid; style plumose above, exserted at or above the middle or near the tips, GRAIN oblong in outline; embryo rather more than 1/2 as long as the grain. PEDICELLED SPIKELET i-flowered or sterile, falling entire from the pedicel, LOWER GLUME oblong, truncate, pilose; upper ciliate above, membranous, LEMMA I or absent, if present, then hyaline, empty or subtending a male flower. Perennials; culm smooth or finely striate, villous at the nodes; leaf sheath shorter than the internode of the culm, strongly ribbed, glabrous, sometimes pilose within at the junction with the blade; ligule membranous, sometimes with long hairs from the back; blade narrow-linear with a distinct or prominent midrib, sometimes scabrid to the touch, glabrous or sparsely covered with bulbous-based hairs; inflorescence of 3-9 spikelike racemes arranged digitately or subdigitately at the end of the culm or arranged up a main rachis. 1.

Dichanthium annulatum (Forsk.) Stapf in Prain, Fl. Trop. Afr. 9:178 (1917). Andropogon annulatus Forsk., Fl. Aegypt.-Arab. 173 (1775). A. ischaemum Roxb. ex Wight, Cat. (1834) nomen.

2.

D. aristatum (Poir.) C.E. Hubb., Kew Bull. 1939:654 (1939). Andropogon nodosus (Willem.) Nash, N. Amer. Fl. 1 7 : 1 2 2 (1912). A. caricosus L. subsp. mollicomus Hack. var. mollicomus (Kunth) Hack, in DC., Monogr. Phan. 6:569 (1889). D. nodosum Willm. in Usteri, Neue Ann. Bot. 1 8 : 1 1 (1796).

3.

D. sericeum (R. Br.) A. Camus, Bull. Mus. Hist. Natl. 27:549 (1921). Andropogon sericeus R. Br., Prodr. Fl. Nov. Holl. 1:201 (1810).

Eremochloa Btise in Miquel, PI. Junghn. 1:357 ( 1 §54)One species recorded in Hawaii, described as follows: 1.

Eremochloa ophiuroides (Munro) Hack, in DC., Monogr. Phan. 6:261 (1889).

Low perennial, creeping by thick short-noded leafy stolons; racemes spikelike, smooth, subcylindric, terminal and axillary on slender pedicels, 2 to 6 cm long; rachis flat, not thickened, the first glume of sessile spikelet winged at summit.

PANICOIDEAE -

ANDROPOGONEAE

203

Erianthus Michx., Fl. Bor. Amer. 1:54 (1803). SPIKELETS all alike, in pairs along a slender axis, one sessile, the other pedicelled, the rachis disarticulating below the spikelets, the rachis joint and pedicel falling attached to the sessile spikelet. GLUMES coriaceous, equal, usually copiously clothed, at least at the base, with long silky spreading hairs, STERILE LEMMA hyaline, FERTILE LEMMA hyaline, the midnerve extending into a slender awn. PALEA small, hyaline. Perennial reedlike grasses, with elongate flat blades and terminal, oblong, usually silky panicles.

1.

Erianthus r a v e n n a e (L.) Beauv., Ess. Agrost. 14, 162, 177(1812).

E u l a l i a Kunth, Rev. Gram. 160 (1829). SPIKELETS alike or nearly so, one pedicelled, the other sessile, hairy or glabrous, FLOWERS 1-2; when 2, the lower male and the upper bisexual. GLUMES equal; lower membranous, muticous, often truncate, dorsally flattened or concave or longitudinally grooved, more or less 2-keeled, with inflexed margins; upper often narrower, keeled, acute, sometimes with a fine straight mucro. LEMMAS usually 2; lower hyaline, muticous, sometimes reduced or quite suppressed, sterile or subtending a male flower; upper hyaline, 2-lobed, with a fine or rather stout or more or less kneed awn from the sinus, subtending a bisexual flower, PALEA small, hyaline or lacking, STAMENS 3. STYLES distinct; stigmas plumose. GRAIN oblong in outline; hilum punctiform; embryo rather large. Annuals or perennials; culm faintly ribbed, hairy below the nodes; leaf sheath ribbed, tuberculed, hairy; blade with a thin midrib and obscure lateral veins, villous; internodes of rachis similar to the pedicels, articulated below the spikelets; inflorescence a terminal panicle; branches spikelike, simple, fascicled or digitately arranged, with a readily disarticulating rachis.

1.

Eulalia fulva (R. Br.) O. Ktze. Rev. Gen. PI. 2:775 ( i 8 9 0 Pollinia fulva (R. Br.) Benth., Fl. Austral. 7:526 (1778), non Spreng. (1815).

H a c k e l o c h l o a O. Ktze., Rev. Gen. PI. 2:776 (1891). SPIKELETS in pairs, one sessile, the other pedicelled, in the notches of an articulated secund rachis, the lower containing a bisexual floret, the upper male or sterile, SESSILE SPIKELET: GLUMES 2, the lower globular or subglobular, deeply concave, very obtuse, thickly crustaceous,

204

GRASSES

OF

HAWAII

rugose on the back or conspicuously pitted or tuberculed, abaxial; upper glume ovate-lanceolate, smaller than the lower glume, thin but rigid and more or less chartaceous, 7-nerved, immersed in the rachis and filling the opening, L E M M A S 2, the lower hyaline, small, with or without a palea, neuter; upper lemma thin and hyaline, much shorter than the glumes, subtending a bisexual floret, PALEA I , corresponding to the upper lemma, often minute, L O D I C U L E S 2, broadly cuneate. S T A M E N S 3, the anthers usually very small, STYLES distinct, the stigmas short, laterally exserted. G R A I N subglobular, included in the hardened lower glume, P E D I C E L L E D SPIKELET usually reduced to two lanceolate or ovate-lanceolate glumes, or sometimes with a lemma subtending a male floret. Erect annuals; culm much branched; leaf blade flat, more or less cordate at the base; ligule membranous, soon lacerating. Inflorescence of axillary and terminal spikelike racemes subtended by spathes, in a leafy panicle, the rachis readily disarticulating. 1.

Hackelochloa granulans (L.) (1891). [Fig. 80]

O. Ktze.,

Rev. Gen. PI. 2:776

Heteropogon POT., Syn. PI. 2:533 (1807). SPIKELETS in pairs, one sessile, the other pedicelled; spikelets on lower portion of racemes (one to many pairs) alike in sex and shape; spikelets on upper portion of raceme differing in sex and shape; spikelets of homogamous pairs male or neuter, long persistent, SESSILE SPIKELET of heterogamous pair 1-flowered, falling with the adjacent internode of the rachis; callus long, pungent, densely bearded upwards, L O W E R G L U M E linear oblong, obtuse or truncate, flat on the back, with reflexed margins, pilose; upper linear, acute, prominently keeled, L E M M A S 2, hyaline; lower empty; upper stipitiform from a very slender hyaline base, passing into a usually stout kneed awn about 6-10 times as long as the lemma, subtending a female or bisexual flower, PALEA none. L O D I C U L E S large or more or less reduced, STAMENS 3. O V A R Y oblong in outline; style plumose low down, exserted terminally or laterally, G R A I N more or less linear in outline; embryo somewhat exceeding the middle of the grain, PEDICELLED SPIKELET often somewhat twisted, usually 1 flowered; lower glume lanceolate, acuminate, acute, flat on the back, winged on one side; upper thinner in texture, lanceolate, acuminate, acute, keeled, L E M M A S 2, hyaline, 1-nerved, well-developed, or more or less reduced; upper empty; lower sometimes subtending a male flower. PEDICEL linear, tapering to the base, glabrous.

PANICOIDEAE -

ANDROPOGONEAE

205

FIG. 80. HackelocMoa granulans (L.) O. Ktze. (From D. Meredith, T h e Grasses and Pastures of South Africa.)

206

GRASSES

OF

HAWAII

Perennials; culm smooth, glabrous; leaf sheath generally shorter than the internode of the culm, striate, glabrous; ligule very short, ciliate; blade narrow-linear, flat or longitudinally folded, with or without a distinct midrib, with fine lateral veins, smooth or scabrid, glabrous or sometimes sparsely hairy; inflorescence a solitary terminal spikelike raceme, with the awns very long and usually twisted together. i.

Heteropogon contortus (L.) Beauv. ex Roem. et Schult., Veg. 2:836(1817). [Fig. 81] Andropogon contortus L . , Sp. PI. 1045 (1753).

Syst.

Hyparrhenia Anderss. in Nov. Act. Soc. Sci. Upsal. ser. 3, 2:254 (1856) nomen: ex Fourn., M e x . PI. Gram. 51, 67 (1886) descr. SPIKELETS in pairs, one sessile, the other pedicelled, those of each pair differing in sex and more or less in shape, or those of the lowest pair or pairs homogamous, sometimes male, SESSILE S P I K E L E T of the heterogamous pair falling with the adjacent internode of the rachis and the accompanying pedicel of the pedicelled spikelet. F L O W E R I . GLUMES equal, more or less chartaceous; lower lanceolate, truncate or acute, sometimes 2-toothed, rarely produced into 2 short awns, flat or rounded on the back, ciliate, usually pilose; upper narrowly and shallowly boatshaped, rounded on the back below, very rarely awned, finely keeled toward the tip, 3-nerved. L E M M A S 2, ciliate or ciliolate, one or both glabrous; lower hyaline, 2-nerved, empty; upper stipitiform, delicate at the base, gradually hardened above, 2-toothed, awned from the sinus and with the awn 3 - 1 2 times as long as the lemma, subtending the bisexual flower; awn distinctly kneed, more or less hirtellous or subplumose. P A L E A none, rarely a small hyaline scale, L O D I C U L E S 2. S T A M E N S 3. O V A R Y more or less ovoid; styles plumose above, laterally exserted. G R A I N oblong in outline; embryo about 1/2 as long as the grain. PEDICELLED SPIKELET 1-flowered or sterile, falling entire from the pedicel. G L U M E S similar to the glumes of the sessile spikelet. L E M M A S usually both developed, rarely 1 or o ; lower lemma empty; upper lemma subtending a male flower, more rarely a bisexual flower, or empty, PEDICEL linear, more or less obliquely truncate and sometimes obscurely auricled at the apex, long-ciliate.

Perennials, slender or very robust; culm branched, smooth, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous, sometimes pilose within at the junction with the blade; ligule scarious; blade usually narrow-linear, more rarely short and linear-lanceolate,

PANICOIDEAE -

ANDROPOGONEAE

207

FIG. 8 1 . Heteropogon contortus (L.) Beauv. Plant, x 1/6. 1 , Ligule, x 1 ; 2, detached sessile spikelet, X 4; 3 and 4, its lower glume, front and back view, x 5; 5, its upper glume, X 5; 6, grain, showing hilum, X 5 5 7 , pedicelled spikelet, X 5; 8, its upper glume, x 5. (From N. L . Bor, The Grasses of Burma, Ceylon, India and Pakistan.)

208

GRASSES OF H A W A I I

with midrib prominent beneath and sometimes almost 1/3 the width of the blade, with the lateral veins close and usually distinct, often scabrid to the touch; inflorescence in pairs of spikelike racemes subtended by a spathe; two or more pairs of such racemes subtended by a common spathe and the whole forming a spatheate panicle; pairs of racemes terminating long branches of the main rachis and exserted from the spathes, or short or almost enclosed by the spathes, sometimes racemes very short and not visibly paired; spathes green or reddish-brown, ribbed, glabrous; internodes of rachis similar to the pedicel of the pedicelled spikelet. 1.

Hyparrhenia hirta (L.) (1919).

Stapf in Prain, Fl. Trop. Afr. 9:315

2.

H. rufa (Nees) Stapf in Prain, Fl. Trop. Afr. 9:304 (1919). [Fig. 82] Trachypogon rufus Nees, Agrost. Bras. 345 (1829). Andropogon rufus (Nees) Kunth, Rev. Gram. 1, Suppl. 39 (1830); Enum. PI. 1:492 (1833).

Imperata Cyr., PI. Rar. Neap. 2:26 (1792). SPIKELETS all alike, generally in pairs and unequally pedicelled, disarticulating from the pedicels, FLOWER usually solitary, sometimes a lower male flower present, GLUMES subequal, membranous, 3-9-nerved, rarely nerveless, enveloped by very long silky hairs from their bases and from the obscure callus, LEMMAS 2, generally much smaller than the glumes, hyaline, awnless; upper smaller than the lower, subtending the bisexual flower; lower empty, rarely subtending a male flower, PALEA hyaline, broad, nerveless, LODICULES none, STAMENS 1-2. STYLES connate below; stigmas linear, exserted from the top of the spikelet. GRAIN oblong in outline; embryo 1/2 the length of the grain or more. Perennials; basal leaves crowded and, like those of the innovation shoots, long; inflorescence a spikelike or narrowly ovate silvery-silky panicle, with the branches continuous and bearing the pedicelled spikelets.

1.

Imperata cylindrica (L.) (1812).

Beauv., Ess. Agrost. 165, 166, 177

Ischaemum L., Sp. PI. 1049 (1753). SPIKELETS in pairs, both alike or differing only in sex, or more or less heteromorphous with one pedicelled and the other sessile, SESSILE SPIKELETS dorsally compressed, often rather broad, deciduous with the

2IO

GRASSES OF

HAWAII

adjacent internode of the rachis and pedicel, FLOWERS 1-2, when 2 then the lower male and the upper usually bisexual, GLUMES equal or subequal; lower dorsally flattened or somewhat convex, usually coriaceous below, chartaceous and markedly nerved upwards, rarely concave and more or less chartaceous throughout, more or less 2-keeled, with inflexed margins; upper boat-shaped, keeled at least above, sometimes awned. LEMMAS 2, rigidly membranous to hyaline, lower muticous, empty or subtending a male flower; upper usually bifid and awned from the sinus, rarely mucronate or muticous, subtending a bisexual flower, PALEAS more or less equalling their lemmas, hyaline, LODICULES 2, cuneate. STAMENS 3, sometimes smaller or rudimentary in the fertile flower, STIGMAS linearoblong, laterally exserted. GRAIN oblong or lanceolate in outline, dorsally compressed; embryo reaching to the middle of the grain, PEDICELLED SPIKELET either like the sessile in sex and form or male or more or less reduced, sometimes apparently laterally compressed with a median keel owing to the more or less complete suppression of one side, falling entire from its pedicel. Generally perennials; culm smooth or ribbed, glabrous or sometimes hairy at the nodes; leaf sheath ribbed, glabrous or pilose; blade flat with an evident but not very prominent midrib and with distinct lateral ribs, glabrous or pilose; ligule membranous; inflorescence of compressed binate, digitate, or fascicled racemes; rachis readily disarticulating; internodes flattened or subconcave on the inner side, often stout; pedicels similar to the internodes. 1.

2. 3.

Ischaemum byrone (Trin.) Hitchc., Mem. Bernice P. Bishop Mus. 8(3): 213 (1922). [Fig. 83] Spodiopogon byronis Trin., Mem. Acad. Sci. Petersb. ser. 6, 2:301 (1832)I. byronis Sprague, Index Kew Suppl. 7:128 (1929). I. lutescens Hack, in DC., Monogr. Phan. 6:221 (1889). I. digitatum Brogn. in Duperrey, Bot. Voy. Coquille 22:70 (1831). I. indicum (Houtt.) Merr., Journ. Arn. Arbor. 19:320 (1938). I. ciliare Retz., Obs. Bot. 6:36 (1791).

Narenga Bor, Indian For. 66:227 ( I 94 0 )SPIKELETS all alike, in pairs, one sessile the other pedicelled on the articulate fragile rachis of panicled racemes, the pedicelled spikelets falling from their pedicels, the sessile together with the contiguous joint

PANICOIDEAE -

ANDROPOGONEAE

211

FIG. 83. Ischaemum byrone (Trin.) Hitchc. Plant, x 1/2; sessile spikelet, x 3; floret, X 5; pedicelled spikelet, X 5. (From O. Degener, Flora Hawaiiensis.)

212

GRASSES OF HAWAII

of the rachis and pedicel, FLORETS 2 ; the lower reduced to an empty lemma; the upper bisexual, GLUMES coriaceous, brown, shining; lower glume flat on the back, margins laxly inflexed, short haired on the margins; upper keeled, membranous at the top. LEMMAS hyaline; upper truncate, awnless, paleate. LODICULES 2. STAMENS 3. STIGMAS laterally exserted. Tall perennial grasses with long flat scabrid leaves. Culms densely bearded at the nodes. Panicle narrow-dense; hairs shorter than the spikelets. See Bor (i960), p. 197 for a discussion of Narenga. 1.

Narenga porphyrocoma (Hance) Bor, Indian For. 66:227 (1940). Saccharum narenga Wall., Cat. no. 8856. Eriochrysis porphyrocoma H. F. Hance, Journ. Bot. 14:294 (1876). Saccharum porphyrocomum (Hance) Hack, in DC., Monogr. Phan. 6:120 (1889).

Polytrias Hack, in Engler and Prantl, Natiirl. Pflanzenf. 2:2, 24(1887). SPIKELETS all alike or the pedicelled reduced, in threes at the nodes of the rachis of the terminal inflorescence, two sessile, one pedicelled, usually covered with golden hairs, SESSILE SPIKELETS: lower glume oblong, concave on the back, the upper somewhat keeled; FLORETS 2, the lower male or empty, the upper bisexual; lower lemma hyaline, truncate, the upper bifid at the tip and furnished with a geniculate awn. PEDICELLED SPIKELET sometimes male or empty.

1.

Polytrias amaura (Biise) O. Ktze., Rev. Gen. PI. 2:788 (1891). P. praemorsa (Nees) Hack, in Engler and Prantl, Natiirl. Pflanzenf. 2:2, 24 (1887).

Saccharum L., Sp. PI. 54 (1753). SPIKELETS in pairs, one sessile, the other pedicelled, all alike, usually awnless, rarely awned, each with a ring of long, silky hairs from the callus, FLOWERS 2, upper bisexual, lower reduced to a lemma, GLUMES somewhat hardened, LEMMAS thinner than the glumes; upper sometimes absent, when present, awnless or more rarely with a terminal, usually straight awn. PALEA small. Robust erect, tufted perennials. Leaf blades various, sometimes reduced to a stout midrib. Inflorescence a panicle; usually large, showy, silkily hairy and plumelike.

PANICOIDEAE -

1.

ANDROPOGONEAE

Saccharum arundinaceum Retz.,

2I3

Obs. Bot. 4:14 (1786).

Erianthus arundinacetis (Retz.) Jeswiet, Arch. Suikerind. N e d . - I n d . 33 (Meded.): 399 (1925). E. maximum Brongn. in Duperrey, Bot. Voy. Coquille 2:97 (1831). 2.

S. barberi Jeswiet, Arch. Suikerind. Ned.-Ind. 33 (Meded.): 404

3.

S. bengalense Retz., Obs. Bot. 5:16 (1789).

(!925)S. arundinaceum Hook, f., Fl. Brit. Ind. 7 : 1 1 9 (1896) in part, non Retz. 4.

S. officinarum L., Sp. PI. 54 (1753). S. edule Hassk. Flora 25, Beibl 3 (1842).

5.

S. robustum Brandes et Jeswiet ex Grassl., Journ. Arb. 27:234 (1946).

6.

S. sinense Roxb., PI. Coromand. 3 (1819). [5. barberi is usually included in S. sinense. S. sinense is a rather heterogeneous group and represents a recent introgression of 5 . spontaneum with 5 . officinarum.]

7.

S. spontaneum L., Mant. Alt. 2:183 (1771).

Schizachyrium Nees, Agrost. Bras. 331 (1829). SP 1 RELETS in pairs, one sessile, the other pedicelled, differing in sex and more or less in shape and size, SESSILE SPIKELET 1-flowered, falling with the adjacent internode of the rachis and accompanying pedicelled spikelet. GLUMES equal or subequal; lower more or less convex or flat on the back at least from the middle upwards, with sharply inflexed and mostly narrow margins, 2-keeled, with the keels running out into teeth or mucros, chartaceous to subcoriaceous; upper thinner to membranous, narrowly boat-shaped to dorsally flattened, keeled (at least upwards), 1-3 nerved, with the delicate margins ciliolate. LEMMAS 2, hyaline, ciliolate; lower membranous downwards, empty; upper usually bifid or 2-toothed, more rarely bipartite or entire, with an awn continuous with or from the sinus of the lobes, subtending a bisexual flower, PALEA none, or represented by a microscopic hyaline scale, LODICULES 2, glabrous. STAMENS 3, rarely 2. OVARY subglobose; style plumose above, laterally exserted low down, GRAIN narrowly linear in outline or tapering upwards, subterete. PEDICELLED SPIKELET 1-flowered or sterile, LOWER GLUME

214

GRASSES OF

HAWAII

subcoriaceous, awned, with the awn about as long as the glume; upper membranous, acuminate, LEMMAS 2, hyaline, lanceolate, acuminate; both empty or the upper sometimes subtending a male flower, PEDICEL linear-oblong, flattened, slightly concave on the face, long-ciliate. Perennials; culms smooth, glabrous; leaf sheath ribbed, glabrous, sometimes ciliate at the junction with the blade; ligule membranous; blade narrow linear, longitudinally folded, closely ribbed, with the midrib forming a keel; inflorescence a lax panicle formed of long cylindric spikelike racemes, solitary and terminating the branches of the culm, with a spathe at the base of the pedicel of each raceme; internodes of rachis similar to the pedicel of the pedicelled spikelets. 1.

Schizachyrium scoparium (Michx.)

Nash in Small, Fl. South-

east United States 59 (1903). Andropogon scoparius Michx., Fl. Bor. Amer. 1:57 (1803). Sclerostachya (Anderss.)

A. Camus in Lecomte, Fl. Gen. de l ' l n d o -

Chine 7:243 (1922). Sclerostachyum Andropogon

Stapf in Ridley, Fl. Malay Penin. 5:186, 194 (1925).

subgenus

Sclerostachyus

Anderss. ex Hack, in

DC.,

Monogr. Phan. 6:121 (1889). SPIKELETS in pairs, one long-pedicelled, the other short-pedicelled on the tough secondary filiform branches of an erect, terminal panicle, oval-oblong or slightly obovate-oblong; obtuse, deciduous from the pedicels, SHORT-PEDICELLED SPIKELETS; glumes coriaceous, brown, shining, the lower flat on the back, with incurved margins, the upper keeled acute, with margins ciliate above, FLORETS 2, the lower male or empty; the lemma hyaline, ovate-oblong, ciliate at the top, paleate; the upper bisexual; lemma hyaline, awnless; PALEA as long as its lemma. LONG-PEDICELLED SPIKELET similar with the short pedicelled, sometimes male, female or empty.

1.

Sclerostachya fusca (Roxb.) A. Camus in Lecomte, Fl. Gen. de l'lndo-Chine 7:243 (1922). Saccharum fuscum Roxb., Fl. Ind. 1:241 (1820).

Sehima Forsk., Fl. Aegypt.-Arab. 178 (1775). SPIKELETS in pairs, one sessile, the other pedicelled, those of each pair differing in sex and (in the African species) also much in shape, SESSILE

PANICOIDEAE -

ANDROPOGONEAE

215

deciduous together with the adjacent internode of the rachis and the pedicel, FLOWERS 2 ; lower male; upper bisexual, GLUMES equal or subequal; lower deeply grooved, rarely flat, 2-toothed or 2-mucronate, more or less chartaceous, acutely 2-keeled upwards, with inflexed margins, with the keels winged; upper boat-shaped, keeled upwards, with a bristle-like awn. LEMMAS 2, hyaline; lower entire, muticous, subtending the male flower; upper bifid, awned from the sinus, subtending the bisexual flower, PALEAS more or less equalling their lemmas, hyaline. LODICULES 2, cuneate. STAMENS 3. STIGMAS linear oblong, laterally exserted. GRAIN oblong in outline, obtusely 3-angled; embryo reaching to middle of the grain, PEDICELLED SPIKELETS flat, with (in African species) a strongly nerved or ribbed lower glume, tardily separating from its pedicel; pedicel sublinear and parallel with the internode of the rachis. LEMMAS 2; the upper sometimes subtending a male flower; the lower sterile. Perennials; culm smooth, glabrous; leaf sheath ribbed, glabrous; blade flat, narrowed into a setaceous portion, closely ribbed, without an evident midrib, glabrous, scabrid to the touch; ligule a fringe of long hairs; inflorescence a spikelike raceme, usually gently curved; internodes of the rachis similar to the pedicels. SPIKELETS

1.

Sehima nervosum (Rottl.) 1917).

Stapf in Prain, Fl. Trop. Afr. 9:36

Ischaemum laxum R. Br., Prod. Fl. Nov. Holl. 205 (1810). Sorghastrum Nash in Britton, Man. Fl. North. United States 71 (1901). SPIKELETS solitary, sessile, accompanied by a pedicel without a spikelet, 1-flowered, LOWER GLUME lanceolate, somewhat flat on the back, with inturned margins, truncate, 9-nerved, pilose on the back; upper lanceolate, convex on the back, concave on the face, with inturned ciliate margins, 5-nerved, glabrous on the back, LEMMAS 2, hyaline; lower more or less broadly linear, 2-keeled, with the margins on the upper part inflexed and ciliate, 2-nerved, empty; upper linear, 2-lobed, convex on the back, with the margins on the upper part inturned and ciliate, produced into a long awn from the sinus of the lobes, subtending a palea and a bisexual flower, PALEA delicate, hyaline, lanceolate, slightly longer than the ovary, LODICULES 2, cuneate. STAMENS 3. OVARY ellipsoid; styles plumose. T h e ACCOMPANYING NAKED PEDICEL about 3/4 as long as the sessile spikelet, linear, long-pilose.

2l6

GRASSES

OF

HAWAII

Perennials; culm faintly striate, glabrous; leaf sheath finely ribbed, glabrous; ligule produced into 2 lobes; blade rolled, appearing terete, finely ribbed, glabrous; inflorescence a lax panicle; internode of rachis about 3/4 as long as the sessile spike, linear with a cup and ball articulation, long-ciliate. 1.

Sorghastrum nutans (L.) Nash in Small, Fl. Southeast United States 66 (1903). [Fig. 84]

Sorghum Moench, Meth. PI. 207 (1794). Sorgum Adans., Fam. PI. 2:38, 606 (1763) nom. illegit. SPIKELETS in pairs, one sessile, the other pedicelled, those of each pair differing in sex and shape, SESSILE SPIKELET falling with the adjacent internode of the rachis and the accompanying pedicelled spikelet or at least with its pedicel, F L O W E R I . GLUMES equal, coriaceous (at least when mature), rarely permanently chartaceous, muticous; lower with a broad flattened or convex back, with the margins narrowly inflexed near the tips; upper boat-shaped, keeled upwards, with narrow hyaline margins, ciliolate above, sometimes mucronate. L E M M A S 2, hyaline, ciliate; lower 2-nerved or nerveless, empty; upper oblong to ovate, 2-toothed or 2-lobed, with a perfect or variously reduced awn or mucro arising from the sinus, rarely entire and mucronate or muticous, 1-2-nerved, subtending the bisexual flower, L O D I C U L E S 2, sometimes ciliate. PALEA hyaline, often minute or lacking, S T A M E N S 3. O V A R Y ellipsoid; styles terminal or subterminal; stigmas plumose, laterally exserted. G R A I N mostly obovoid (in cultivated forms globose or subglobose). P E D I C E L L E D S P I K E L E T 1-flowered or sterile, GLUMES awnless. L E M M A S 2, hyaline; lower empty; upper subtending a palea and a male flower or empty. PALEA hyaline, linear, PEDICEL linear, semiterete, or flat on the face and convex on the back, shorter than the sessile spikelet.

Annuals or perennials; culm ribbed or smooth, glabrous or sometimes villous at the nodes; leaf sheath ribbed, glabrous or sometimes pilose within at the junction with the blade; ligule scarious, sometimes with long hairs from the back; blade linear, more rarely filiform, distinctly ribbed, with or without a distinct or prominent midrib, sometimes scarious on the margins, glabrous or pilose with bulbous-based hairs; inflorescence an open panicle; primary branches often in whorls on the main rachis terete or angled, glabrous; internodes of ultimate rachis similar to the pedicels.

FIG. 84. Sorghastrum nutans (L.) Nash. 1, Plant, X 1/4; 2, panicle, X 1/2; J, pair of spikelets, X 3. (U.S.D.A. photograph.)

2l8

G R A S S E S OF H A W A I I

For an elaboration of the cultivated races of sorghums, see J. D. Snowdon's, The Cultivated Races of Sorghum (1936). 1.

Sorghum x almum Parodi, Rev. Argent. Agron. 10:361 (1943). A spontaneous hybrid whose putative parents are S. halepense and a member of the series Arundinaceae (Bor, i960).

2.

S. caffrorum Beauv., Ess. Agrost. 1 3 1 , 178 (1812). S. vulgare var. caffrorum (Retz.) Hubb. et Rehdr., Harvard Univ. Bot. Mus. Leaflet No. 1, p. 10 (1932).

3.

S. dochna (Forsk.) Snowdon, Kew Bull. 1935:234 (1935). S. vulgare var. saccharatum (L.) Boeri., Ann. Jard. Bot. Buetenzorg 8:69 (1890).

4.

S. dochna var. technicum Snowdon, Kew Bull. 1935:235 (1935). S. vulgare var. technicum (Koern.) Jav., Magyar Fl. 1:63 (1924).

5.

S. durra Stapf in Prain, Fl. Trop. Afr. 9:126 (1912). S. vulgare var. durra (Forsk.) Hubb. et Rehdr., Harvard Univ. Bot. Mus. Leaflet No. 1, p. 10 (1932).

6.

S. halepense (L.) Pers., Syn. PI. 1, 101 (1805). [Fig. 85] Holcus halepensis L., Sp. PI. 1047 (1753). Andropogon sorghum (L.) Brot, subsp. halepensis Hack, in DC., Monogr. Phan. 6, 501 (1889). A. halepensis (L.) Brot. var. typicus Asch, et Graeb., Syn. Mitteleur. Fl. 2, Abt. i, 47 (1898). A. halepensis (L.) Brot. var. genuinus (Hack.) Stapf in Hook, f., Fl. Brit. Ind. 7:183 (1896). A. halepensis Brot., Fl. Lusit. 1:89 (1804). S. roxburghii Stapf in Prain, Fl. Trop. Afr. 9:126 (1912). Sorghum vulgare var. roxburghii (Stapf) Haines, Bot. Bihar and Orissa 1034 (1934).

7.

8.

S. sudanense (Piper) Stapf in Prain, Fl. Trop. Afr. 9 : 1 1 3 (1917)Andropogon sorghum sudanensis Piper, Proc. Biol. Soc. Wash. 28:33 (i9i5)Holcus sorghum sudanensis (Piper) Hitchc., Proc. Biol. Soc. Wash. 29:128 (1916).

PANICOIDEAE -

ANDROPOGONEAE

219

FIG. 85. Sorghum halepense (L.) Pers. Plant, x 1/2; two views of terminal raceme, x 5. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed.; U . S . D . A . photograph.)

220

GRASSES OF

HAWAII

Spodiopogon Trin., Fund. Agrost. 192 (1820). SPIKELETS laterally compressed or subterete, in pairs, one pedicellate, one sessile in simple branches or paniculate spikes, FLOWERS 2; the upper floret bisexual or female, the lower male, G L U M E S stiff; the outer glume lanceolate, convex or flat, 5-9-nerved, acuminate or toothed, inner glume keeled, membranous, lanceolate, 3-7-nerved. L E M M A S and PALEA very thin and transparent, fertile lemma with a twisted awn. L O D I C U L E S cuneate. ANTHERS linear, S T I G M A S linear, G R A I N free, narrowly fusiform. Tall grasses, leaves often long-petioled.

1.

Spodiopogon aureus Hook, et Am., Bot. Beechey's V o y . 273 (1840). Hitchcock (1922, p. 215) states, 'Spodiopogon aureus Hook. & Arn. is admitted to the Hawaiian flora by Hillebrand on the authority of Monroe but this reference is probably incorrect.'

Themeda Forsk., Fl. Aegypt-Arab. 178 (1775). Anthistiria

Linn, f., Nov. Gram. Gen. 35 (1779).

A P P A R E N T SPIKELET consisting of 4 sessile, sterile or male spikelets which form an involucre and which surround a short rachis bearing 1 sessile and 2 pedicelled spikelets; the whole group of 7 spikelets enclosed in a spathe. I N V O L U C R A L SPIKELETS sterile or 1-flowered, sessile, lanceolate, acuminate, LOWER G L U M E lanceolate or linear-lanceolate, acuminate, more or less flat on the back, glabrous or covered with stiff bulbousbased hairs; upper hyaline or membranous, L E M M A S 2, hyaline; lower empty; upper empty or subtending a male flower, SESSILE SPIKELET 1-flowered, densely villous at the base, usually awned. LOWER G L U M E thin or subchartaceous, oblong-lanceolate or oblong-elliptic, obtuse or truncate, somewhat flat on the back, usually shortly pilose; upper similar or thinner in texture, L E M M A S 2, hyaline, LOWER L E M M A empty; upper reduced to a 2-lobed stipe, usually awned from the sinus, subtending a bisexual flower; awn when present, 6 - 1 3 times as long as the lemma. PALEA hyaline, small or lacking, L O D I C U L E S 2, rather large, glabrous. S T A M E N S 3 . O V A R Y oblong in outline; style plumose above, G R A I N linearobovate in outline, grooved on the face, PEDICELLED SPIKELET 1-flowered or sterile, lanceolate, long-acuminate, LOWER G L U M E rounded on the back, glabrous or covered with stiff bulbous-based hairs; upper similar but narrower and distinctly keeled, PEDICEL terete or flat on the face and convex on the back.

PANICOIDEAE - ANDROPOGONEAE

221

Perennials; culms smooth, glabrous; leaf sheath shorter than the internode of the culm, ribbed, glabrous or with stiff bulbous-based hairs; ligule membranous, finely ciliate; blade usually longitudinally folded, with fine veins and a distinct midrib, often scabrid, glabrous or sometimes pilose at the junction with the sheath; inflorescence formed of groups of spikelets subtended by a spathe; one or more of such groups subtended by a common spathe; the whole forming a lax panicle; groups of spikelets forming the panicle on slender branches of the culm, usually drooping; main spathe passing into a long acuminate point, ribbed, glabrous or covered with scattered bulbous-based hairs. 1.

Themeda arguens (L.)

Hack, in DC., Monogr. Phan. 6:657

(1889). 2.

T. australis (R. Br.) Stapf in Prain, Fl. Trop. Afr. 9:420 (1919).

3.

T. gigantea (Cav.J Hack, in D C . , Mongr. Phan 6:670 (1889).

4.

T. quadrivalvis (L.)

5.

T. triandra Forsk., Fl. Aegypt.-Arab. 178 (1775). [Fig. 86]

Kuntze,

Rev. Gen. PI. 2:794 (1891).

222

GRASSES

OF

HAWAII

FIG. 86. Themeda triandra Forsk. Plant, X 1/4. 1, Ligule. 2, Group of spikelets showing: 2 a, involucral sterile or male spikelets; 2b, bisexual awned spikelet, and 2c, one of two pedicelled spikelets, X 2. 3, Bisexual and pedicelled spikelets, X 4. (From N. L . Bor, T h e Grasses of Burma, Ceylon, India and Pakistan.)

PANICOIDEAE -

MAYDEAE Dumort.,

MAYDEAE

223

Obs. Gram. Belg. 84, 142 (1823).

Spikelets unisexual, dissimilar, monoecious in the same inflorescence or in different inflorescences, when in the same inflorescence the male spikelets above the female. Male spikelets 2-flowered, mostly paired, one sessile, one pedicelled or both pedicelled. Glumes enclosing the florets, chartaceous or membranous. Lemmas hyaline. Paleas, if present, similar. Stamens 3 ; anthers linear. Lodicules 2, fleshy. Female spikelets 2-flowered, with the lower floret sterile, solitary or sometimes paired, sunk in the cavities of a thickened jointed rachis, crowded in rows on a thick spongy rachis, or solitary, enclosed in a bony, hardened false involucre derived from a metamorphosed sheath. Glumes firm or thin, emarginate or lobed. Lemmas membranous, short, unawned; paleas somewhat similar or the lower absent. Lodicules 2 or lacking, GRAIN subglobose or flattened, enclosed in the hardened false involucre or metamorphosed sheath or surrounded by the indurated glumes or enclosed between the hardened lower glume and the adjacent joint of the rachis. Hilum basal, ovate, linear or punctiform. Starch grains simple. Annual or perennial. Leaf blades linear to lanceolate, with panicoid anatomy; silica cells cross- or dumbbell-shaped; microhairs 2-celled, slender, elongate. Ligules well developed. Chromosomes small; basic number 5, 9, (10). Genera :

Coix

Tripsacum

Euchlaena

Zea

Polytoca

KEY TO THE GENERA OF MAYDEAE i.

Female spikelets completely enclosed in a metamorphosed leaf sheath which takes the form of a spherical or cylindrical, osseous or ivory beadlike structure Coix p. 224

1.

Female spikelet not enclosed in a false involucre as above 2.

Female spikelets in crowded longitudinal rows on a very thick axis; male spikelets in ample terminal panicles . . . Zea p. 228

224

GRASSES

2.

OF

HAWAII

Inflorescence, especially the female, not as above 3.

Female spikelets not enclosed partly or wholly in a hardened lower glume, usually sunk in the slotted axis joint; female spikelets solitary at each node . Euchlaena p. 225 Female spikelets enclosed in a structure partly derived from the hardened lower glume and partly from the thickened axis joint or internode

3.

4.

T h e fruit case formed partly by the lower glume and chiefly by the broad thickened internode or joint of the rachis; male and female spikelets in the same spike, the male terminal; sheath not tessellately nerved Tripsacum p. 225

4.

T h e fruit case chiefly formed by the lower glume and in a lesser degree by the narrow joint or internode of the rachis; spikes monoecious Polytoca p. 225

Coix L., Sp. PI. 972 (1753). SPIKELETS unisexual, S T A M I N A T E SPIKELETS 2-flowered, in twos or threes on the continuous rachis, the normal group consisting of a pair of sessile spikelets with a single pedicillate spikelet between, the latter sometimes reduced to a pedicel or wanting, GLUMES membranous, obscurely nerved, L E M M A and PALEA hyaline, S T A M E N S 3. P I S T I L L A T E SPIKELETS 3 together, 1 fertile and 2 sterile at the base of the inflorescence. GLUMES of the fertile spikelet several-nerved, hyaline below, chartaceous in the upper, narrow pointed part, FIRST G L U M E very broad, infolding the spikelet, the margins infolded beyond the two lateral stronger pair of nerves, S E C O N D G L U M E narrower than the first, keeled, STERILE L E M M A similar but a little narrower, FERTILE L E M M A and palea hyaline, STERILE SPIKELETS consisting of a single narrow tubular glume as long as the fertile spikelet, somewhat chartaceous. Tall branched grasses with broad flat blades, the monoecious inflorescences numerous on long stout peduncles, these clustered in the axils of the leaves, each inflorescence consisting of an ovate or oval pearly-white or drab, beadlike, very hard, tardily deciduous involucre (much modified sheathing bract) containing the pistillate lower portion of the inflorescence, the points of the pistillate spikelets and slender portion of the inflorescence protruding through the orifice at the apex,

225

PANICOIDEAE - MAYDEAE

the staminate upper portion of the inflorescence 2-4 cm long, soon deciduous, consisting of several clusters of staminate spikelets. 1.

Coix lacryma-jobi L., Sp. PL

Euchlaena Schrad. in Linnaea

972 (1753).

8 (1833)

[Fig.

87]

Litt.-Bericht,

25 (1833).

STAMINATE SPIKELETS as in Zea, PISTILLATE SPIKELETS solitary on opposite sides, sunken in cavities in the hardened joints of an obliquely articulate rachis. FIRST GLUME indurate, covering the cavity, SECOND GLUME membranous, LEMMA hyaline. Robust annuals and perennials with broad flat blades, terminal panicles of staminate spikelets and axillary spikes or pistillate spikelets. Spikes infolded in foliaceous spathes or husks, two to several of these together enclosed in the leaf sheaths.

1. 2.

Euchlaena mexicana Schrad., Ind. Sem. Hort. Gottingen ( 1 8 3 2 ) , reprinted in Linnaea 8, Litt.- Bericht, 2 5 ( 1 8 3 3 ) . [Fig. 88] E. perennis Hitchc., Journ. Wash. Acad. Sci. 1 2 : 2 0 7 ( I 9 2 2 )-

Polytoca R. Br. in Benn., PL Jav. Rar.

20 ( 1 8 3 8 ) .

SPIKELETS in pairs, one sessile and one pedicelled. MALE SPIKELETS lanceolate, 2-flowered. GLUMES 2-several-nerved, lower glume 2-keeled. LEMMAS hyaline, paleate, triandrous. ANTHERS very long, LODICULES cuneate. FEMALE SPIKELETS imbricate; the pedicelled spikelet imperfect, its pedicel confluent with the rachis. GLUMES of sessile spikelet: the lower oblong, coriaceous, margins inflexed embracing the rachis, at length crustaceous; the upper thin, acuminate, LEMMAS hyaline, STAMINODES and LODICULES lacking, OVARY minute, styles long, stigmas slender, GRAIN very small, orbicular, enclosed in the hardened outer glumes. Tall branching grasses, nodes bearded, flowering branches fascicled, spatheaceous. Leaves long, flat. Spikes solitary or panicled, bisexual and female below, or the upper male only, rachis jointed.

1.

Polytoca macrophylla Benth., Journ. Linn. Soc. Bot. 19:52 (1881).

Tripsacum L., Syst. Nat. ed.

10, 2 : 1 2 6 0 ( 1 7 5 9 ) .

SPIKELETS unisexual, STAMINATE SPIKELETS 2-flowered, in pairs on one side of a continuous rachis, one sessile, the other sessile or pedicellate

FIG. 87. Coix lacryma-jobi L . Terminal portion of flowering culm. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

FIG. 88. Euchlaena mexicana Schrad. Plant much reduced; pistillate inflorescence enclosed in bract (a), and with portion of bract removed (b), X I. (c) Lateral view of rachis joint and fertile spikelet, and (d) dorsal view of same, showing first glume, X 2. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd ed. ; U . S . D . A . photograph.)

228

GRASSES OF H A W A I I

similar to those of Zea. GLUMES firmer, PISTILLATE SPIKELETS solitary (a minute rudiment or a sterile spikelet sometimes found), on opposite sides at each joint of the thick, hard articulate lower part of the same rachis, sunken in hollows in the joints, consisting of one perfect floret and a sterile lemma, FIRST GLUME coriaceous, nearly infolding the spikelet, fitting into and closing the hollow of the rachis. SECOND GLUME similar to the first but smaller, infolding the remainder of the spikelet. STERILE LEMMA, FERTILE LEMMA, and PALEA very thin and hyaline, these progressively smaller. Robust perennials, with usually broad flat blades and monoecious terminal and axillary inflorescences of 1-3 racemes, the pistillate part below, breaking up into bony, seedlike joints, the staminate above on the same rachis, deciduous as a whole. 1.

Tripsacum dactyloides (L.) L., Syst. Nat. ed. 10, 2:1261 (1759). [Fig. 89]

2.

T. laxum Nash, N. Amer. Fl. 17:81 (1909).

Zea, L., Sp. PI. 971 (1753). Monoecious annual; MALE SPIKELETS in racemes, GLUMES subequal, lanceolate in outline, acute or acuminate, 7-11-nerved. LEMMAS 2, subequal to the glumes, hyaline, 3-5-nerved, and like the glumes ciliate toward the tips, PALEAS subequal to the lemmas, those of the upper florets longer than those of the lower, shorter than their lemmas. LODICULES 2, fleshy, truncate, STAMENS 3, anthers long, FEMALE SPIKELETS closely sessile, densely imbricated on the thick axis of the spike, RACHIS inarticulate and spongy, GLUMES broad and fleshy at the base, membranous or hyaline upwards, or rarely subherbaceous, short and broad, obtuse or emarginately 2-lobed, the upper margins ciliate. LEMMAS broad and hyaline, the lower sterile, the upper fertile, PALEAS membranous, short and broad, STAMINODIA none, STYLE simple, much elongated, filiform, shortly bifid at the apex, the stigmas papillose, GRAIN hard, embryo oblong. Tall monoecious annuals; leaf blades broad, usually undulate; male panicles on long peduncles; female spikes solitary in the axis of the leaves, conspicuous from the numerous long exserted pendulous styles, enveloped by the broad leaf bases. 1.

Zea mays L., Sp. PI. 971 (1753). [Fig. 90]

FIG. 89. Tripsacum dactyloides (L.) L. 1, Plant, x 1/4; 2, inflorescence showing staminate (upper portion) and pistillate (lower portion) spikelets, x 1 2 ; 3, portion of pistillate spikelet, x 10; 4, pair of staminate spikelets, x 10. (U.S.D.A. photograph.)

FIG. 90. Zea mays L . Pistillate inflorescence (ear) and 2 branches of staminate inflorescence (tassel), x 1/2; pair of pistillate spikelets attached to rachis (cob) with mature grains, the second glume showing, X 2; single pistillate spikelet soon after flowering, X 4; staminate spikelet, X 2. (From Hitchcock and Chase, Manual of the Grasses of the United States, 2nd éd.; U . S . D . A . photograph.)

PANICOIDEAE -

231

PANICEAE

PANICEAE R. Br. in Flinders, Voy. Terra Australis 2:582 (1814). Spikelets usually similar, bisexual, rarely unisexual, solitary or paired, usually deciduous from the pedicels at maturity, 2-flowered, with the lower floret male or sterile, and the upper bisexual. Glumes membranous or herbaceous usually unequal, the lower rarely reduced to a mere thickened rim or absent altogether, rarely both absent; lower lemma of similar texture to the glumes, paleate or not; upper lemma of firmer texture than the lower, smooth or rarely rugose; palea of the same texture as the lemma; lemma awnless or with a very short stout awn from the tip; rachilla not or very rarely produced. Lodicules usually 2. Stamens 3. Ovary glabrous; styles 2, stigmas plumose. Grain firmly enclosed by the lemma and palea; hilum usually basal, punctiform; embryo 1/2 the length of the grain; starch grains simple. Annual or perennial herbs, rarely woody. Leaf blades linear to lanceolate, rarely ovate, tapering or rounded at the base, rarely cordate, usually flat, with panicoid anatomy; silica cells nodular, dumbbell- or cross-shaped; microhairs 2-celled, filiform; others club-shaped in some species of Digitaria. First foliage leaf of seedling flat and horizontal. Spikelets arranged in effuse or contracted panicles or in a spikelike inflorescence and then frequently surrounded by reduced bristle-like branches. Ligules membranous, a row of hairs or rarely absent. Chromosomes small; basic numbers 9, 10, rarely 7, 15, 17, 19. Genera:

Alloteropsis Axonopus Brachiaria Cenchrus Digitaria Dissochondrus

Echinochloa Eriochloa Ixophorus Melinis Oplismenus

Panicum Paspalidium Paspalum Pennisetum Rhynchelytrum

Sacciolepis Setaria Stenotaphrum Trichachne Urochloa

K E Y T O T H E G E N E R A OF P A N I C E A E

1.

Spikelets finely awned or mucronate from the notched tips of the upper and lower glume and lower lemma, delicately pedicelled, panicled; lower glume minute; upper lemma rigidly membranous

232

GRASSES OF

2.

HAWAII

Upper glume markedly gibbous, 5-nerved; lower glume separated from the upper by a space more than 0.6 mm long Rhynchelytrum

2.

1.

P- 295 Upper glume not gibbous, 5-7-nerved; spikelets awned, not compressed laterally Melinis p. 260 Spikelets not awned or if awned then subsessile in false secund, variously arranged spikes and with awns from the entire tips of the upper glumes and lower lemmas (Echinochloa spp.) or from the tips of both glumes (Oplismenus) ; upper lemma mostly crustaceous, rarely chartaceous

3.

Spikelets falling singly, not subtended by bristles, or if so, then the bristles persisting after the spikelets have fallen (Setaria) 4.

Spikelets arranged in more or less open panicles, or with the panicles contracted and spikelike 5.

Spikelets not subtended by bristle-like branches 6.

Spikelets arranged in open or contracted panicles; upper glume as long as the spikelet

6.

5.

.

.

.

Panicum p. 263

Spikelets usually arranged in cylindrical spikelike panicles ; upper glume inflated or not; upper lemma and palea indurated and closed at the apex . . Sacciolepis p. 300 Spikelets subtended by one or more bristle-like branches

7.

Both florets of the spikelet bisexual; lower and upper lemmas coriaceous, smooth . . Dissochondrus

7.

Florets unalike as to sex

P- 253

8.

Lower lemma membranous, the upper chartaceous, rugose; lower floret male or neuter, the upper bisexual Setaria p. 302

8.

Upper lemma 5-nerved, punctate-scabrous, flattened and with 2 grooves on the back; lower floret male or neuter, the upper bisexual Ixophorus p. 260

P A N I C O I D E A E - PANICEAE

233

Spikelets arranged in one-sided spikes or spikelike racemes; spikes or racemes digitate or scattered, rarely solitary

4. 9.

Lemma of the upper floret more or less crustaceous or coriaceous, usually with narrow inrolled margins, exposing much of the palea 10. Lower glume and lowest internode of the rachilla not forming a swollen callus at the base of the spikelet 11. Lower glume (when present) turned away from the rachis of the racemes or spike, the back of the upper lemma facing it, i.e., spikelets abaxial 12. Lower glume developed, although sometimes small 13. Spikelets not sunken in hollows in a thickened corky rachis 14. Glumes acuminate or awned, rarely only acute; upper lemma not mucronate 15. Leaf blades linear; racemes dense; culms erect or suberect Echinochloa P- 255 15. Leaf blades lanceolate to ovate; racemes loose to moderately dense; culms creeping and ascending . . . . Oplismenus p. 2 6 2 14. Glumes awnless, if acuminate, then with the upper lemma mucronate 16. Upper lemma acute, not mucronate

Paspalidium

p. 285

16. Upper lemma obtuse, mucronate or very short-awned . . . . Urochloa P- 3 1 1 13. Spikelets sunken in hollows in a thickened rachis, the latter disarticulating at maturity

Stenotaphrum

p. 306

GRASSES OF

234

HAWAII

12. Lower glume usually absent; spikelets plano-convex Paspalum p. 285 11.

Lower glume turned toward the rachis, the back of the upper lemma turned away from it, i.e., spikelets adaxial

17. Lower glume present

.

17. Lower glume absent

.

.

.

Brachiaria p. 236 Axonopus

P- 235 10. Lower glume and lowest internode of the rachilla forming a swollen callus at the base of the spikelet; upper lemma mucronate or short-awned . . Eriochloa p. 258 9.

Lemma of the upper floret thinly cartilagenous, usually with flat hyaline margins 18. Spikelets awnless 19.

Spikelets enveloped in long silky hairs

Trichachne p. 308 19. Spikelets often hairy, but not surrounded by long silky hairs Digitaria p. 246 18. Spikelets awned 3.

.

.

.

.

Alloteropsis

P- 235 Spikelets with an involucre of bristles or subtended by a solitary bristle and falling with or without the bristles at maturity, solitary or in clusters 20. Upper lemma smooth; bristles caducous 21. Involucre of free, naked or plumose bristles 22. Lemmas heteromorphous; lower often 3-lobed; upper shorter, shining; upper floret readily disarticulating; rachis with decurrent wings below the insertion of the pedicels . . . . Pennisetum (Eriochaeta) P- 293 22. Lemmas more or less alike or lower much reduced; upper floret not deciduous; rachis ribbed but not conspicuously so, not winged Pennisetum P- 293

PANICOIDEAE -

PANICEAE

235

21.

Involucre of spines or rigid bristles united at the base into a hard cup Cenchrus p. 238 20. Upper lemma transversely rugose; bristles persistent Setaria p. 302 Alloteropsis J.S. Presl ex C.B. Presl, Rel. Haenk. 343 (1830), emend. Hitchc., Contr. U. S. Natl. Herb. 12:210 (1909). SPIKELETS in pairs or threes together or fascicled, secund and abaxial (i.e., the back of the lower glume and sterile lemma turned away from the axis), ovate or elliptical to lanceolate-oblong in outline, acute or acuminate, slightly or conspicuously dorsally compressed, mostly awned, falling entire below the glumes, FLOWERS 2, the lower usually male, the upper bisexual, GLUMES unequal, the lower smaller, membranous to hyaline, very acute or mucronulate, 3-1-nerved, the upper equal or subequal to the spikelet, membranous to chartaceous, 5-nerved, densely ciliate along the nerves and sometimes the margins expanded into broad wings, LEMMAS 2, the lower subtending a male floret, resembling the upper but not ciliate, minutely truncate at the apex, the upper chartaceous, mucronate or produced into a straight awn, glabrous, ciliolate upwards, 5-nerved. PALEAS with broadly auricled 'flaps,' the lower short, deeply bifid, the upper equal to its lemma and 2-keeled. LODICULES 2, broadly cuneate. STAMENS 3. STYLES distinct; stigmas laterally exserted. GRAIN enclosed by the lemma and palea, elliptical-oblong in outline, much compressed; hilum basal, punctiform. Perennials or annuals; leaf blade flat or more or less convolute; ligule membranous, ciliate or ciliolate, short or reduced to a mere rim; inflorescence of more or less spikelike digitate or subdigitate racemes; racemes sessile or peduncled, the rachis triangular.

1.

Alloteropsis cimicina (L.J Stapf in Prain, Fl. Trop. Afr. 9:487 (1919). Coridochloa cimicina (L.) Nees, Edinburgh New Phil. Journ. 15:381 (1833)-

Axonopus Beauv., Ess. Agrost. 12, 154 (1812). SPIKELETS solitary, secund, depressed-biconvex, not turgid, adaxial, subsessile and alternate in two rows on one side of a 3-angled rachis. FLORET 1. GLUME 1, the lower glume absent, the upper oblong-elliptical, membranous or chartaceous, 2-4-nerved, oblong-elliptical in outline,

236

GRASSES OF H A W A I I

rather obtuse, LEMMAS 2, the lower sterile, without a palea, resembling the upper glume in shape, size and consistency, glabrous or sparsely appressed silky-pubescent near the margin; fertile lemma crustaceous, indurated, pale, glabrous, without nerves, obtuse, the margins inrolled and embracing the palea. PALEA I , indurated, smooth, not keeled, equal in size and shape to the fertile lemma and embraced by its margins but ultimately free, LODICULES 2, broadly cuneate. STAMENS 3, dark colored. STYLES distinct, the stigmas laterally exserted near the tip of the spikelet. Stoloniferous or tufted perennials with usually flat or folded rather obtuse leaf blades; ligule membranous; inflorescence of spikelike racemes, digitate or racemose along the main axis. 1. 2.

Axonopus affinis Chase, Journ. Washington Acad. Sci. 28:180 (1938). A. compressus (Sw.) Beauv., Ess. Agrost. 12, 154, 167 (1812). [Fig. 91] Milum compressum Sw., Prodr. Veg. Ind. Occ. 24 (1728). Paspalum compressum (Sw.) Rasp., Ann. Sci. Nat. Bot. 1. 5:301 (1825).

Brachiaria Griesb. in Ledeb., Fl. Ross. 4:469 (1853). SPIKELETS solitary or in pairs, more rarely in fascicles of three or more, pedicelled, secund and abaxial, falling entire, FLOWERS 1-2. GLUMES dissimilar and unequal in length, membranous; lower smaller than the upper, ovate-lanceolate to ovate-orbicular, rounded at the apex, 3 - 1 1 nerved, villous or glabrous; upper elliptic, usually shortly acuminate or with acumen 1/3 as long as the glume, rounded on the back, 5-7nerved, densely villous or with a few scattered hairs or glabrous, LEMMAS 2, very dissimilar, LOWER LEMMA resembling the upper glume, 5-7nerved, subtending a palea and sometimes a male flower, PALEA almost as long as its lemma, membranous or hyaline, UPPER LEMMA ovate-elliptic, obtuse, acute or subacute, sometimes shortly acuminate and sometimes minutely rugose on the back, faintly 5-nerved, subtending a palea and a bisexual flower, PALEA similar to and embraced by its lemma, with 2 marginal flaps at the base, LODICULES 2, broadly cuneate. OVARY subovoid; styles distinct, plumose above, laterally exserted from the upper part of the spikelet. GRAIN tightly enclosed by the more or less hardened lemma and palea, broadly oblong or elliptic in outline; embryo 1/2 to more than 3/4 as long as the grain. Perennials or annuals; culm ribbed, sparsely pubescent or finely villous, usually villous at the nodes; leaf sheath longer or shorter than

PANICOIDEAE -

PANICEAE

237

238

GRASSES OF H A W A I I

the internode of the culm, prominently ribbed, sparsely covered with tubercule-based hairs sometimes almost villous; ligule long-ciliate; blade flat, lanceolate or linear-lanceolate, or long-linear and tapering to a fine point, closely veined, with a distinct midrib and sometimes with distinct lateral veins, with often undulating margins or margins sometimes cartilaginous, sometimes toothed, often scabrid to the touch, hispid or finely villous or covered with tubercule-based hairs; inflorescence of spikelike racemes arranged on a main rachis; rachis semiterete and pilose with scattered hairs, or rounded on the back and concave on the face and with ciliate margins. 1.

Brachiaria brizantha (Hochst. ex A. Rich.) Stapf in Prain, Fl. Trop. Afr. 9:531 (1919). [Fig. 92]

2.

B. ciliatissima (Buckl.) Chase in Hitchc., U. S. Dept. Agr. Bull. 772, 221 (1920).

3.

B. dictyoneura (Fig. et Be Not.) Stapf in Prain, Fl. Trop. Afr. 9:512 (1919).

4.

B. mutica (Forsk.) Stapf in Prain, Fl. Trop. Afr. 9:526 (1919). [Fig- 93] Panicum muticum Forsk., Fl. Aegypt.-Arab. 20 (1775). P. purpurascens Raddi, Agrost. Bras. 47 (1823). P. barbinode Trin., Mem. Acad. Sci. Petersb. ser. 6, 3:256 (1834). P. guadaloupense Steud., Syn. PI. Glum. 1:61 (1854).

5.

B. plantaginea (Link) Hitchc., Contr. U. S. Natl. Herb. 1 2 : 2 1 2 (1909).

6.

B. reptans (L.) Gard. et C.E. Hubb. in Hook., Icones PI. (1938) sub. tab. 3363. Panicum reptans L., Syst. Nat. ed. 10, 2:870 (1759). P. caespitosum Sw., Fl. Ind. Occ. 1:146 (1797).

Cenchrus L., Sp. PI. 1049 (1753). SPIKELETS 1-5, enclosed in a spiny involucre and falling entire with the involucre; involucre pedicelled, of several hardened connate spinous bracts which appear as two single bracts covered with hard spines. FLOWERS 1-2. GLUMES unequal, usually small and hyaline or membranous ; lower sometimes suppressed; upper ovate, acuminate, obtuse, rarely 1/2 as long as the spikelet, if longer, then up to 7-nerved. LEMMAS 2; lower ovate-lanceolate, acuminate, empty or subtending a palea and

239

FIG. 92. Brachiaria brizantha ( H ö c h s t , ex A . R i e h . ) S t a p f . Plant, x 1/2. 1, L i g u l e ; 2, axis of r a c e m e ; 3, s p i k e l e t ; 4, l o w e r g l u m e ; 5, u p p e r g l u m e ; 6, l e m m a of u p p e r floret; 7, p a l e a ; all x 4. ( F r o m N . L . Bor, T h e G r a s s e s of B u r m a , C e y l o n , India and Pakistan.)

24°

FIG. 93. Brachiaria mutica (Forsk.) Stapf. Plant, x 1; part of spike, x 3; two views of spikelets, x 10. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

241

PANICEAE

a male flower; upper ovate-acuminate, subacute, with widely inflexed margins, 5-7-nerved, membranous to chartaceous, subtending a palea and a bisexual flower, PALEA similar to its lemma, LODICULES none. STAMENS 3 . OVARY ovoid; style distinct or connate at the base, plumose above, GRAIN elliptic, obovate or oblong in outline; embryo 2 / 3 as long as the grain. Annuals or perennials; culm finely striate, glabrous; leaf sheath shorter than the internode of the culm, ribbed, pilose at the junction with the blade, otherwise glabrous; ligule ciliate; blade flat or longitudinally folded, closely ribbed, with a distinct midrib, glabrous; inflorescence consisting of the involucres surrounding the spikelets arranged on a central axis. 1.

Cenchrus agrimonioides Trin., Gram. Pan. 7 2 ( 1 8 2 6 ) . [Fig. C. calyculatus var. uniflorus Hillebr., Fl. Haw. Is. 5 0 5 ( 1 8 8 8 ) .

2.

C. agrimonioides var. laysanensis F.B.H. Brown in Christ, and Caum., Bull. Bernice P. Bishop Mus. 8 1 : 2 0 ( 1 9 3 1 ) . [Leeward Islands, Hawaii.] C. agrimonioides Hitchc., Mem. Bernice P. Bishop Mus. 8:211 ( 1 9 2 2 ) pro parte.

3.

C. biflorus Roxb., Fl. Ind. 1 : 2 3 8

4.

C. ciliaris L., Mant. Alt. PI. 2 : 3 0 2 (1771). [Fig. 9 5 ] Pennisetum cenchroides (L.) L. Rich, in Pers., Syn. PI. 1:72 P. ciliare (L.) Link, Hort. Berol. 1 : 2 1 3 ( 1 8 2 7 ) .

5.

6.

94]

(1820).

C. echinatus L., Sp. PI. 1 0 5 0 ( 1 7 5 3 ) . [Fig. C. pungens H.B.K., Nov. Gen. et Sp. 1 : 1 1 5 C. brevisetus Fourn., Mex. PI. 2 : 5 0 ( 1 8 8 6 ) .

(1805).

96] (1816).

C. echinatus var. hillebrandianus (Hitchc.) F.B.H. Brown, Bull. Bernice P. Bishop Mus. 8 4 : 6 5 ( 1 9 3 1 ) . C. hillebrandianus Hitchc., Mem. Bernice P. Bishop Mus. 8:211 (1922).

7.

C. pedunculata Degener et Whitney in Degener, Fl. Hawaiiensis, Family 4 7 (Cenchrus), fig. 2 5 ( 1 9 3 6 ) . [Fig. 9 7 ]

8.

C. setigerus Vahl, Enum. PI. 2 : 3 9 5 ( 1 8 0 6 ) . C. montanus Nees ex Royle, 111. Bot. Him. 4 1 6

(1840)

nom. nud.

242

GRASSES

OF

HAWAII

FIG. 94. Cenchrus agrimonioides Trin. Plant, x 1/2; spikelet, x 3; upper portion of spikelet showing the sharp awns. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

PANICEAE

243

FIG. 95. Cenchrus ciliaris L . 1 , Plant, X 1/2. 2, S p i k e l e t w i t h s u p p o r t i n g bristles (sterile b r a n c h e s ) ; 3, l o w e r g l u m e ; 4, u p p e r g l u m e ; 5, l o w e r l e m m a ; 6, u p p e r l e m m a ; 7, palea o f u p p e r l e m m a ; all X 4. ( F r o m N . L . Bor, T h e G r a s s e s of B u r m a , C e y l o n , India and Pakistan.)

244

GRASSES OF

HAWAII

FIG. 96. Cenchrus echinatus L . Plant, x 1 jz; spikelet x 3 ; terminal portion of bristles X 5. a, b, c, and d, Parts of the floret, all x 5. (From O . Degener, Flora Hawaiiensis.)

PANICOIDEAE -

PANICEAE

245

FIG. 97. Cenchrus pedunculata Degener et Whitney. Plant, X 1/2; spikelet, X 3; upper portion of spikelet showing bristles, X 5. (From O. Degener, Flora Hawaiiensis.)

246

GRASSES OF H A W A I I

Digitaria Heister ex Fabricus, Enum. Method. PI. ed. 1, 207 (1759). Syntherisma Walt., Fl. Carol. 76 (1788). SPIKELETS solitary or usually in pairs or threes, more rarely in fours and sixes, secund and abaxial, pedicelled, with the pedicels usually unequal. FLOWER 1 . GLUMES very dissimilar; lower a minute membranous scale or a very delicate readily disintegrating hyaline membrane, sometimes quite suppressed; upper linear-oblong, linear-lanceolate, or lanceolate, obtuse or acute, sometimes ciliate, villous or pilose, very rarely reduced or only 1/2 as long as the lower lemma, usually 3-nerved, with the nerves more or less parallel, membranous, LEMMAS 2, dissimilar; lower lanceolate and determining the outline and size of the spikelet, flat on the back, 5-7-nerved, very rarely 3-nerved, with the nerves parallel, sometimes ciliate, densely villous or pilose, rarely reduced to a small scale, subtending a minute palea; upper lanceolate, obtuse sometimes shortly acuminate, with the margins inflexed, faintly 3-nerved, subtending a palea and a bisexual flower, PALEA similar to its lemma, often completely enclosed by it. LODICULES 2, broadly cuneate. STAMENS 3. OVARY ovoid; styles distinct, plumose above, laterally exserted near the top of the spikelet. GRAIN tightly enclosed by the slightly hardened lemma and palea; embryo usually 1/2 as long as the grain.

Annuals or perennials, sometimes creeping and rooting at the nodes; culm smooth, glabrous; leaf sheath longer or shorter than the internode of the culm, ribbed, softly villous or pilose with bulbous-based hairs or glabrous; ligule membranous, sometimes with hairs from the back; blade flat or rolled, mostly long-linear, very rarely lanceolate or linear-lanceolate, sometimes with cartilaginous and undulate margins, with or without a distinct midrib, with the lateral veins evident or not, sparsely pilose with bulbous-based hairs; inflorescence of narrow spikelike racemes, arranged digitately or subdigitately at the end of the culm, more rarely the racemes solitary; rachis flattened, usually with a raised keel on the inner face and with the spikelets arranged on either side of the keel. The reader is referred to Henrard (1950) for an excellent discussion on the nomenclature of the genus. 1.

2. 3.

Digitaria adscendens (H.B.K.) Henr., Blumea 1:92 (1934). [Fig. 98] D. henryi Rendle, Journ. Linn. Soc. Bot. 36:323 (1904). D. argyrograpta (Nees) Stapf in Dyer, Fl. Cap. 7:374 (1898). D. bicornis (Lamk.) Roem. et Schult. ex Loud., Hort. Brit. 24 (1830).

PANICOIDEAE - PANICEAE

247

FIG. 98. Digitarla adscendens (H.B.K.) Henr. Plant, x 1/2; portion of spike, x 4; spikelet, X 10. (From O. Degener, Flora Hawaiiensis.)

248

G R A S S E S OF H A W A I I

4. D. brownii (Roem. et Schult.) Hughes, Kew Bull. 1923: 313 (1923). 5. D. debilis (Desf.) Willd., Enum. PI. 91 (1809). Panicum debile Desf., Fl. Atlant. 1:59 (1798). Syntherisma debilis (Desf.) Skeels, U. S. Dept. Agr. Bur. PI. Ind. Bull. 261, 30 (1912). 6. D. decumbens Stent, Bothalia 3:150 (1930). [Fig. 99] 7. D. diagonalis (Nees) Stapf in Dyer, Fl. Cap. 7:381 (1898). 8. D. didactyla Willd., Enum. PI. Hort. Berol. 91 (1809). 9. D. divaricatissima (R. Br.) Hughes, Kew Bull. 1923:314 (1923). 10. D. diversinervis (Nees) Stapf in Dyer, Fl. Cap. 7:379 (1891). 1 1 . D. eriantha Steud., Fl. Bot. Zeit. Jahrb. 12:468 (1829). 12. D. eriostachya Mez., Bot. Jahrb. 56 (no. 125) :8 (1921). 13. D. exilis (Kippist) Stapf, Kew Bull. 1915:385 (1915). 14. D. filiformis (L.) Koeler, Descr. Gram. 26 (1802). Panicum filiforme L., Sp. PI. 57 (1753). 15. D. fuscescens (Presl) Henr. in Meded., Rijks Herb. Leiden no. 61, p. 8 (1930). D. pseudo-ischaemum Büse in Miquel, PI. Junghn. 382 (1854). 16. D. glauca A. Camus, Bull. Soc. Bot. France 73:914 (1927). 17. D. herpoclados Pilger in von Rosen, Wiss. Ergeb. Schwed. Rhod.Kongo Exped. 1:200 (1915). 18. D. horizontalis Willd., Enum. PI. 92 (1809). 19. D. iburua Stapf, Kew Bull. 1915:382 (1915). 20. D. ischaemum (Schreb.) Schreb. ex Muhl., Descr. Gram. Plant. Calam. 131 (1817). 21. D. kilimandscharica Mez., Bot. Jahrb. 57:194 (1921). 22. D. longiflora (Retz.) Pers., Syn. PI. 1:85 (1805). Paspalum longiflorum Retz., Obs. Bot. 4, 15 (1786). Syntherisma longiflora (Retz.) Skeels, U. S. Dept. Agr. Bur. PI. Ind. Bull. 261, 30 (1912). 23. D. macroglossa Henr., Monograph Digitaria 419 (1950).

PANICOIDEAE - PANICEAE

Serv. Circ. 342.)

249

250

G R A S S E S OF H A W A I I

24. D. microbachne (Presi) Henr. in Meded., Rijks Herb. no. 16, p. 13 (1930). Panicum microbachne J.S. Presi in C.B. Presi, Rei. Haenk. 11298 (1830). Syntherisma microbachne (Presi) Hitchc., Mem. Bernice P. Bishop Mus. 8 (3): 177 (1922). 25. D. milanjiana (Rendle) Stapf in Prain, Fl.Trop. Afr. 9:430 (1919). 26. D. milanjiana (Rendle) Stapf ssp. eylesiana Henr., Monograph Digitaria 459 (1950). 27. D. monodactyla (Nees) Stapf in Dyer, Fl. Cap. 7:373 (1898). 28. D. natalensis Stent, Bothalia 3 : 1 5 2 (1930). 29. D. pentzii Stent, Bothalia 3:147 (1930). 30. D. phaeothrix (Trin.) Parodi, Physis. 9:13, 19 (1928). 31. D. pruriens (Trin.) Biise in Miquel, PI. Junghn. 379 (1854). [Fig. 100] Panicum pruriens Fisch. ex Trin., Gram. Pan. 77 (1826). D. consanguinea Gaud, in Freyc., Voy. Uran. Bot. 410 (1830). Paspalum sanguinale var. pruriens (Trin.) Hook, f., Fl. Brit. Ind. 7 : 1 5 (1896). Syntherisma consanguinea (Gaud.) Skeels, U. S. Dept. Agr. Bur. PI. Ind. Bull. 282, 33 (1913). S. pruriens (Trin.) Arthur, Torreya 19:83 (1919). 32. D. sanguinalis (L.) Scop., FI. Cam. ed. 2, 1 :$2 (1772). [Fig. 101] Panicum sanguinale L., Sp. PI. 57 (1753). Syntherisma sanguinalis (L.) Dulac, Fl. Haut. Pyr. 77 (1867). Paspalum sanguinale (L.) Lamk., Tab. Encycl. Meth. Bot. 1 : 1 7 6

im*)33. D. seriata Stapf in Prain, Fl. Trop. Afr. 9:432 (1919). 34. D. smutsii Stent, Bothalia 1:268 (1924). 35. D. swazilandensis Stent, Bothalia 3:150 (1930). 36. D. timorensis (Kunth) Bai. in Morot, Journ. de Bot. 4:138 (1890). 37. D. tricholaenoides Stapf in Dyer, Fl. Cap. 7:381 (1898). 38. D. uniglumis (Rich.) Stapf in Prain, Fl. Trop. Afr. 9:474 (1919). 39. D. valida Stent, Bothalia 3:148 (1930).

PANICOIDEAE — PANICEAE

251

FIG. 100. Digitaria pruriens (Trin.) Biise. Plant, x 1/2; two views of part of spike, x 3 ; pair of spikelets, X s ; spikelet, floret, and grain, all x 10. (From O. Degcner, Flora Hawaiiensis.)

252

GRASSES

OF

HAWAII

FIG. IOI, Digitarla sanguinalis (L.) Scop. Plant, X 1/2; part of spike, spikelet, floret, and grain, all x 4. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

PANICEAE

253

40. D. violascens Link, Hort. Berol. 1:229 i 1 ^ 2 ! ) D. chinensis (Nees) A. Camus, Not. Syst. Lecomte 4:48 (1923), non Horneman (1819). Paspalum minutiflorum Steud., Syn. PI. Glum. 1 : 1 7 (1854), non Desv. (1831). P. chinensis Nees in Hook, et Arn., Bot. Beechey's Voy. 231 (1826). Syntherisma chinensis (Nees) Hitchc., Mem. Bernice P. Bishop Mus. 8:173 (1922). S. helleri Nash, Minn. Bot. Stud. 1:798 (1897). Panicum violascens (Link) Kunth, Rev. Gram. 1:33 (1829). Dissochondrus (Hillebr.) Kuntze, Rev. Gen. PI. 2:770 (1891). One species recorded in Hawaii, described as follows: Dissochondrus biflorus (Hillebr.) Kuntze ex Hack in Engl, and Prantl, Planzenfam. Nachtr. 1:41 (1897). [Fig. 102] Setaria biflora Hillebr., Fl. Haw. Is. 503 (1888). Culms 60-120 cm tall, compressed, puberulent at the nodes; sheaths glabrous, compressed, keeled, extending at the summit beside the petiole into 2 membranous, more or less deciduous or fragile auricles as much as 4 cm long, especially prominent on the innovations; ligule short, scarious; blades flat, firm, 15-30 cm long, 1 - 2 cm wide, gradually acuminate, contracted at the base into a petiole, glabrous, scaberulous on the margin and toward the tip; panicle cylindrical, slender, spikelike, 10-20 cm long, somewhat loosely flowered, more or less interrupted at the base, the branches short and appressed, the axis puberulent; bristle flexuous, antrorsely scabrous, 5 - 1 0 mm long, solitary below each spikelet or wanting below some of the spikelets; spikelets 2.5-3 mm long, pointed, glabrous; first glume broad, obtuse, 1/4-1/3 as long as the florets, puberulent, scarcely nerved; second glume pointed beyond the florets, puberulent, faintly 5-nerved; florets yellowish, smooth and polished, about 2 mm long, slightly pointed, the tip minutely puberulent. According to Hitchcock (1922), 'The genus is based on Setaria biflora Hillebr., the only species, and was first described as a subgenus of Setaria by Hillebrand. Kuntze raised this subgenus to generic rank but inadvertently gave the species as Dissochondrus bifidus O K = Setaria bifida Hillebr.' Chase and Niles in their Index to Grasses (1962) note the following: 'Dissochondrus bifidus (Hillebr.) Kuntze, Rev. Gen. PI. i 2 : 7 7 o . 1891.

254

GRASSES OF HAWAII

FIG. 102. Dissochondrus biflorus (Hillebr.) Kuntze. (From A. S. Hitchcock, The Grasses of Hawaii.)

PANICOIDEAE - PANICEAE

255

Based on "Setaria (bifida)" Hillebrandt. Error for biflora Hillebr. Corrected by Hackel in Engler & Prantl, Pflanz. Nachtr. 2 : 4 1 . 1 8 9 7 . ' Echinochloa Beauv., Ess. Agrost. 5 3 ( 1 8 1 2 ) . SPIKELETS solitary or in pairs, or rarely in threes, sessile, subsessile, or pedicelled, falling entire, FLOWERS 1 - 2 . GLUMES unequal, herbaceous; lower usually less than 1/2 as long as upper, ovate from a clasping base, subacute, sometimes shortly acuminate, often distinctly 3-5-nerved; upper ovate, subacute, sometimes acuminate, convex on the back, 3-7nerved, pubescent or hispid, LEMMAS 2, dissimilar; lower similar to the upper glume, sometimes awned, subtending a palea and occasionally also a male flower; awn, when present, as long as or 3 times as long as the lemma, PALEA hyaline, subequalling its lemma, UPPER LEMMA ovate to lanceolate-ovate, acute, subacute, sometimes shortly acuminate, convex, smooth and polished on the back, chartaceous to crustaceous subtending a palea and a bisexual flower, PALEA similar to its lemma, with infolded margins, LODICULES 2, cuneate, fleshy, STAMENS 3. STYLES distinct, plumose above, exserted from near the tip of the spikelet. GRAIN broadly elliptic in outline; embryo more than 1/2 as long as the grain. Annuals or perennials, culm sometimes stout, ribbed or smooth, glabrous; leaf sheath equalling or shorter than the internode of the culm, ribbed, glabrous; ligule lacking or represented by a fringe of hairs; blade linear or long-lanceolate, flat or more rarely folded, finely and closely veined, with a distinct midrib; inflorescence of spikes or spikelike subracemes arranged on a main rachis; rachis simple, more rarely branched, flattened or rounded on the back, with a narrow raised keel on the inner face, with bristle-like hairs from the nodes. 1.

2.

3.

Echinochloa colonum (L.) Link, Hort. Berol. 2 : 2 0 9 (I^33)[Fig- 1 0 3 ] Panicum colonum L., Syst. Nat. ed. 10, 2 : 8 7 0 (1759). P. crusgalli var. colonum Coss., Richt. PI. Eur. 1 : 2 6 (1890). E. crusgalli (L.) Beauv., Ess. Agrost. 53, 1 6 1 ( 1 8 1 2 ) . [Fig. 104] Panicum crusgalli L., Sp. PL 56 (1753). E. crusgalli f. longiseta (Trin.) Farzvell, Mich. Acad. Sci. Rept. 2 1 : 3 4 9 (1920).

4.

Panicum crusgalli var. longisetum Trin., Gram. Icon. 2 : 1 6 2 (1826). E. cruspavonis (H.B.K.) Schult., Syst. Veg. 2, Mant. 269 ( 1 8 2 4 ) . Oplismenus cruspavonis H.B.K., Nov. Gen. et Sp. 1 : 1 0 8 ( 1 8 1 5 ) .

FIG. 103. Echinochloa colonum (L.) Link, (a) Culm, leaf sheath, and ciliate leaf base, x 3; (b) spikelet, X 1 2 ; (c) floret, adaxial view, showing indurated palea, x 1 2 ; (d) floret, abaxial view, showing indurated lemma, x 12; (e) habit, showing decumbent stems rooting at the nodes, x 1/4. (From H. L . Mason, A Flora of the Marshes of California.)

FIG. 104. Echinochloa crusgalli (L.) Beauv. (From D . Meredith, T h e Grasses and Pastures of South Africa.)

258

GRASSES

OF

HAWAII

E. crusgalli var. cruspavonis ( H . B . K . ) Hitchc., Contr. U . S. Natl. Herb. 22:148 (1920). 5.

E. frumentacea (Roxb.)

Link, Hort. Berol. 1:204

Panicum frumentaceum Roxb., Hort. Beng. 7 (1814) without descr., Fl. Ind. 1:307 (1820), non Salisb. (1796). E. colonum var. frumentacea

(Roxb.) Ridley, Fl. Malay Penin.

5:223 (1925). 6.

E. polystachya (H.B.K.)

Hitchc., Contr. U . S. Natl. Herb. 22:135

(1920). Oplismenus polystachyus H . B . K . , Nov. Gen. et Sp. 1:107 (1815). E. polystachya var. spectabilis (Nees) Martinez Crovetlo, Rev. Arg. Agron. 9:318 (1942). Panicum spectabile Nees in Trin., Gram. Pan. 138 (1826). 7.

E. stagnina (Retz.)

Beauv.,

Ess. Agrost. 53, 161, 171 (1812).

[Fig. 105] Panicum stagninum Retz., Obs. Bot. 5:17 (1789). 8.

E. walteri (Pursch)

Heller, Cat. N . Amer. PL ed. 2, 21 (1900).

Eriochloa Kunth in Humb. et Bonpl., Nov. Gen. et Sp. 1:94 (1816). in pairs or solitary, pedicelled, falling entire, FLOWERS 1 - 2 . very unequal; lower reduced to a small nerveless scale clasping the base of the spikelet or a minute cupular rim clasping the callously thickened beadlike lower internode; upper as long as the lemmas, ovate-elliptic, mucronate, sometimes shortly awned, faintly 3-5-nerved, glabrous or pilose, L E M M A S 2 ; lower similar in shape and texture to the upper glume, empty or subtending a palea and a male flower, PALEA, when present, usually similar to its lemma, UPPER L E M M A elliptic, with inflexed margins, abruptly mucronate, finely rugose, glabrous, subtending a palea and bisexual flower, PALEA similar in texture to its lemma, elliptic, obtuse, with inflexed margins, L O D I C U L E S 2, broadly cuneate, sometimes quite suppressed, STAMENS 3. STYLES distinct or united at the base, plumose above, laterally exserted near the tip of the spikelet. G R A I N enclosed by the upper lemma and palea, falling with them and the tightly appressed lower lemma and upper glume. Perennials or annuals; culm strongly ribbed or striate, glabrous; leaf sheath as long as or shorter than the internode of the culm, ribbed, glabrous; ligule a ciliated rim; blade linear, flat, with distinct lateral SPIKELETS

GLUMES

PANICOIDEAE -

FIG. 105. Echinochloa stagnina (Retz.) Beauv. O. Degener, Flora Hawaiiensis.)

PANICEAE

259

Plant, x 1/8; spikelct, x 5/2. (From

z6o

GRASSES OF H A W A I I

veins, with or without a distinct midrib; inflorescence of narrow racemes arranged up the main rachis; main rachis angled, grooved; racemes more or less erect, with the rachis flat on the back and with a raised keel on the face. 1.

Eriochloa polystachya H.B.K., Nov. Gen. et Sp. 1:95 (1815). E. subglabra (Nash) Hitchc., Contr. U. S. Natl. Herb. 12:208 (1909).

2.

E. procera (Retz.) C.E. Hubb., Kew Bull. 1930:256 (1930).

3.

E. punctata (L.) Desv. ex Hamilt., Prodr. PI. Ind. Occ. 5 (1825).

Ixophorus Schlecht., Linnaea 31:420 (1861-62). SPIKELETS 2-flowered, the upper flower bisexual or female, the lower staminate, arranged in two rows along the branches of a simple panicle; the pedicels as well as the main axis and primary branches, produced beyond the spikelets into slender, smooth and viscid awnlike bristles, which equal or exceed the spikelets in length, GLUMES shorter than the lemmas; lower glume very short, 3-nerved; upper glume, many-nerved, somewhat shorter than the lowest lemma, LEMMAS 2, longer than the glumes; the lower 5-nerved and much longer than the fertile lemma; fertile lemma 5-nerved, punctate-scabrous, flattened and with two grooves on the back, with a distinct scar near the base and mucronate or short awn-pointed, PALEA of male floret equalling its lemma, at first hyaline, the margins becoming broad winglike and cartilaginous in fruit. STAMENS 3 . STIGMAS plumose; styles long, distinct, GRAIN oblong-obtuse, compressed, free within its lemma and palea.

Rather broad-leaved, annual or perennial, branching grasses with a simple paniculate inflorescence of unilateral racemes. 1.

Ixophorus unisetus (Presl) Schlecht., Linnaea 311420, 747 (1862). [Fig. 106]

Melinis Beauv., Ess. Agrost. 54 (1812). SPIKELETS solitary, pedicelled, falling entire, FLOWER I . LOWER GLUME reduced to a minute scale; upper oblong-elliptic, truncate, shortly 2-lobed, mucronate or shortly awned from the sinus, prominently 7-nerved, with the nerves not anastomozing, glabrous or pilose above. LEMMAS 2; lower elliptic, 2-lobed, 5-nerved, empty, awned from the sinus; awn 3 times as long as the lemma; upper ovate, obtuse, minutely 2-lobed, obscurely 3-1-nerved, membranous, shining, subtending a palea and the bisexual flower, PALEA similar to its lemma, LODICULES 3,

PANICOIDEAE -

PANICEAE

2ÖI

FIG. 106. Ixophorus unisetus (Presl) Schlecht. Plant, x 1 ; pistillate and bisexual florets, x 20; glumes, x 10; grain, x 10. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

2Ó2

GRASSES

OF

HAWAII

very delicate, STAMENS 3. O V A R Y ellipsoid, style free, plumose above, laterally exserted. G R A I N terete, linear-ovate in outline ; embryo about 1 ¡2 as long as the grain. Perennials or annuals, culm smooth or faintly ribbed, leaf sheath longer or shorter than the internode of the culm, ribbed, pilose with bulbous-based hairs; ligule ciliate; blade flat, usually linear-lanceolate, tapering to a fine point, distinctly ribbed, with a more distinct midrib, sometimes subcrenulate on the margins, pilose with bulbous-based hairs; inflorescence a contracted and narrow or open and spreading panicle; ultimate branches of main rachis more or less capillary, pedicel of spikelet capillary. 1.

Melinis minutiflora Beauv.,

Ess. Agrost. 54 (1812).

Oplismenus Beauv., Fl. Oware et Benin 2:14 (1807). SPIKELETS solitary, in pairs, or in small clusters, pedicelled, secund and abaxial, falling entire, FLOWER usually 1, sometimes 2. GLUMES equal, elliptic, 3-5-nerved, awned, membranous, sometimes ciliate, glabrous; awn of the lower glume 2 - 4 times as long as the glume, L E M M A S 2, dissimilar; lower lemma exceeding the glumes and resembling them, 5-9-nerved, muticous or mucronate or very shortly awned, empty or sometimes subtending a palea and sometimes also a male flower, PALEA, when present, subequal to its lemma, 2-nerved. UPPER L E M M A lanceolate, shortly apiculate or obtuse, chartaceous, subtending a palea and a bisexual flower, PALEA very similar to its lemma, LODICULES 2, broadly cuneate, often very delicate, STAMENS 3. O V A R Y ellipsoid; styles distinct, long, plumose, terminally or subterminally exserted. G R A I N oblong in outline, tightly enclosed by the somewhat hardened lemma and palea; hilum 1/3-1/2 the length of the grain. Annuals or more usually perennials, sometimes with a creeping base and rooting at the nodes; culms slender, striate, hairy or glabrous; leaf sheath longer or shorter than the internode of the culm, ribbed, ciliate on the margins and at the junction with the blade, glabrous; ligule very short, membranous; blade lanceolate, acute, sometimes acuminate, with a distinct midrib and evident lateral ribs, scabrid, glabrous; inflorescence subspicate, with distant spikelets and/or with the spikelets on very short distant branches of the main rachis; primary branches, when present, sharply 3-angled, scantily pilose.

1.

Oplismenus burmannii (Retz.) 169 (1812).

Beauv.,

Ess. Agrost. 54, 168,

PANICOIDEAE -

PANICEAE

263

2.

O. compositus (L.) Beauv., Ess. Agrost. 54, 168 (1812).

3.

O. hirtellus (L.) Beauv., Ess. Agrost. 54, 168, 170 (1812). [Fig. 107]

Panicum L., Sp. PI. 55 (1753). SPIKELETS usually solitary, more rarely in pairs, and unequally pedicelled, lanceolate, oblong, elliptic, or orbicular in outline, symmetric in profile, rarely somewhat oblique, falling entire or almost so, 1-flowered rarely 2-flowered. GLUMES unequal, more rarely equal, herbaceousmembranous; lower elliptic or broadly ovate, obtuse, very rarely 3-5fid, sometimes small and hyaline; upper as long as the lemmas, elliptic, ovate, or lanceolate, acute or obtuse, more rarely 3-4-fid or -toothed, rounded on the back, distinctly 5-9-nerved. LEMMAS 2; lower very similar to the upper glume, subtending a palea and rarely also a male flower, PALEA somewhat thinner in texture than the lemma, equalling it in length, UPPER LEMMA elliptic, ovate-elliptic or rarely oblong, obtuse or subacute, deeply concave, usually with inflexed margins, smooth or rugose, varying from membranous to subcrustaceous, glabrous, subtending a palea and a bisexual flower, PALEA similar to and subequalling its lemma, often with marginal flaps at the base, LODICULES 2, broadly cuneate. STAMENS 3. OVARY ellipsoid; styles distinct, plumose above, laterally exserted. GRAIN tightly enclosed by the hardened lemma and palea, embryo about 1/2 as long as the grain.

Annuals or perennials; culm very slender or somewhat robust, finely ribbed or smooth, sometimes channelled down one side, glabrous or pubescent, sometimes villous at the nodes; leaf sheath shorter than the internode of the culm, ribbed, glabrous, pubescent or pilose from tubercle-based hairs; ligule various, sometimes a minute line of hairs and almost obsolete, sometimes a dense fringe of hairs, or membranous and long-ciliate, or more rarely hyaline and finely ciliate; blade linearlanceolate or long-linear with a distinct or prominent midrib or the midrib not differentiated, with the side nerves distinct but sometimes distant, or closely ribbed, usually flat, more rarely longitudinally folded, or convolute and more or less terete; inflorescence a contracted or open panicle; main rachis terete or angled; primary branches often whorled on the main rachis, crowded or remote, linear and grooved, sometimes terete and ribbed or smooth, often almost capillary; ultimate branches terete, sometimes almost capillary, or linear, usually glabrous, sometimes pubescent, or with long club-shaped glandular hairs, or pilose with long scattered hairs.

264

GRASSES OF

FIG. 107. Oplismenus hirtellus (L.) Beauv. Grasses and Pastures of South Africa.)

HAWAII

Spikelets, x 5. (From D . Meredith, T h e

PANICOIDEAE -

PANICEAE

265

1. Panicum alakaiense Skottsb., Acta Hort. Gothob. 15:293, 294 ( I 944)2. P. antidotale Retz., Obs. Bot. 4:17 (1786). 3. P. beecheyi Hook, et Arn., Bot. Beechey's Voy. 100(1832). [Fig. 108] P.ramosius Hitchc., Journ. Bot. Brit, and For. 71:6 (1933). According to Chase and Niles, Index to Grasses 2:570 (1962), 'Panicum Beecheyi Hook, et Arn. misapplied by Hitchc., Mem. Bernice P. Bishop Mus. 8:184, £ 73 ( I 9 2 2 )-' P. kauaiense Hitchc., Mem. Bernice P. Bishop Mus. 8:187 (1922). 4. P. P. 5. P. 6. P.

bulbosum H.B.K., Nov. Gen. et Sp. 1:99 (1815). nodosum Willd. in Steud., Nom. Bot. ed. 2, 2:260 (1841). capillare L., Sp. PI. 58 (1753). carteri Hosaka, Occas. Papers Bernice P. Bishop Mus. 17:67 (1942). [Fig. 109]

7. P. colliei Endl., Ann. Wien Mus. Naturgesch. 1:157 (1836) without descr., based on P. affine Hook, et Arn., Bot. Beechey's Voy. 100 (1841), non Poir (1816), non Nees (1829). [Fig. n o ] P. lanaiense Hitchc., Mem. Bernice P. Bishop Mus. 8:189 (1922). 8. P. coloratum L., Mant. PI. 1 =30 (1767). 9. P. conjugens Skottsb., Acta Hort. Gothob. 15:298, 299 (1944). 10. P. cynodon Reich., Sitzungsb. Akad. Wiss. Math. Naturw. (Wien) 76:724 (1878). 11. P. degeneri Potztal, Mitteil, Bot. Gart. Mus. Berlin-Dahlem 1:1 (i953)- [Fig- i n ] 12. P. dichotomiflorum Michx., Fl. Bor. Amer. 1:48 (1803). 13. P. fauriei Hitchc., Mem. Bernice P. Bishop Mus. 8:182 (1922). [Fig. 112] 14. P. forbesii Hitchc., Mem. Bernice P. Bishop Mus. 8:199 (1922). [Fig- 113] 15. P. hillebrandianum Hitchc., Mem. Bernice P. Bishop Mus. 8:197 (1922). [Fig. 114] P. monticola Hillebr., Fl. Haw. Is. 500 (1888), non P. monticolum Hook. f. (1864). 16. P. hillebrandianum var. gracilius Skottsb., Acta Hort. Gothob. 15:296, 299 (1944). 17. P. heupueo St. John, Pacific Sei. 13:156-158 (1959).

FIG. 108. Panicum beecheyi Hook, et Arn. Plant, X 1/4; terminal portion of raceme, x 4; spikelet, X 10; sterile lemma, fertile lemma, palea with grain, all x 10. (From O. Degener, Flora Hawaiiensis.)

FIG. 109. Panicum carteri Hosaka. (a) Habit; (b) node enlarged; ( c ) ligule enlarged; (d) spikelet; (e) first glume, dorsal view; ( f ) second glume, dorsal view; ( g ) sterile lemma, dorsal view; (h) floret. (From E. Y. Hosaka, 1942. A New Hawaiian Panicum. Occas. Papers, Bernice P. Bishop Museum 17, No. 5. Reprinted with permission of The Bernice P. Bishop Museum.)

268

FIG. IIO.

GRASSES OF H A W A I I

Panicum colliei Endl. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

PANICOIDEAE -

PANICEAE

269

i2

h

to y

FIG. H I . Panicum degeneri Potztal. Plant, x 1; spikelet, x 12; outer and inner glumes, sterile lemma and palea, fertile lemma and palea, all x 10. (From O. Degener, Flora Hawaiiensis.)

270

FIG. 112.

GRASSES OF

HAWAII

Panicum fauriei Hitchc. (From A . S. Hitchcock, T h e Grasses of Hawaii.)

272

GRASSES OF H A W A I I

FIG. I 14. Panicum hillebrandianum Hitchc. (From A . S. Hitchcock, 19Z2. T h e Grasses of Hawaii.)

PANICOIDEAE -

PANICEAE

273

18. P. imbricatum Hillebr., Fl. Haw. Is. 501 (1888). [Fig. 115] 19. P. imbricatum f. minus Skottsb., Acta Hort. Gothob. 15:289, 290 (1944). 20. P. imbricatum var. molokaiense Skottsb., Acta Hort. Gothob. 15:290, 291 (1944). 21. P. isachnoides Munro, Journ. Bot. 7:178 (1869) nom. nud. ex Hillebr., Fl. Haw. Is. 501 (1888). [Fig. 116] P. conchoideum Hillebr., Fl. Haw. Is. 501 (1888). 22. P. isachnoides var. kilohanae Skottsb., Acta Hort. Gothob. 15:288 (1944). 23. P. kaalaense Hitchc., Mem. Bernice P. Bishop Mus. 8:191, 193 (1922). [Fig. 1 1 7 ] 24. P. konaense Whitney et Hosaka, Occas. Papers Bernice P. Bishop Mus. 125:3 (1936). [Fig. 118] 25. P. koolauense St. John et Hosaka, Occas. Papers Bernice P. Bishop Mus. 1 1 : 3 (1935). 26. P. laevifolium Hack., Bull. Herb. Boiss. 3:378 (1895). 27. P. maximum Jacq., Collect. Bot. 1:76 (1786); Ic. PI. Rar. 1 : 2 (1781-1786). [Fig. 119] 28. P. maximum var. trichoglume Eyles, Trans. Roy. Soc. South Afr. 5:300 (1916) without descr.; Robyns, Mem. Inst. Col. Beige 1, 6:31 (1932). 29. P. miliaceum L., Sp. PI. 58 (1753). 30. P. molokaiense Degener et Whitney, in Degener, Flora Hawaiiensis, Family 47 (Gramineae), Oct. 25 (1936). [Fig. 120] 31. P. nephelophilum Gaud., in Freyc., Voy. Uran. Bot. 4 1 1 (1830). P. havaiense Reich., Sitzungsb. Akad. Wiss. Math. Naturw. (Wien) 76:723 (1878). [Fig. 1 2 1 ] 32. P. nephelophilum beta var. tenuifolium (Hook, et Arn.) Hillebr., Fl. Haw. Is. 497 (1888). P. nephelophilum beta var. rhyacophilum (Hook, et Arn.) Hillebr., Fl. Haw. Is. 498 (1888). 33. P. niihauense St. John, Occas. Papers Bernice P. Bishop Mus. 9:5 ( I 93 1 )-

o

C/3

< S

o u fe

s 3

££ o É

FIG. 117. Panicum kaalaense Hitchc. (From A. S. Hitchcock, The Grasses of Hawaii.)

FIG. 118. Panicum konaense Whitney et Hosaka. (a) Habit; (b) ligule; (cj node; (d) spikelet; (e) first glume, dorsal view; ( f ) second glume, dorsal view; (g) sterile lemma, dorsal view; (h) floret. (From L . D . Whitney and E. Y . Hosaka. Occas. Papers, Bernice P. Bishop M u s e u m 12, No. 5. Reprinted with permission of T h e Bernice P. Bishop Museum.)

278

FIG. 119. Panicum maximum Jacq. Plant, x 1/2. 1, L i g u l e ; 2, spikelet; 3, lower glume; 4, upper glume; 5, back view of upper lemma; 6, floret; all x 4. (From N. L . Bor, T h e Grasses of Burma, Ceylon, India, and Pakistan.)

PANICOIDEAE -

PANICEAE

279

FIG. 120. Panicum molokaiense Degener et Whitney. Plant, x 1/3; portion of panicle, X 4; spikelet, x 10; glumes, lemma, and grain, all x 10. (From O. Degener, Flora Hawaiiensis.)

28O

FIG. 121. Panicum Hawaii.)

GRASSES OF H A W A I I

nephelophilum

Gaud. (From A. S. Hitchcock, The Grasses of

PANICOIDEAE -

PANICEAE

34. P. nubigenum Kunth, Enum. PI. 1 :g8 (1833). [Fig. 122] P. montanum Gaud., Voy. Uran. Bot. 411 (1830), non Roxb. (1820). Neurachne montanum Gaud., Voy. Uran. Bot. Atlas, pi. 26 (1830). 35. P. obtusum H.B.K., Nov. Gen. et Sp. 1:98 (1815). P. oahuaense Steud., Nom. Bot. ed. 2, 2:260 (1840) nom. nud. 36. P. oreoboloides (Whitney) Skottsb., Acta Hort. Gothob. 15:292, 293 (1944)P. imbricatum Hillebr. var. oreoboloides Whitney, Occas. Papers Bernice P. Bishop Mus. 13, 16:173 (1937). 37. P. oreoboloides var. subimbricatum Skottsb., Acta Hort. Gothob. 15:292, 293 (1944). 38. P. pellitoides Brown et St. John, Occas. Papers Bernice P. Bishop Mus. 1012:3 (1934) 39. P. pellitum Trin., Gram. Pan. 198 (1826), Gram. Icon. 2 (1829). P. pellitum var. pseudagrostis (Trin.) Nees, Nov. Act. Acad. Caes. Leop. Carol. 19, Suppl. 1:37 (1841), 169 (1843). P. gossypinum Hook, et Arn., Bot. Beechey's Voy. 100 (1832). P. pseudagrostis Trin., Gram. Pan. 197 (1826). 40. P. pilosum Sw., Prodr. Veg. Ind. Occ. 22 (1788). P. distichum Lamk., Tab. Encycl. Meth. Bot. 4:731 (1798). 41. P. prolutum F. Muell., Trans, and Proc. Vic. Inst. (Melbourne) 1854/1855:46 (1855). 42. P. radiatius (Vickery) St. John, Pacific Sci. 8:158, 159 (1959). Paspalidium radiatum Vickery, N.S. Wales Natl. Herb. Contr. 1 (6):332~334 (1950) non P. radiatum R. Br., Prodr. Fl. Nov. Holl. 192 (1810). 43. P. repens L. Sp. PI. ed. 2, 1:87 (1762). 44. P. scribnerianum Nash, Bull. Torrey Bot. Club 22:421 (1895). 45. P. tenuifolium Hook, et Am., Bot. Beechey's Voy. 101 (1832). [Fig. 123] 46. P. texanum Buckl., Prelim. Rept. Geol. Agr. Surv. Tex. App. 3 (1866). 47. P. torridum Gaud, in Freyc., Voy. Uran. Bot. 411 (1830). [Fig. 124] P. cinereum Hillebr., Fl. Haw. Is. 500 (1888). 48. P. virgatum L., Sp. PI. 59 (1753).

FIG. 123. Panicum tenuifolium Hook, et Arn. (From A . S. Hitchcock, T h e Grasses of Hawaii.)

284

GRASSES

OF

HAWAII

FIG. 124. Panicum torridum Gaud. Plant, x 1/2; spikelet, outer glume, sterile lemma, fertile lemma, and grain, all x 10. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

49. P. xerophilum (Hillebr.) 8:193

i^22)-

[Fig-

PANICEAE

Hitchc., M e m . Bernice P. Bishop M u s .

I25]

P. nephelophilum beta var. xerophilum Hillebr., Fl. Haw. Is. 498 (1888). Paspalidium Stapf in Prain, Fl. T r o p . Afr. 9:582 (1920). SPIKELET solitary, sessile, falling entire, FLOWERS 2. LOWER GLUME very much shorter than the spikelet, transversely oblong, hyaline; upper glume resembling the lower, LEMMAS 2 ; lower more or less ovate-elliptic, rounded on the back, deeply concave, subacute, 5-nerved, membranous, subtending a palea and a male flower, PALEA elliptic-lanceolate, flat and 2-keeled on the back, with margins inturned. LODICULES 2, broadly cuneate, fleshy, STAMENS 3. UPPER LEMMA broadly elliptic, rounded on the back, deeply concave, with inturned margins and 2 flaps at the base, scabrid on the back, LODICULES 2, cuneate. STAMENS 3. OVARY ellipsoid; styles connate at the base, plumose above, exserted from the tip of the spikelet. GRAIN tightly enclosed by the more or less hardened lemma and palea. Semiaquatic or terrestrial perennials; leaf sheath longer than the internodes of the culm, striate, glabrous; ligule a ciliated rim; blades linear, obtuse and minutely notched at the apex, with a distinct midrib, distinctly nerved, glabrous; inflorescence spikelike, consisting of spikes on a central flattened rachis; rachis of spikes flat on the back, with a much raised zigzag keel on the face; spikelets arranged alternately on either side of the keel, sometimes short secondary spikes on the rachis.

1.

Paspalidium jubiflorum (Trin.)

Hughes,

K e w Bull. 1923:317

(!923)Panicum jubiflorum Trin., Gram. Pan. 130 (1826). Paspalum L., Syst. Nat. ed. 10, 2:855 (1759). SPIKELETS solitary or paired, subsessile or unequally pedicelled, secund, and abaxial, orbicular to oblong, usually decidedly plano-convex, falling entire, FLOWER I . LOWER GLUME typically suppressed, very rarely represented by a small scale; upper conforming to the general shape of the spikelet, 3-nerved, with the nerves curved, glabrous or softly pilose. LEMMAS 2; lower equal and similar to the glume, empty; upper chartaceous to crustaceous, obtuse, emucronate, faintly nerved, sometimes with inflexed margins, sometimes minutely punctiform, faintly nerved,

286

GRASSES OF HAWAII f.

FIG. 125. Panicum xerophilum (Hillebr.) Hitchc. (From A. S. Hitchcock, T h e Grasses of Hawaii.)

PANICOIDEAE -

PANICEAE

287

glabrous, subtending a palea and the bisexual flower, PALEA similar to its lemma and tightly embraced by it, with the margins inflexed as two ears, L O D I C U L E S 2, cuneate. O V A R Y ellipsoid; styles distinct or connate at the base, plumose above, exserted near the tip of the spikelet. G R A I N tightly enclosed by the slightly hardened lemma and palea, more or less biconvex; scutellum orbicular-elliptic, shorter than 1/2 the length of the grain. Perennials; culm smooth or ribbed, glabrous; leaf sheath longer than the internode of the culm, closely ribbed, sometimes long-pilose at the junction with the blade, otherwise glabrous; ligule long, membranous; blade long-linear, flat, rarely convolute, with a distinct midrib and distant but distinct, rarely obscure, lateral veins, glabrous; inflorescence spikelike and solitary at the end of the culm or consisting of a pair of terminal spikes or spikelike racemes or of spikelike racemes arranged up a main rachis; rachis flat on the back, with a raised midrib on the face, bearing the spikelets alternately on either side of the midrib, glabrous. 1. Paspalumbertoniii/acft.inFedde,Repert. S p . N o v . 10:165 (1911). 2. P. commersonii Lamk., T a b . Encycl. Meth. Bot. 1:175 (1791). [Fig. 126] P. scrobiculatum L . var. commersonii (Lamk.) Stapf in Prain, Fl. T r o p . Afr. 9:573 (1919). 3. P. conjugatum Berg., [Fig. 127]

Act. Helv. Phys. Math. 7:129

(1772).

4. P. dilatatum Poir. in Lamk., T a b . Encycl. Meth. Bot. 5:35 (1804). [Fig. 128] P. platense Spreng., Syst. Veg. 1:247 ( J 825). P. ovatum Nees in Trin., Gram. Pan. 113 (1826). P. dilatatum var. decumbens Vasey, Bull. Torrey Bot. Club 13:166 (1886). Digitaria dilatata (Poir.) Coste, Fl. France 3:553 (1906). 5. P. distichum L., Syst. Nat. ed. 10, 2:855 (^SQ)- [Fig- 129] 6. P. fimbriatum H.B.K.,

Nov. Gen. et Sp. 1:93 (1815). [Fig. 130]

7. P. floridanum Michx.,

Fl. Bor. Amer. 1:44 (1803).

8. P. hartwegianum Fourn., Mex. PI. 2:12 (1886). 9. P. lividum Trin. in Schlect., Linnaea 26:383 (1854). 10. P. malacophyllum Trin., Gram. Icon. 3:271 (1831). 11. P. notatum Fliigge, Monogr. Pasp. 106 (1810).

288

GRASSES

OF

HAWAII

F i g . 126. Paspalum commersonii Lamk. Plant, x 1/4. 1, Ligule; 2, apical portion of a raceme showing: 2a, very short pedicels and 2b, winged axis; 3, winged axis from above, X 3. 4, Upper glume; 5, lower lemma; 6, upper lemma; 7, palea; 8, grain, all X 6. (From N . L . Bor, T h e Grasses of Burma, Ceylon, India and Pakistan.)

PANICOIDEAE - PANICEAE

289

Fig. 127. Paspalum conjugatum Berg. Plant, x 1/2; three views of a raceme showing arrangement of spikelets, x 3; two views of a floret, X 10. (From O. Degener, Flora Hawaiiensis.)

FIG. 128. Paspalum dilatatum Poir. (a) Floret, showing lemma, X 8; (b) floret, showing palea, X 8; (c) rachis, showing the 2 rows of hairy spikelets, X 4; (d) habit, showing the noticeably pubescent lowest sheaths, the arching leaves, and the spreading racemes, X 1/5; (e) upper leaf sheath, pubescent only around ligule and on base of blade, x 4. (From H. L . Mason, A Flora of the Marshes of California.)

FIG. 129. Paspalum distichum L . (a) Floret, showing palea, x 8; (b) floret, showing lemma, X 8; (c) rachis, showing 2 rows of spikelets, X 4; (d) habit, showing the decumbent rooting base, the flat leaf blades, and the paired racemes, x 1/4; (e) leaf sheath, ligule, and node, densely pubescent, x 2 J . (From H. L . Mason, A Flora of the Marshes of California.)

292

GRASSES

OF

HAWAII

FIG. 130. Paspalum fimbriatum H . B . K . Plant, X 1 / 2 ; portion of raceme showing arrangement of spikelets, X 3 ; two views of a pair of spikelets, X 1 0 ; fertile lemma with grain, X 10. ( F r o m O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

PANICEAE

293

12. P. orbiculare G. Forst., Fl. Ins. Austr. Prodr. 7 (1786). [Fig. 1 3 1 ] 13. P. plicatulum Michx., Fl. Bor. Amer. 1:45 (1803). P. plicatum Pers., Syn. PI. 1:86 (1805), error for plicatulum. 14. P. racemosum Lamk., Tab. Encycl. Meth. Bot. 1:176 (1791). P. stoloniferum Bosc., Trans. Linn. Soc. 2:83 (1794). 15. P. stramineum Nash in Britton, Man. Fl. North. United States and Canada 74 (1901). 16. P. urvillei Steud., Syn. PI. Glum. 1:24 (1854). P. larranagai Arech., Ann. Mus. Nac. Montevideo 1:60 (1894). P. vaseyanum Scrib., U. S. Dept. Agr. Div. Agrost. Bull. 17, 32, f. (1899). 17. P. vaginatum Sw., Prodr. Veg. Ind. Occ. 21 (1788). 18. P. virgatum L., Sys. Nat. ed. 10, 2:855 (1759). Pennisetum L. C. Rich., in Pers., Syn. PI. 1:72 (1805). SPIKELETS solitary or in clusters of 2-4, subtended by and deciduous with an involucre of bristles; bristles sessile or pedicelled, simple, rarely branched, naked, scabrid, or plumose, FLOWERS 1-2. GLUMES usually small and hyaline, unequal; lower glume usually suppressed; upper glume rarely 1 /2 as long as the spikelet, if longer, then up to 7-nerved. LEMMAS 2 ; lower lemma lanceolate or ovate-lanceolate, acuminate, acute, empty or subtending a palea and a male flower, PALEA subequal to its lemma, or more or less reduced, or absent, UPPER LEMMA resembling the lower, 5-7-nerved, membranous to chartaceous, subtending a palea and a bisexual flower, PALEA subequal to its lemma and of similar texture, 2-nerved. LODICULES small, usually in the front of and outside the lemma or absent, STAMENS 3, with the tips of the anthers naked or minutely penicillate. OVARY ellipsoid; style free or connate below, GRAIN broadly oblong in outline, enclosed by the more or less hardened lemma and palea; embryo 1/2-3/4 a s l° n g a s the grain.

Annuals or perennials; culm striate, glabrous; leaf sheath shorter than the internode of the culm, glabrous; ligule usually reduced to a ciliated rim or a fringe of hairs, rarely membranous; blade narrow-linear, flat or folded, closely veined, with a prominent midrib, scabrid, usually glabrous, more rarely with scattered bulbous-based hairs; inflorescence a solitary terminal cylindrical or more or less oblong subspike; main rachis straight or flexuous, furrowed.

294

GRASSES OF

HAWAII

F i g . 131. Paspalum orbiculare Forst. Plant, x 1/2; floret, fertile lemma and grain, X 10. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

1.

2.

PANICEAE

295

Pennisetum clandestinum Höchst, ex Chiov., Ann. Ist. Bot. Roma 8:41 (1903), and in Leeke, Zeitschr. Naturw. 79:23 (1907). [Fig- 132] P. complanatum (Nees) Hemsl., Biol. Centr. Amer. Bot. 3:507 (1885).

3.

P. macrostachyum (Brongn.) ser. 6, 1 : 1 7 7 (1834).

Trin., Mem. Acad. Sei. Petersb.

4.

P. Orientale (Willd.) L. C. Rich, in Pers., Syn. PI. 1 7 2 (1805). Panicum Orientale{Rich.) Willd., Enum. PI. Hort. Berol. 1031 (1809).

5.

P. purpureum Schumach., Beskr. Guin. PI. 44 (1827). [Fig. 133]

6.

P. schimperi Steud., Nom. Bot. ed. 2, 298 (1841) nom. nud., Steud. ex A. Rich., Tent. Fl. Abyss. 2:381 (1851). P. merkeri Leeke, Zeitschr. Naturw. 79:27 (1907).

7.

P. setaceum (Forsk.) Chiov., Soc. Bot. Italy Bull. 1 9 2 3 : 1 1 3 (1923). [Fig. 134] P. asperifolium (Desf.) Kunth, Rev. Gram. 1:49 (1829). P. ruppelii Steud., Syn. PI. Glum. 1 : 1 0 7 (1854). P. setosum (Sw.) L. C. Rich, in Pers., Syn. PI. 1 : 7 2 (1805). [Fig- 13S] Cenchrus setosus Sw., Prodr. Veg. Ind. Occ. 26 (1788). P. typhoides (Burm.) Stapf et C.E. Hubb., Kew Bull. 1933:271

8.

9.

(!933)See Bor (i960), pp. 350-351 for a discussion on the current state of confusion about the proper name for the 'pearl millet.' There is a very good possibility that the proper name for pearl millet is P. americanum (L.) K . Schum. instead of P. typhoides (Burm.) Stapf et C.E. Hubb. P. typhoideum L . C . Rich, in Pers., Syn. PI. 1 ¡72 (1805). 10. P. villosum R. Br., in Fresn., Mus. Senckenb. 2 : 1 3 4 (1837); in Salt, Voy. Abyss. App. 62 (1814) nomen. Rhynchelytrum Nees in Lindl., Nat. Syst. ed. 2, 446 (1836). SPIKELETS solitary, pedicelled, falling entire, often completely enveloped by long shining hairs; pedicels capillary or sometimes narrowlinear, with thickened tips, FLOWERS 1 - 2 . GLUMES very dissimilar or at least unequal; lower glume reduced to a minute scale or obsolete, sometimes remote from the upper glume and 5-(rarely 7~)nerved; upper glume more or less dorsally gibbous and laterally compressed, sometimes

FIG. 132. Pennisetum clandestinum Höchst, ex Chiov. A, Plant, X 1/2; B, terminal flowering branch, x 1 ; C, dissected panicle, X 1.5; D, fascicle, X 1.5. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

FIG. 133. Pennisetum purpureum Schumach. Plant, X 1/2. i, supporting bristles, X 3; J, spikelet ( j a , lower glume, 3b, lemma of upper floret, X 3; 5, palea, X 3; 6, stamens and bearded at the tip of the anther cells, x 10. (From H . L . Bor, Ceylon, India and Pakistan.)

Ligule; 2, spikelet and upper glume), X 3; 4, ovary, x 5; 7, stamen, T h e Grasses of Burma,

298

FIG. 134. Pennisetum setaceum (Forsk.) Chiov. A, Plant, x 1/4; B, fascicle, X 3; C, spikelet, x 10. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

FIG. 135. Pennisetum setosum (Sw.) L. C. Rich. A, Plant, X 1/4; B, fascicle, x 3; C, floret, x 10; D, spikelet, x 10. (From an original drawing in the files of the Dept. of Agronomy and Soil Science, Univ. of Hawaii.)

3oo

GRASSES OF H A W A I I

evenly curved to almost straight on the keel, ovate, oblong-ovate, or lanceolate, 2-toothed, sometimes mucronate or shortly awned from the sinus, 5-nerved, hairy with long silky hairs, 5-nerved; awn, when present, 2/3 length of the glume, LEMMAS 2; lower lemma more or less like the upper glume, subtending a palea and sometimes also a male flower, PALEA subequalling its lemma, dissimilar or more or less similar to it, 2-nerved, hyaline, UPPER LEMMA ovate-oblong, obtuse or acute, more rarely truncate and minutely notched, chartaceous, subtending a palea and a bisexual flower, PALEA similar to its lemma, 2-nerved, chartaceous. LODICULES 2. STAMENS 3. OVARY ellipsoid; styles free,

plumose, laterally exserted. GRAIN closely embraced by the lemma and palea; embryo about 1/2 the length of the grain. Perennials or annuals; culm striate, pilose with bulbous-based hairs or glabrous, sometimes villous at the nodes; leaf sheath shorter than the internode of the culm, ribbed, pilose with bulbous-based hairs; ligule ciliate; blade flat or rolled, sometimes setaceous, usually narrowlinear, more rarely linear-lanceolate, distinctly or indistinctly veined, pilose or glabrous; inflorescence a lax or more or less contracted panicle; rachis linear, pilose. 1.

Rhynchelytrum repens (Willd.) C.E. Hubb., K e w Bull. 1 9 3 4 : 1 1 0 (1934). [Fig. 136] Saccharum repens Willd., Sp. PI. 1 : 3 2 2 (1798). Tricholaena rosea Nees, Ind. Sem. Hort. Vratisl. (1835), and in Linnaea 1 1 :Litt.-Bericht 129 (1837). Tricholaena repens (Willd.) Hitchc., Man. Grasses W. Indies, U. S. Dept. Agr. Misc. Publ. 2 1 3 , 331 (1936). Rhynchelytrum roseum (Nees) Stapf et C.E. Hubb. ex Bews, T h e World's Grasses 223 (1929), and in Prain, Fl. Trop. Afr. 9:880

( I 93 0 )-

Sacciolepis Nash ex Britton, Canada 89 (1901).

Man. Fl. North. United States and

SPIKELETS solitary, pedicelled, usually more or less oblique at the base, falling entire; pedicel short, finely filiform, FLOWERS 1 - 2 . GLUMES very unequal; lower reduced to a minute hyaline scale; upper curved or basally gibbous or saccate on the back, concave, usually 7-9-nerved, sometimes 5-13-nerved, with hyaline margins or tip. LEMMAS 2, dissimilar; lower resembling the upper glume but with a straight back, subtending a palea and sometimes a male flower, PALEA linear, hyaline.

PANICOIDEAE -

PANICEAE

FIG. 136. Rhynchelytrum repens (Willd.) C . E. H u b b . floret, x 3. (From O . Degener, Flora Hawaiiensis.)

301

Plant, X 1/2; spikelet, X 3;

302

GRASSES OF H A W A I I

LEMMA oblong or elliptic, convex, chartaceous, becoming crustaceous, with firm narrowly involute margins, obscurely 5-nerved, subtending a palea and a bisexual flower, PALEA more or less similar to its lemma, LODICULES 2, broadly cuneate. STAMENS 3. OVARY ellipsoid; styles distinct, plumose near the ends, exserted from the apex of the spikelet. GRAIN tightly enclosed by the somewhat hardened lemma and palea, elliptic in outline. Annuals or perennials; culm slender, ribbed, glabrous; leaf sheath shorter than the internode of the culm, prominently ribbed, glabrous; ligule a very narrow minutely ciliate rim; inflorescence a narrow evident panicle. UPPER

1.

Sacciolepis indica (L.) Chase, Proc. Biol. Soc. Washington 21:8 (1908). Aira spicata L., Sp. PI. 63 (1753). A. indica L., Sp. PI. ed. 2, 94 (1762); indica is a change of name for A. spicata Sp. PI. 63 (1753). Panicum contractum Wight et Arn. ex Nees in Linneae io:Litt.Bericht 1 1 7 (1836). Sacciolepis contracta (Wight et Arn.) Hitchc., Mem. Bernice P. Bishop Mus. 8:199 (1922).

2.

S. striata (L.) Nash, Bull. Torrey Bot. Club 30:383 (1903). Panicum aquaticum Muhl., Descr. Gram. 126 (1817), non Poir. (1816).

Setaria Beauv., Ess. Agrost. 51 (1812) nom. cons., non Michx. (1803). Chaetochloa Scribn., U. S. Dept. Agr. Div. Agrost. Bull. 4, 38 (1897). SPIKELETS solitary or clustered, subsessile, falling entire, subtended by one or more persistent bristles, more rarely some spikelets without bristles, BRISTLES shorter than the spikelet or up to 4 times as long, smooth or scabrid. FLOWERS 1-2. GLUMES unequal or almost equal, membranous; lower generally much smaller than the upper, ovate from a clasping base, usually 3-5-(rarely 1- or 7-)nerved; upper usually 3-9-nerved. LEMMAS 2, dissimilar; lower more or less exceeding and resembling the upper glume, empty or subtending a palea and sometimes also a male flower, UPPER LEMMA elliptic, deeply concave, subacute, transversely rugose or pitted, 5-nerved, chartaceous to coriaceous, subtending a palea and a bisexual flower, PALEA similar to its lemma

PANICOIDEAE - PANICEAE

303

and equalling it. LODICULES 2, broadly cuneate. STAMENS 3. OVARY ellipsoid; styles distinct, plumose above, laterally exserted. GRAIN oblong or elliptic in outline, tightly enclosed by the hardened lemma and palea; embryo about 1 ¡2 as long as the grain. Perennials or annuals, sometimes stoloniferous; culm smooth or ribbed, sometimes channelled down one side, glabrous; leaf sheath longer or shorter than the internode of the culm, ribbed, with membranous margins, sometimes ciliate, glabrous, more rarely pilose; ligule ciliate; blade usually flat and narrow-linear, sometimes rolled or longitudinally folded, closely veined, with a prominent or distinct midrib, smooth or scabrid, glabrous; inflorescence either a cylindrical spikelike panicle, or an open panicle, or more rarely subspicate; rachis of open inflorescence angled, minutely pubescent; main rachis of spikelike inflorescence sometimes villous; bristles golden, tawny, or dark purple in color. 1. Setaria barbata (Lamk.) Kunth, Rev. Gram. 1:47 (1829). Panicum barbatum Lamk., Tab. Encycl. Meth. Bot. 1:171 (1791). Chaetochloa barbata (Lamk.) Hitchc. et Chase, Contr. U. S. Natl. Herb. 18:348 (1917). 2. S. geniculata (Lamk.) Beauv., Ess. Agrost. 51, 169, 178 (1812). [ F i g- J 37] Panicum imberbe Poir. in Lamk., Tab. Encycl. Meth. Bot. Suppl. 4:272 (1816).

3.

P. geniculatum Poir. in Lamk., Tab. Encycl. Meth. Bot. 4:727 (err. tyP- 737) (!798). S. purpurascens H.B.K., Nov. Gen. et Sp. 1:110 (1815). Chaetochloa geniculata (Lamk.) Millsp. et Chase, Field Mus. Bot. 3:37 ( I 9°3)S. glauca (L.) Beauv., Ess. Agrost. 51, 169, 178 (1812). [Fig. 138] Chaetochloa glauca(L.) Scribn., U. S. Dept. Agr. Div. Agrost. Bull. 4, 39 (1897).

C. lutescens (Weig.) Stunz, U. S. Dept. Agr. Bur. Plant Indus. Inventory Seeds 31:36, 86 (1914). Pennisetum glaucum (L.) R. Br., Prodr. Fl. Nov. Holl. 1:195 (1810). Panicum lutescens Weig., Obs. Bot. 20 (1772). S. lutescens (Weig.) F.T. Hubb., Rhodora 18:232 (1916). Panicum glaucum L., Sp. PI. 56 (1753). See Bor (I960), pp. 360, 361; Reeder, Rhodora 53:27 (1951); and Stapf, Kew Bulletin 147 (1928) for reasons why the name S.

FIG. 137. Setaria geniculata (Lamk.) Beauv. (a) Spikelet, showing fertile lemma and the few upwardly barbed bristles on branchlet, X 1 2 ; (b) floret, showing palea, x 12; (c) spikelet, showing first glume and sterile lemma, x 12; (d) rachilla, the bristles remaining, the spikelets having fallen off from the branchlets, x 6; (e) leaf sheath, showing long-ciliate ligule and the sparsely set, long hairs at base of the scabrous blade, x 6; ( f ) habit, upper part, showing the slender linear panicles, x 2/5; (g) habit, lower part, showing the knotty, branching rhizomes and the erect, ascending leaf blades, X 2/5. (From H. L . Mason, A Flora of the Marshes of California.)

FIG. 138. Setaria glauca (L.) Beauv. (a) Spikelet, showing first glume and sterile lemma, x 10; (b) spikelet, showing fertile lemma and the upwardly barbed, slender bristles on branchlet, x 10; (c) floret, showing palea, x 10; (d) leaf sheath and ciliate ligule, x 6; (e) habit, upper part of culm, showing spikelike panicle, x 2/5; (f) habit, lower part, showing the leaf blades with villous base above sheath, x 2/5. (From H. L. Mason, A Flora of the Marshes of California.)

3O6

GRASSES OF H A W A I I

lutescens (Weig.) F.T. Hubb., as the species is known in the United States, should be dropped in favor of S. glauca (L.) Beauv. 4.

S. italica (L.) Beauv., Ess. Agrost. 51, 170, 178 (1812).

5.

S. longiseta Beauv., Fl. Oware 2:81 (1818).

6.

S. palmifolia (Koen.) Stapf, Journ. Linn. Soc. Bot. 42:186 (1914). Panicum nervosum Roxb., Hort. Beng. 8 (1814) without descr., Fl. Ind., ed. Carey, 1 1 3 1 4 (1820), non Lamk. (1798). Panicum neurodes Schult., Syst. Veg. 2: Mant. 228 (1824). Chaetochloa palmifolia (Willd.) Hitchc. et Chase, Contr. U. S. Natl. Herb. 18:348 (1917).

7.

S. sphacelata (Schumach.) Stapf et C.E. Hubb. ex M.B. Moss, Kew Bull. 1929:195 (1929) without descr., Stapf and C.E. Hubb. in Prain, Fl. Trop. Afr. 9:795 (1930).

8.

S. verticillata (L.) Beauv., Ess. Agrost. 51, 171, 178 (1812). [Fig- 139] Panicum verticillatum L., Sp. PL ed. 2, 1:82 (1762). Chaetochloa verticillata (L.) Scribn., U. S. Dept. Agr. Div. Agrost. Bull. 4:39 (1897). Pennisetum verticillatum (L.) R. Br., Prodr. Fl. Nov. Holl. 195 (1810).

Stenotaphrum Trin., Fund. Agrost. 175 (1820). SPIKELETS solitary or in fascicles of 2-5, subsessile or on short appressed branches, immersed in hollows on the rachis or very closely appressed to the upper side of the rachis, falling entire, FLOWERS 1-2. GLUMES unequal; lower glume varying from very minute and scale-like to nearly 1/2 the length of the spikelet, broadly truncate, obtuse, or emarginate, nerveless; upper glume oblong-obovate, subobtuse, very convex on the back, as long as or nearly as long as the spikelet, 5-9nerved, glabrous, LEMMAS 2, dissimilar, lanceolate or ovate-lanceolate, acute, glabrous; lower lemma chartaceous to coriaceous, 3-9-nerved, subtending a palea and usually a male flower, PALEA as long as its lemma, membranous, UPPER LEMMA chartaceous to coriaceous, or membranous, subtending a palea and a bisexual flower, PALEA equal to and tightly embraced by its lemma, faintly 2-nerved. LODICULES 2, quadrate-cuneate. STAMENS 3. OVARY ellipsoid; style free or almost so, plumose above, laterally exserted. GRAIN oblong in outline, plano-convex; embryo 1/2 as long as the grain.

FIG. 139. Setaria verticillata (L.) Beauv. (a) Spikelet, showing short first glumes, and the single, downwardly barbed bristles on branchlets, X 16; (b) floret, showing l e m m a , X 1 6 ; (c) auricles, leaf sheath, and ciliate ligule, X 4; (d) panicle, X 4/5; (e) floret, showing palea, X 16; ( f ) habit, upper part, showing panicles, X 1/5; (g) habit, lower part, showing the lax, arching leaf blades and roots at the nodes, X 1/5. (From H. L . Mason, A Flora of the Marshes of California.)

3O8

GRASSES OF H A W A I I

Creeping or prostrate perennials, rooting at the nodes, sometimes tufted annuals; culm erect, striate, glabrous; leaf sheath shorter than the internode of the culm, longitudinally folded, closely ribbed, glabrous or pilose at the junction with the blade; ligule a fringe of short hairs; blade flat, linear, closely veined, with a distinct midrib, scabrid, glabrous; inflorescence spikelike, solitary and terminal or lateral from the upper leaves; rachis compressed, flat on the back, hollowed on the inner face; branches short, terminated by an oblique appendage. i.

Stenotaphrum secundatum (Walt.) Kuntze, Rev. Gen. PI. 2:794 (1891). [Fig. 140] Ischaemum secundatum Walt., Fl. Carol. 249 (1788). S. americanum Schrank, PI. Rar. Hort. Monaco, pi. 98 (1822). S. glabrum Trin., Fund. Agrost. 176 (1820). Rottboellia stolonifera Poir. in Lamk., Tab. Encycl. Meth. Bot. 6:310 (1804). S. secundum (Osphalm.) Kuntze, Rev. Gen. PI. 2:794 i 1 ^ 1 ) -

Trichachne (L.) Nees, Agrost. Bras. 86 (1829). Valota Adans., Fam. 2:495 ( I 7^3) nom. nud. SPIKELETS lanceolate, in pairs, short-pedicelled, in two rows along one side of a slender rachis. FIRST GLUME minute, glabrous, SECOND GLUME and sterile lemma about as long as the fruit, 3-5-nerved, copiously silky, FERTILE LEMMA cartilaginous, lanceolate, acuminate, usually brown, the flat white hyaline margins broad. Perennials with slender erect or ascending racemes, approximate to rather distant along a slender main axis, forming a white to brownish silky panicle. 1.

Trichachne californica (Benth.) Chase, Journ. Washington Acad. Sei. 23:455 (1933). Panicum lachnanthum Torr., U. S. Expl. Miss. Pacif. Rept. 7:21 (1858), non Höchst. (1855).

2.

T. insularis (L.) Nees, Agrost. Bras. 86 (1829). [Fig. 141] Andropogon insularis L., Syst. Nat. ed. 10, 2:1304 (1759). Panicum lanatum Rottb., Act. Lit. Univ. Hafn. 1:269 (1778). P. leucophaeum H.B.K., Nov. Gen. et Sp. 1:97 (1815). Valota insularis (L.) Chase, Biol. Soc. Washington Proc. 19:188 (1906).

FIG. 140. Stenotaphrum secundatum (Walt.) Kuntze. inflorescence, X 5. (U.S.D.A. photograph.)

1, Plant, X 1/2; 2, portion of

3io

GRASSES

OF

HAWAII

FIG. 1 4 1 . Trichachne insularis (L.) Nees. Plant much reduced; ligule, x 1/2; spikelet, x 5. (From O. Degener, Flora Hawaiiensis.)

PANICOIDEAE -

PANICEAE

3"

Urochloa Beauv., Ess. Agrost. 52 (1812). SPIKELETS solitary, in pairs, or in fascicles of 3-4, sessile, subsessile, or pedicelled, secund and abaxial on the rachis, falling entire from the pedicels; pedicels usually much reduced and disc-tipped or, if in pairs, then the primary slightly longer, FLOWERS 1-2. GLUMES similar and subequal or more often dissimilar and very unequal in length; the lower a small nerveless hyaline scale, 3-5-nerved; upper equalling the lemmas, ovate, acuminate, rounded on the back, deeply concave, obtuse, or acute, 5-7-nerved, pilose, pubescent, or glabrous, LEMMAS 2, very dissimilar; lower very much like the upper glume and more or less equalling it, subtending a palea and sometimes a male flower, PALEA subhyaline, equalling its lemma, UPPER LEMMA broadly elliptic to suborbicular, abruptly mucronate, with inflexed margins, concave and rugose on the back, crustaceous, subtending a palea and a bisexual flower; mucro sometimes almost 1/2 as long as the lemma, PALEA subsimilar to its lemma but rounded at apex, with 2 marginal flaps at the base, LODICULES 2, broadly cuneate. STAMENS 3. OVARY subglobose; styles distinct, plumose above, laterally exserted. GRAIN tightly enclosed by the more or less hardened lemma and palea; embryo 1/2-3/4 a s l ° n g a s the grain. Annuals or perennials; culm ribbed, sometimes channelled down one side, glabrous or sometimes sparsely pilose, villous at the nodes, leaf sheath ribbed, sometimes pilose with bulbous-based hairs; ligule ciliate; blade lanceolate to linear-lanceolate, finely veined, with or without a distinct midrib, often with undulated margins, often long-ciliate, scabrid, sparsely pilose with bulbous-based hairs; inflorescence of 3 - 1 2 spikes or spikelike racemes arranged alternately on main rachis; rachis flattened on the back, with a raised narrow or broad keel on the inner face, minutely ciliate on the margins, usually with a few long hairs at the nodes.

1.

Urochloa bolbodes (Steud.) Stapf in Prain, Fl. Trop. Afr. 9:593 (1920). Panicum numidianum Hack., Bull. Herb. Boiss. 4: App. 3, 14 (1896), non Lamk. (1791).

2.

U. panicoides Beauv., Ess. Agrost. 53 (1812), var. panicoides Bor 372 (i960).

3.

U. pullulans Stapf in Prain, Fl. Trop. Afr. 9:590 (1920).

4.

U. stolonifera (Goossens) L. Chippindall, Grasses and Pastures S. Afr. 381-382 (1955). Brachiaria stolonifera Goossens, Kew Bull. 1934:195 (1934).

COMMON NAMES, ORIGINS, AND DATES OF FIRST RECORD OF HAWAIIAN GRASSES In Table 3 are listed in alphabetical order approximately 500 species of grasses found in Hawaii, together with the native country or putative origin of each, its common names, and the date when its presence was first recorded in the islands. An attempt has been made to list all of the local common names, including especially any Hawaiian names, for each grass. T h e date of first record in Hawaii is given, rather than an approximate date of introduction, since in many cases it is impossible to determine when the grass was originally introduced. Many were brought in on different occasions and under various names; others came in as impurities in seed purchased for use in the islands. Since these records are incomplete, a date given in the table may represent an actual date of introduction, a first collection date, an approximate date when first observed, etc. This information has been compiled from the records and collections in the herbaria of the Hawaii Agricultural Experiment Station, T h e Bernice P. Bishop Museum, the Hawaiian Sugar Planters' Association, as well as from private individuals who were responsible for the introduction or the collection of certain species. A question mark (?) in this column indicates that there is no date of first record for that particular species. A native country is not listed for a number of the grasses, inasmuch as there is often considerable doubt as to the actual place of origin. In these cases a geographic area is indicated instead. A knowledge of the area of origin is of much value in predicting the ability of a species to become widespread in different environments. The reader is referred to the bibliography, in particular to Hitchcock (1922) and Whitney et al. (1939) for further information about dates of introduction, places of first collection, distribution, etc.

C O M M O N NAMES, O R I G I N S , A N D FIRST RECORDS

3r3

TABLE 3 L I S T OF HAWAIIAN GRASSES WITH COMMON NAMES, ORIGIN, AND DATE OF FIRST RECORD PUTATIVE BOTANICAL NAME

COMMON NAME

Agropogon littoralis

FIRST

NATIVE COUNTRY

RECORD

Europe

Agropyron cristatum dasystachyum

crested wheatgrass thickspike wheatgrass

desertorum elongatum inerme

desert wheatgrass tall wheatgrass beardless wheatgrass

intermedium repens smithii spicatum

intermediate wheatgrass quackgrass, quitchgrass, couchgrass western wheatgrass bluebunch wheatgrass

trachycaulum

slender wheatgrass

trichophorum

stiffhair wheatgrass, pubescent wheatgrass

Agrostis alba avenacea

DATE OF

ORIGIN OR

red top, herdsgrass heu-pueo

borealis canina microphylla fallax palustris sandwicense scabra

Arctic bent velvet bent spike ledtop

semiverticillata stolonifera tenuis

water bent marsh bent colonial bent

creeping bent Hawaiian bent

Eurasia northwestern United States Russia Europe western United States Europe Eurasia United States western United States western United States

? 1936 1922 ? ? J

939 ?

J

939 1911

1911 1911

Europe Eurasia Australia and through Polynesia United States Europe North America Hawaiian Islands Eurasia Hawaiian Islands Canada and northern United States Europe Europe Europe

!9!S ?

1912 1906 1938 ?

1864 1913 *933

3H

GRASSES OF

TABLE 3

BOTANICAL NAME

Aira caryophyllea

HAWAII

(continued)

COMMON NAME

silver hairgrass

Alloteropsis cimicina

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Europe

1906

India, tropical Africa

1916

Alopecurus pratensis

meadow foxtail

Eurasia

I

Ammophila arenaria

European beachgrass

Europe

!9!3

Andropogon bicornis gerardi

bluejoint turkeyfoot

1955

bushy beardgrass turkeyfoot, sand bluestem seacoast bluestem broomsedge, yellow bluestem

Latin America western United States North America United States United States eastern United States

sweet vernalgrass

Europe

1906

Europe

1938

glomeratus halli littoralis virginicus Anthoxanthum odoratum Apera interrupta

9I3

1916 1964 1940 ? 1932

Aristida adscensionis

sixweeks threeawn

American tropics

1938

Arrhenatherum elatius

tall oatgrass

Europe

1906 ?

Arundinella holcoides Arundo donax Astrebla elymoides lappacea pectinata Avena barbata byzantina

Burma giant reed, Spanish reed

Old World

1906

mitchellgrass

Australia Australia Australia

1938 !939 W3

slender wildoat

Europe Europe

1906 1935

COMMON N A M E S , O R I G I N S , AND F I R S T RECORDS

TABLE 3

BOTANICAL NAME

315

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

fatua sativa Axonopus affinis compressus Beckmannia syzigachne Bothriochloa ambigua barbinodis caucasico insculpta intermedia ischaemum perforata pertusa saccharoides Bouteloua chondrosioides

wildoat culivated oat

Europe Europe

1906 1909

narrow-leaved carpetgrass

southern United States

1912

America

l

broad-leaved carpetgrass, carpetgrass

939

American sloughgrass

fuzzy top

silver beardgrass, silver bluestem gramagrass

curtipendula

side-oats grama

eriopoda

black grama

filiformis

slender grama

gracilis

blue grama

heterostega hirsuta

hairy grama

parryi

Parry grama

1922 Australia Mexico Eurasia tropical Africa Australia India, North Africa southwestern United States and Mexico Old World tropics

? 1906 ? 1940 1906 1935 1918 1916

Jamaica western United western United western United western United western United Antilles western United western United

States

1937

States

1906

States

X

States

J

States

1922

93S

937 ?

States

1935

States

J

937

3i6

GRASSES OF H A W A I I

TABLE 3

(continued) PUTATIVE

BOTANICAL NAME

rothrockii Brachiaria brizantha ciliatissima dictyoneura mutica plantaginea reptans

COMMON NAME

Rothrock grama

palisadegrass perennial signalgrass paragrass, panicumgrass, californiagrass

Brachypodium distachyon

FIRST

NATIVE COUNTRY

RECORD

western United States

Africa Africa cosmopolitan tropics

1902 1916 I 937

Europe

1936 1932 1840

Europe Europe

Bromus breviaristatus

short-awned bromegrass

western United States Pacific coast of United States Europe Europe Europe Europe North America Europe Europe western United States Europe Europe Europe Europe Europe Europe Europe South America

commutatus erectus hordeaceus inermis marginatus molliformis mollis polyanthus

hairy chess meadow brome

racemosus rigidus var. gussonei rubens squarrosa sterilis tectorum unioloides

smooth brome soft chess

ripgutgrass foxtail chess downychess, cheatgrass rescuegrass

93S

1938 1 935

big quakinggrass little quakinggrass

California brome

I

tropical Africa southwestern United States tropical Africa

Briza maxima minor

carinatus

DATE OF

ORIGIN OR

1938

1936 !939 9I3

I

p 1910 J 939 1906 1906 !939 1906 1906 1906 1906 1906 1920 1888 1906

COMMON NAMES, O R I G I N S , A N D FIRST RECORDS

TABLE 3

(continued) PUTATIVE

BOTANICAL NAME

COMMON NAME

vulgaris Buchloè dactyloides Calamagrostis canadensis var. scabra expansa hillebrandii langsdorffii

317

American buffalograss

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

western United States

1922

western United States

1936 ?

large Hawaiian reedgrass Hillebrand's reedgrass

United States Hawaiian Islands Hawaiian Islands Europe (?)

kamanomano, kumanomano, mauu-kuku agrimonioides Hawaiian Islands agrimony sandbur Leeward Islands var. laysanensis biflorus India, North Africa birdwoodgrass ciliaris buffelgrass Africa echinatus sandbur, umealu, konpeitoquaa tropical America Hillebrand's sandbur, hillebrandianus Hawaiian Islands hairy sandbur Hawaiian Islands woolly Waianae sandbur pedunculata northwest India, setigerus northeast tropical Africa

?

Cenchrus

Chloris barbata

ntauu-lei, swollen fingergrass

berroi ciliata culcullata distichophylla divaricata gayana polydactyla

weeping chloris stargrass rhodesgrass

tropical America South America southwestern United States and Mexico southwestern United States South America Australia Africa South America

!939 1935 1867

1906 1936

?

!939 1932 1906 1914

3I8

GRASSES OF H A W A I I

TABLE 3

BOTANICAL NAME

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

radiata truncata virgata Chrysopogon aciculatus

radiate fingergrass Australian fingergrass feather fingergrass

tropical America Australia tropical America

1851 1906 1916

East Indies, Hawaiian Islands tropics of Asia and Africa

!9!3

Job's tears, ohe puoheoke, pupu-kolea

East Indies

1895

pampasgrass

South America

J

lemongrass citronellagrass barbwiregrass, soapgrass

East Indies India Australia North Africa, India

1906 1936 1936 ?

South India, Southeast Asia Madras

J

African bermudagrass

Old World Africa Africa Africa

1835 1961 !937 1961

crested dogtail

Europe

1913

orchardgrass, cocksfoot

Europe

1906

beach wiregrass

Old World tropics

1906

hairy oatgrass

Australia Australia Australia

1910 1937 1903

pilipili-ula, piipii, maniania-ula manienie-ula

fulvus Coix lacryma-jobi Cortaderia selloana Cytnbopogon citratus nardus refractus schoenanthus Cynodon arcuatus barberi dactylon magennisii plectostachyus transvaalensis Cynosurus cristatus Dactylis glomerata Dactyloctenium aegyptium Danthonia pilosa racemosa semiannularis

bermudagrass, manienie, mahiki, manienie-haole

wallabygrass

925

1918 937

COMMON NAMES, ORIGINS, AND FIRST RECORDS

TABLE 3

BOTANICAL NAME

319

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

Deschampsia australis f. haleakalensis ssp. nubigena var. gracilis var. gracilis f. major var. gracilis f. media var. gracilis f. minor caespitosa

tufted hairgrass

elongata

slender hairgrass

flexuosa Dichanthium annulatum aristatum sericeum

exilis filiformis ftiscescens glauca

Islands Islands Islands Islands Islands Islands Islands

crinkled hairgrass

temperate regions of Old World U. S. (Pacific northwest) Eurasia

1935 I9J3

angletongrass wildergrass Australian bluegrass

Africa, India Asia Australia

1927 I9I3 1912

South Pacific New Zealand

?

Dichelachne montana sciurea Digitaria adscendens argyrograpta bicornis brownii debilis decumbens diagonalis didactyla divaricatissima diversinervis eriantha eriostachya

Hawaiian Hawaiian Hawaiian Hawaiian Hawaiian Hawaiian Hawaiian

Henry's crabgrass

pangolagrass blue couchgrass

creeping kukae-puaa

Formosa Africa tropical Asia Australia Africa Africa South Africa Mascarene Islands Australia Africa Africa South America, Paraguay Africa Java Madagascar

I9I3

1952 1932 ?

? ?

I93I I9SO ?

1916 195° 1964 1939 ?

1923 1936 1938 ?

32°

GRASSES OF

TABLE

3

HAWAII

(continued) PUTATIVE

BOTANICAL NAME

herpoclados horizontalis iburua ischaemum kilimandscharica longiflora macroglossa microbachne milanjiana ssp. eylesiana monodactyla natalensis pentzii phaeothrix pruriens sanguinalis seriata smutsii swazilandensis timorensis tricholaenoides uniglumis valida violascens

COMMON NAME

smooth crabgrass

kukae-puaa, liilii

woolly fingergrass

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Africa New World tropics Nigeria, Africa cosmopolitan in temperate regions, Africa India Africa probably Philippine Islands Africa Africa South Africa South Africa Africa Uruguay, South America

kukae-puaa, mauu-kukaeJava or Sumatra puaa kukae-puaa, large crabgrass Europe tropical Africa Africa swazigrass, Swaziland Africa fingergrass Southeast Asia, Polynesia South Africa Abyssinia Africa giant pangolagrass tropical Asia kukae-puaa-uka

Dissochondrus biflorus Echinochloa colonum crusgalli cruspavonis frumentacea polystachya

DATE OF

I

943 1916 ?

1964 1917 I 93S 1964 p ?

1935

1826 1864 ? 1948 1948 195° 1964 ? T

935 1917

Hawaiian Islands jungle-rice barnyardgrass

India Eastern Hemisphere Venezuela Uruguay

1864 1864

COMMON N A M E S , O R I G I N S , AND F I R S T RECORDS

TABLE 3

BOTANICAL NAME

32I

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

Africa eastern United States

1920

Africa Africa

1936 1936

Old World tropics

J

India

1840

I93S

blue wild-rye beardless wild-rye Virginia wild-rye

North America western United States North America North America United States

pappusgrass

Arabia

1936

tropical Africa

1959

stagnina walteri Ehrharta calycina erecta Eleusine coracana indica Elymus canadensis condensatus glaucus (riticoides virginicus Enneapogon brachystachyus

perennial veldtgrass panic veldtgrass

goosegrass, wiregrass, manienie-alii Canada wild-rye giant wild-rye

Enteropogon macrostachya Eragrostis atropioides brownei chloromelas cilianensis curvula deflexa elongata falcata fosbergii grandis var. oligantha var. polyantha hosakai intermedia

hard-stemmed lovegrass Brown's lovegrass, sheepgrass Boer lovegrass stinkgrass weeping lovegrass

Fosberg's lovegrass large Hawaiian lovegrass

plains lovegrass

r

935

939

1912 1922 1936

Hawaiian Islands Australia South Africa Europe South Africa Hawaiian Islands India Australia Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands southwestern United States

1906

I95 1

1864 1916 1949 .

?

?

322

GRASSES OF H A W A I I

TABLE 3

(continued) PUTATIVE

BOTANICAL NAME

lehmanniana leptophylla leptostachya tnauiensis mexicana

COMMON NAME

Lehmann lovegrass mountain lovegrass Maui lovegrass Mexican lovegrass

montícola neomexicana niihauensis paupera pectinacea

Niihau lovegrass

pilosa plana

India lovegrass

Carolina lovegrass

poaeoides superba tef tenella trichodes unioloides variabilis whitneyi var. caumii

Tef lovegrass Japanese lovegrass sand lovegrass

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Africa Hawaiian Islands Australia Hawaiian Islands Central America, West Indies Hawaiian Islands southwestern United States, Mexico Hawaiian Islands Hawaiian Islands eastern United States Europe tropical Africa and South Africa India to Mediterranean region tropical Africa Africa Old World tropics central United States Southeast Asia

x

935 ?

?

1940

1906

1916 ? ?

1906 ?

!939 p

variable lovegrass, emoloa, kalamalo, kawelu Hawaiian Islands

Hawaiian Islands

Eremochloa ophiuroides

centipedegrass

China

1920

Erianthus ravennae

ravennagrass

Europe

I

Eriochloa polystachya procera punctata

caribgrass cupgrass Lousiana cupgrass

West Indies tropical Asia American tropics

1916 1936 1916

Euchlaena mexicana perennis

teosinte Mexican teosinte

Latin America Latin America

1909 1940

939

COMMON

NAMES,

ORIGINS,

TABLE 3

AND FIRST

RECORDS

(continued) PUTATIVE

BOTANICAL NAME

Festuca arundinacea elatior hawaiiensis idahoensis kashmiriana ovina pratensis rubra tenui/olia viridula

323

COMMON NAME

reed, fescue, alta fescue tall fescue Hawaiian fescue Idaho fescue

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Europe, temperate Asia Europe Hawaiian Islands western United States India

sheep fescue red fescue greenleaf fescue

Garnotia sandwicensis

Eurasia Europe Europe northwestern United States

1931 1911

1938 ? 1910 ? 1912 1913 ?

Hawaiian Islands

Gastridium phleoides ventricosum

nitgrass

Chile Europe

? 1906

Glyceria fluitans grandis

mannagrass American mannagrass

Eurasia North America

1906 1922

Hackelochloa granulans Heteropogon contortus Hilaria belangeri

cenchroides jamesii mutica

cosmopolitan tropics piligrass, twisted beardgrass, tanglehead curly mesquite

galleta tobosagrass

India, Polynesia southwestern United States and northern Mexico Mexico North America western United States

J

939 ?

l

939

1936

GRASSES OF

324

TABLE 3

HAWAII

(continued.) PUTATIVE

BOTANICAL NAME

COMMON NAME

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Holcus lanatus

velvetgrass, Yorkshire fog

Europe

1906

Hordeum brachyantherum bulbosum murinum nodosum vulgare

meadow barley bulbous barley wild barley meadow barley barley

North America Eurasia Europe Old World Old World

!939 1906 r 93S 1906

Hyparrhenia hirta rufa

thatchinggrass, jaragua

Africa, Middle East Africa

Imperata cylindrica Isachne distichophylla myosotis pallens Ischaemum byrone digitatum indicum

ohe

1916 *933

Europe

?

Hawaiian Islands Philippine Islands Hawaiian Islands

?

Baron's grass

Hawaiian Islands New Guinea India, Southeast Asia

1955 ?

Ixophorus unisetus

mexicangrass

Mexico

1921

Lamarckia aurea

goldentop

Mediterranean region

1937

Leptochloa virgata

juddgrass

tropical America

1906

Lepturus cinereus repens Lolium multiflorum perenne rigidum subulatum temulentum

Russell Islands Polynesia to Australia Italian ryegrass perennial ryegrass Wimmera ryegrass darnel

Europe Europe Europe Europe Europe

1906 1911 1936 I93I

1906

COMMON

NAMES,

ORIGINS,

TABLE 3

AND FIRST

RECORDS

(continued) PUTATIVE

BOTANICAL NAME

Lycurus phleoides

COMMON NAME

Microlaena stipoides Muhlenbergia microsperma

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

1940

western United States

1936

molassesgrass

Africa

!9!3

puu lehua, meadow ricegrass

Australia, Polynesia

littleseed muhly

porteri

bush muhly

racemosa repetís

marsh muhly creeping muhly

Narenga porphyrocoma Oplismenus burmannii compositus hirtellus

DATE OF

North America

wolftail

Melica californica Melinis minutiflora

325

southwestern United States to Peru southwestern United States, northern Mexico North America southwestern United States, Mexico

India

1

935

1936 1922

1

937

1930 p

India, Burma Ceylon

p

basketgrass, honohonokukui

tropical America

1841

Oryza sativa

rice, laiki

Asia

1856

Oryzopsis coerulescens hymenoides

Indian ricegrass

miliacea

smilograss

North Africa western United States Mediterranean region

? 1936 1918

326

GRASSES OF

TABLE 3

BOTANICAL NAME

Panicum alakaiense antidotale beecheyi bulbosum

capillare carteri colliei coloratum conjugens cynodon degeneri dichotomiflorum fauriei forbesii heupueo hillebrandianum var. gracilius imbricatum f. minus var. molokaiense isachnoides var. kilohanae kaalaensis konaense koolauense laevifolium malacophyllum maximum var. trichoglume miliaceum molokaiense nephelophilum niihauense nubigenum

HAWAII

(continued)

COMMON NAME

giant panicgrass Beechey panicgrass bulb panicum

witchgrass

blue panicgrass

fall panicum Faurie's panicgrass Forbes' panicgrass Hillebrand's panicgrass

Kaala panicgrass, konakona Kona panicgrass Koolau panicgrass

guinea grass green panicgrass broomcorn millet Molokai panicgrass konakona Niihau panicgrass

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Hawaiian Islands India Hawaiian Islands southwestern United States, Mexico United States Hawaiian Islands Hawaiian Islands tropical Africa Hawaiian Islands Hawaiian Islands Hawaiian Islands eastern United States Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands South Africa southern United States Africa Africa Eurasia Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands

1916

?

1922 1942 1938

I9SO

1916 ?

1880 p 1923

COMMON N A M E S , O R I G I N S , AND F I R S T

TABLE 3

BOTANICAL NAME

obtusum

virgatum xerophilum

COMMON NAME

vine mesquite

torrid panicgrass, kakonakona switchgrass kakonakona, konakona

Paspalidium jubiflorum Paspalum almum bertonii commersonii conjugatum dilatatum distichum elegans fimbriatum floridanum hartwegianum

Comb's grass

327

(continued)

oreoboloides var. subimbricatum pellitoides pellitum pilosum prolutum coolah grass radiatius repens quackgrass, torpedograss scribnerianum tenuifolium mountain pili, konakona texanum Texas millet torridum

RECORDS

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

southwestern United States, Mexico Hawaiian Islands Hawaiian Islands Hawaiian Islands Hawaiian Islands West Indies Australia Hawaiian Islands Old World North America Hawaiian Islands southwestern United States

1936

? 1936 1906 1939 !939

Hawaiian Islands United States Hawaiian Islands

I93S

Australia

1939

South America South America Old World tropics

1939 1916 1938

Hilo grass, mauu-Hilo, mauu malahini, sour paspalum American tropics dallisgrass, paspalum grass, Australian watergrass South America knotgrass West Indies

1906 1906 ?

Panama paspalum

1906

tropical America southern United States southwestern United States and Mexico

1840

1939 1939

328

GRASSES OF

TABLE

BOTANICAL NAME

3

HAWAII

(continued)

COMMON NAME

lividum

longtom

malacophyllum notatum orbiculare

ribbed paspalum Bahia grass ricegrass, mauu-laiki

plicatulum

browntop millet, brownseed millet

racemosum stramineum urvillei vaginatum

vaseygrass biscuitgrass, seashore paspalum

virgatum Pennisetum clandestinum complanatum macrostachyum orientale purpureum schimperi setaceum setosum typhoides villosum Phalaris arundinacea californica caroliniana coerulescens commutata minor paradoxa tuberosa

kikuyugrass

elephantgrass, napiergrass fountaingrass feathery pennisetum pearl millet feathertop reed canarygrass California canarygrass Carolina canarygrass

large canarygrass

PUTATIVE

D A T E OF

O R I G I N OR

FIRST

NATIVE COUNTRY

RECORD

warmer areas of both Americas Latin America tropical America probably southern China (by early Hawaiians?) southern United States and central South America Peru southwestern United States South America

1939 1906

tropical America Central America

!937 1914

Africa tropical America East Indies India, North Africa

1924 1921 1906 1938

Africa Africa North Africa tropical America India, tropical Africa Africa

1912 1918 1926 1921 ?

Northern Hemisphere Pacific coast of United States United States Mediterranean region Europe, Italy Mediterranean region Mediterranean region Mediterranean region

1922

I

939

!939 !9!3

1935 Ï9Ï3

1938

1906 r

939 ?

1912 iyuu 1906 1906 1915

C O M M O N N A M E S , O R I G I N S , AND F I R S T

TABLE 3

BOTANICAL NAME

RECORDS

329

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

Phleum pratense subulatum Poa ampia annua arachnifera bulbosa compressa flabellata mannii nemoralis nervosa pratensis sandvicensis scabrella siphonoglossa trivialis

timothy

Eurasia 1912 Mediterranean region 1913

big bluegrass annual bluegrass Texas bluegrass bulbous bluegrass Canada bluegrass

North America Europe Europe Europe Europe South America Hawaiian Islands Europe western United States Europe Hawaiian Islands western United States Hawaiian Islands

Mann's bluegrass wood bluegrass Wheeler bluegrass Kentucky bluegrass Hawaiian bluegrass pine bluegrass rough bluegrass, roughstalk bluegrass

Polypogon interruptus ditch polypogon monspeliensis rabbitfoot grass Polytoca macrophylla Polytrias amaura Rhynchelytrum repens Natal redtop Saccharum arundinaceum barberi bengalense edule (=officianarum) officinarum sugar cane, ko robustum wild cane sinense spontaneum

?

1864 I9I2

937 1906 1916 x

I

9I3

*935 1906 1936

Europe

r

Venezuela Venezuela

1871 1906

South Pacific Islands

1940

Southeast Asia

1916

Africa

1895

New Caledonia India Bengal

1948 1930 1920

New Guinea early Hawaiians New Guinea China, Japan Old World

1964

9r3

1929 I 9°S 1918?

33°

GRASSES OF

TABLE 3

HAWAII

(continued) PUTATIVE

BOTANICAL NAME

COMMON NAME

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

Sacciolepis indica

glenwoodgrass

East Indies

1906

Schedonnardus paniculatus

tumblegrass

Western Hemisphere

1916

Schizachyrium scoparium

prairie beardgrass

United States

1922

N e w Caledonia

1948

Old World

1906

Australia

1936

Sclerostachya fusca Secale cereale

rye, huapalaoa-eleele

Sehitna nervosum Setaria barbata geniculata

glauca

italica longiseta palmi/olia sphacelata verticillata

? tropical Asia yellow foxtail, knotroot, bristlegrass, mauukaleponi

tropical America native to warm temperature zone of Old World foxtail millet, Italian millet Eurasia Africa palmgrass tropics of Old World tropical and South Africa bristly foxtail, Old World mauu-pilipili

1851

1910 x

959

1906 1958 i860

Sorghastrum nutans

indiangrass

North America

!939

Sorghum almum halepense sudanense vulgare

columbusgrass johnsongrass sudangrass sorghum

Argentina Mediterranean region Africa Africa

1906 1937 i860

cordgrass, Townsend cordgrass

Europe

1932

Spartina x townsendii

COMMON N A M E S , O R I G I N S , AND F I R S T RECORDS

TABLE 3

(continued) PUTATIVE

BOTANICAL NAME

Sphenopholis obtusata

COMMON NAME

prairie wedgegrass

Spodiopogon aureus Sporobolus africanus airoides cryptandrus diander elongatus fimbriates flexuosus indicus virginicus wrightii Stenotaphrum secundatum

Stipa cornata leucotricha papposa viridula

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

North America

1935

China rattail grass, smutgrass alkali sacaton sand dropseed Indian dropseed mesa dropseed

South Africa western United States North America India Australia South and East Africa western United States tropical America

? 1906 1936 ? 1906 1925 1939 1936 1912

beach dropseed, seashore rushgrass, manienie-maoli akiaki, manienie-akiaki tropical America 1864 saccaton western United States 1936 buffalograss, St. Augustine grass, manienie-mahiki, akiaki-haole, southeastern manienie-akiaki (-haole) United States needle-and-thread grass Texas needlegrass green needlegrass

Themeda arguens australis gigantea quadrivalvis triandra

331

kangaroograss

western United States 1922 southwestern United States and Mexico 1936 Uruguay, Argentina north central United States and adjacent 1922 Canada Southeast Asia, tropics in general Australia Philippines warm and tropical areas of Old World

1939 1936 1936 x 939 J

939

332

GRASSES OF H A W A I I

TABLE 3

BOTANICAL NAME

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY RECORD

Tragus berteronianus

burgrass

Old World

1935

Trichachne californica

cottontop

insularis

sourgrass

North and South America tropical America

*939

Trichoneura elegans

southwestern United States

Tridens flavus

purpletop

Tripsacum dactyloides laxum

eastern gamagrass Guatamala grass

Triraphis mollis Trisetum flavescens glomeratum inaequale Triticum aestivum

?

!939 Central America Central America

1940 1919

Australia yellow trisetum, yellow oatgrass heu-pueo (name used on island of Hawaii), pili-uka

wheat, huapa-laoa

Urochloa bolbodes panicoides pullulans Vulpia megalura myuros octaflora

1906

foxtail fescue

Europe

?

!9J3

Hawaiian Islands Hawaiian Islands Asia

1792

tropical Africa India, Africa Africa

r

North America Europe United States

?

945

1942

I860

1938 1936

COMMON N A M E S , O R I G I N S ,

TABLE 3

BOTANICAL NAME

Zea mays

Zizania latifolia Zoysia japonica matrella tenuifolia

AND F I R S T RECORDS

333

(continued)

COMMON NAME

PUTATIVE

DATE OF

ORIGIN OR

FIRST

NATIVE COUNTRY

RECORD

maize, corn, kulina (Indian-corn), kulina-ono (sweetcorn), kulinapohapoha (popcorn) America

1792

wildrice

Asia

1936

Japan Asian tropics

1961 1916

Mascarene Islands

1906

manilagrass Japanese lawngrass, Korean velvetgrass, velvetgrass, mauukepani

GLOSSARY ABAXIAL. Turned away from the axis, in contrast to adaxial. A C U M I N A T E . Gradually tapering to a sharp point. ACUTE. Sharp-pointed, but less tapering than acuminate. ADAXIAL. Facing or against the axis, in contrast to abaxial. A G G R E G A T E . Collected together in tufts, groups, or bunches. A L T E R N A T E . Placed at different levels, not opposite. A N N U A L . Within one year; applied to grasses which live less than one year. A N T H E R . Sac-like portion of the stamen containing the pollen. A N T H E S I S . The period during which a flower is open. ANTRORSE. Directed upward or forward. APICULATE. Having a minute pointed tip. APPRESSED. Lying against an organ. ARACHNOID. Composed of slender entangled hairs; cobwebby. ARISTATE. Awned; provided with a bristle at the end or at the back or edge of an organ. A R T I C U L A T E . Jointed; joined by a line of demarcation between the two parts which at maturity separate by a clean-cut scar. A S C E N D I N G . Sloping upward. A T T E N U A T E . Gradually narrowed to a slender apex or base. AURICLE. An ear; applied to earlike lobes at the base of blades and to the small lobes at the summit of the sheath. A U R I C U L A T E . Provided with ears. AWN. A slender bristle at the end or on the back or edge of an organ. AXIL. T h e angle between an organ and its axis. AXILLARY. In, or arising from, an axil. BEAK. A hard prolonged tip, usually applied to the ends of seeds and fruits. BEARDED. Hairy, the hairs not scattered. BI-. A Latin prefix meaning two. BIENNIAL. A plant completing its life cycle within two years. BIFID. Two-cleft or two-lobed. BLADE. The part of the leaf above the sheath. BRACT. A reduced leaf forming part of the inflorescence; examples are glumes, lemmas, and paleas, in grasses. BRISTLE. A stiff slender appendage, like the bristles of a brush. CADUCOUS. Falling off very early. CALLUS. T h e basal extension of the mature lemma appearing as a hard, often hairy, protuberance. CANESCENT. Gray-pubescent or hairy. CAPILLARY. Very slender or hairlike. CARINATE. Having a keel. C A R T I L A G I N O U S . Firm and tough; with cartilage-like texture.

336

GRASSES OF H A W A I I

C A R Y O P S I S . The grain or fruit of grasses having the pericarp united with the wall of the ovary; in a few grasses the pericarp is free from the seed. C A U D A T E . Tailed or bearing a tail-like appendage. C E N T I M E T E R (cm), i/ioo of a meter; 0.39 inches. C E S P I T O S E . Having low stems growing in close clusters, forming tufts. C H A R T A C E O U S . Papery or parchment-like in texture. C H L O R E N C H Y M A . Parenchyma tissue containing chloroplasts. C I L I A T E . Marginally fringed with hairs. C I L I O L A T E . Minutely ciliate. C I R C U M S C I S S L E . Dehiscing by a transverse fissure around the circumference. C L E I S T O G A M O U S . Having flowers fertilized by their own pollen within the closed bud. C O L L A R . In grasses, the zone at the junction of the sheath and blade on the outer (under, abaxial) surface of the leaf. C O L U M N . In grasses, the usually twisted lower part of awns that are bent about halfway along their length. C O M P R E S S E D . Flattened. C O N N A T E . United or joined, particularly when two or more like parts are united. C O N V O L U T E . Rolled with one margin wrapped around the other. C O R D A T E . Heart shaped; having a rounded lobe on either side at the base. C O R I A C E O U S . Leathery in texture. C U L M . The main stem in grasses, including the inflorescence. C U N E A T E . Wedge-shaped, the narrow end below. C Y M B I F O R M . Boat-shaped. D E C I D U O U S . Falling off at a particular stage of growth, in contrast to persistent. D E C U M B E N T . Erect or curving upward from a horizontal base; in grasses, applied to the culm. D E H I S C E . T o open. D E N T A T E . Toothed, usually with the teeth pointed outward. D E P A U P E R A T E . Small, usually implying impoverishment; below normal size or dwarfed. D I C H O T O M O U S . Divided in pairs, e.g., dichotomously branched. D I G I T A T E . Arranged like fingers; in grasses, applied to inflorescences comprising three or more racemes at the apex of the culm. D I O E C I O U S . Unisexual, the male and female on different plants. D I S A R T I C U L A T I N G . Separating at the joints or nodes at maturity. D I S T I C H O U S . Arranged conspicuously in two ranks. D I V A R I C A T E . Spreading widely. D O R S A L . Relating to the back. E F F U S E . Spreading.

GLOSSARY

337

E L L I P S O I D . An elliptic solid. E L L I P T I C . Shaped like an ellipse; widest at about the middle, gradually narrowed to both ends, and approximately twice as wide as long. E L O N G A T E . Long and narrow. EMARGINATE. Shallowly notched at the apex. EMBRYO. The rudimentary plant formed in the seed. E N D E M I C . Growing only in a particular geographic location. EROSE. Irregularly notched at the apex or on the margin, as if gnawed. EXSERTED. Projecting beyond, protruding. FASCICLE. A small dense cluster or bundle. F E R T I L E . Producing viable seed and having functional stamens, in contrast to sterile. FIBRILLOSE. Composed of fibers or fiber-like organs. F I L A M E N T . The stalk supporting the anther, and the two making up the stamen. F I L I F O R M . Threadlike. FIMBRIATE. Fringed along the margins. FLABELLATE. Fan-shaped or wedge-shaped. FLEXUOUS. Bending alternately in opposite directions; zigzag. F L O R E T . In grasses, the lemma and palea with enclosed flower (stamens and pistil); florets may be bisexual (perfect, hermaphrodite), unisexual (either male or female), sterile (neuter, barren, empty), or reduced to the lemma. F U S I F O R M . Thickest near the middle and tapering toward each end; spindle-shaped. G E N I C U L A T E . Kneed; bent abruptly like a knee; in grasses, applied to culms bent at the nodes, or to bent awns. GLABROUS. Without distinct hairs, but not necessarily smooth. GLAND. A protuberance or depression that secretes, or appears to secrete, a fluid. GLAUCOUS. Blue-green, often owing to a white wax or white skin covering the green parts of the plant. GLOBOSE. Nearly spherical. G L U M E . An empty bract at the base of the spikelet. GRAIN. The caryopsis or naked seed in grasses. HERBACEOUS. Thin in texture and green in color, in contrast to woody. H E T E R O G A M O U S . Bearing two types of flowers, such as male and female. H E T E R O M O R P H O U S . Variation from the normal structure, or having organs of different forms. H I L U M . The point of attachment of the ovule to the ovary wall in the grain. H I R S U T E . Covered with stiff rigid hairs. H I R T E L L O U S . Minutely hirsute. HISPID. Covered with stiff rigid hairs.

33»

GRASSES

OF

HAWAII

H I S P I D U L O U S . Minutely hispid. H O M O G A M O U S . Having one kind of flower (floret). H Y A L I N E . T h i n and transparent, or nearly so. I M B R I C A T E . Overlapping, like shingles on a roof. I N C L U D E D . Enclosed by and not protruding beyond surrounding organs. I N D U R A T E . Hardened or becoming tough. I N F L E X E D . Turned abruptly inward at the margins. I N F L O R E S C E N C E . Flower head; in grasses, a panicle, raceme, or spike, or reduced to one spikelet. I N T E R N O D E . T h e portion between two successive nodes on an axis. I N V O L U C R E . A ring or circle of bracts below the flower or flower cluster; in grasses, applied to the cluster of bristles or sterile branchlets below the spikelets in Pennisetum and a few other genera. I N V O L U T E . Having the margins rolled inward. J O I N T . An articulation. K E E L . A sharp fold or ridge; in grasses, applied usually to a compressed leaf, glume, lemma, or palea. L A C E R A T E . T o r n or cleft irregularly, like the ligule in some grasses. L A N C E O L A T E . Broadest below the middle and tapering toward both ends; shaped like a lance head. L A T E R A L . Belonging to the side. L E M M A . A bract of the spikelet above the glumes; the outer bract subtending the flower. L I G U L E . A n outgrowth (either a membrane or a fringe of hairs) on the inner (upper, adaxial) surface at the junction between the sheath and blade in the leaves of grasses. L I N E A R . Long and narrow with parallel sides, usually at least ten times as long as wide, like the leaf blades in many grasses. L O B E . A segment or division; in grasses, applied especially to the segments of a lemma that is divided at the apex. L O B E D . Divided into apical segments. L O D I C U L E . A small scale at the base of the stamens and the pistil in most grasses. M E M B R A N O U S ( M E M B R A N A C E O U S ) . Like skin in texture; thin, dry, and often translucent. M E T E R (m). T h e standard unit of linear measurement of the metric system; about 39.37 inches. M I D R I B . T h e midvein or main rib of a leaf. M O N O E C I O U S . Unisexual, the two kinds of flowers on the same plant. M U C R O . A sharp point, minute awn. M U C R O N A T E . Having a mucro. M U T I C O U S . Blunt, lacking a point. N E R V E . In grasses, a vein in the leaf, glume, lemma, or palea.

GLOSSARY

339

N E U T E R . Without stamens or pistils; said of florets or spikelets. N O D E . The point on a stem or other axis at which a leaf or bract arises. OB-. A Latin prefix meaning reversed. O B S O L E T E . Not evident, or verging upon complete suppression. O B T U S E . Blunt or rounded at the apex. O V A R Y . The female part of the flower enclosing the ovule, which develops into the seed after fertilization. O V A T E . Egg-shaped, with the broader end downward. O V O I D . An egg-shaped solid. P A L E A . The upper of the two bracts enclosing the flower in grasses, sometimes called the valvule or pale. P A N I C L E . A branched inflorescence; a compound flower cluster with a main axis and subdivided branches. P A P I L L O S E . With small nipple-shaped projections. P E C T I N A T E . Comblike; used especially with some species of Bouteloua where the spikelets are set close together, parallel and divergent from the rachis like the teeth of a comb. P E D I C E L . In grasses, the stalk of the spikelet. P E D I C E L L A T E . Borne on a pedicel. P E D U N C L E . In grasses, applied to the stalk of a raceme or cluster of spikelets. P E N D U L O U S . Somewhat drooping. P E N I C I L L A T E . Tipped with a tuft of fine hairs, as the stigmas of some grasses. P E R E N N I A L . Of more than two years duration; used either as a noun or an adjective. P E R F E C T . A bisexual flower; having both pistil and stamen. P E R S I S T E N T . Remaining attached to the axis, in contrast to deciduous. PERICARP. The skin of the grass fruit; ripened wall of the ovary. P I L O S E . Bearing long soft hairs. P I S T I L . The female part of the flower, consisting, when perfect, of an ovary, style, and stigma. P L A N O - C O N V E X . Flat on one side and curved on the other. P L I C A T E . Folded into longitudinal plaits. P L U M O S E . Feathery with soft hairs; resembling a plume or feather. P R I C K L E . More or less slender, short, sharp, hard outgrowth of the epidermis. P R O C U M B E N T . Lying on the ground without rooting at the nodes and with ascending tips. P U B E R U L E N T . Covered with short, fine, or almost imperceptible hair. P U B E S C E N T . Hairy; usually a general term for hairiness. P U L V I N U S . A cushion-like swelling at the base of branches which causes them to spread.

340

GRASSES OF

HAWAII

P U N C T A T E . Dotted with minute depressions. P U N C T I F O R M . In the form of a dot or point. P U N G E N T . Piercing; having a sharp rigid point, like the leaf blades of some grasses. RACEME. An unbranched inflorescence or part of an inflorescence, with the spikelets pedicelled directly on the axis. RACHILLA (or RHACHILLA). The axis of the spikelet in grasses. RACHIS (or RHACHIS). In grasses, the main axis of a raceme or spike. RADIATE. Spreading from or arranged around a common center. RETRORSE. Pointing backward or downward. RHIZOME. An underground stem, bearing scalelike leaves. RHOMBOIDAL. More or less quadrangular with lateral obtuse angles. R U D I M E N T . An imperfectly developed organ or part. RUGOSE. Wrinkled, ridged, like the upper floret in some genera of the Paniceae. SACCATE. Bearing distended, sac-like or pouch-like bodies. S A G I T T A T E . Lobed at the base, like the barbed head of an arrow. SCABERULOUS (SCABRIDULOUS). Slightly scabrid. SCABRID. Rough to the touch, covered with minute spines. SCABROUS. The same as scabrid. SCALE. A miniature leaf without blades; in grasses, sometimes applied also to a reduced glume, lemma, or palea. SCARIOUS. Thin and dry, appearing as if shrivelled. SCURFY. Appearing as though covered with minute scales. SECUND. One-sided, or arranged along one side. SESSILE. Without a stalk. SETACEOUS. Bristle-like. SHEATH. The part of the leaf below the blade, usually surrounding the culm. S I N U A T E . Having strongly wavy margins. SINUS. T h e angle or recess formed by two lobes. SPATHE. A bladeless leaf or sheathing bract below the inflorescence or racemes in some grasses, especially genera of the Andropogoneae. SPIKE. An unbranched inflorescence or part of an inflorescence, with the spikelets sessile and borne directly on the axis. S P I K E L E T . The unit of the grass inflorescence, usually composed of two glumes and one or more flowers each borne in the axil of a lemma and a palea. S T A M E N . The male (pollen-bearing) part of the flower, consisting of an anther and usually a filament. S T A M I N O D E . A sterile reduced stamen. STEM. T h e main axis of the plant, bearing the leaves and inflorescence; in grasses, called the culm.

GLOSSARY

341

S T E R I L E . Not producing viable seed or pollen, in contrast to fertile. S T I G M A . T h e female part of the flower that receives the pollen. S T I P E . A minute stalk-like support of an organ. S T O L O N . A stem that creeps along the surface of the ground (in contrast to a rhizome), rooting from the nodes and there giving rise to vegetative shoots and culms. S T R I A T E . Marked with parallel lines or ridges. S T Y L E . T h e connection between the stigma and the ovary. S U B - . A Latin prefix used to denote almost, somewhat, or slightly; for example, glumes that are almost but not quite equal in length are said to be subequal, and if they are somewhat blunt, they are said to be subobtuse. S U B C U C U L L A T E . Somewhat hood-shaped. S U B T E N D . T o be below, like the bristles below the spikelets in Setaria and Pennisetum. S U B U L A T E . Awl-shaped. T E R E T E . Circular in section; rounded, not angled or grooved. T O M E N T O S E . Densely covered with cottony or woolly hairs. T O M E N T U L O S E . Minutely tomentose. T R U N C A T E . Ending abruptly in a nearly square apex, as if cut off at the end. T U B E R C U L A T E . Bearing small pimple-like projections. T U F T E D . Growing in close clusters. T U S S O C K . A very dense bunch or tuft. U N C I N A T E . Bent or hooked inward at the apex. U N I S E X U A L . Of one sex. V E R T I C E L L A T E . With organs arranged in whorls, as several leaves or branches at a node. V I L L O U S . Covered with long soft hairs.

BIBLIOGRAPHY Bews, J. W. 1929. The World's Grasses. Longmans, Green & Co., London. 408 pp., illus. Bor, N. L. i960. The Grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, London. 767 pp., illus. Brown, W. V. 1958. Leaf anatomy in grass systematics. Bot. Gaz. 119: 170-178. Bryan, E. H., Jr. 1931. Checklist of the plants of the Hawaiian Islands. Revised by H. St. John, 1931. Unpublished manuscript. Chase, A., and C. D. Niles. 1962. Index to Grass Species. G. K. Hall & Co., Boston. 3 vols. Cheeseman, T. F. 1925. Manual of the New Zealand Flora. New Zealand Board of Science and Arts. Wellington. 1161 pp., illus. Degener, O. 1946. Flora Hawaiiensis or New Illustrated Flora of the Hawaiian Islands. 2nd ed., Books 1-4. 1192 pp., illus. and I. Degener. 1963. Flora Hawaiiensis or New illustrated Flora of the Hawaiian Islands. Book 6. 200 pp., illus. Gardner, C. A. 1952. Flora of Western Australia. Vol. 1, part 1, Gramineae. William H. Wyatt, Govt. Printer, Perth, Western Australia. 400 pp., illus. Henrard, J. Th. 1950. Monograph of the Genus Digitaria. Universitaire Pers Leiden, Leiden. 999 pp., illus. Hillebrand, W. F. 1888. Flora of the Hawaiian Islands. B. Westermann & Co., New York. 674 pp. Hitchcock, A. S. 1922. The grasses of Hawaii. Mem. Bernice P. Bishop Museum 8:101-230, illus. 1936. Manual of the Grasses of the West Indies. U. S. Dept. Agr. Misc. Pubi. 243, 439 pp., illus. 1950. Manual of the Grasses of the United States. 2nd ed., revised by A. Chase. U. S. Dept. Agr. Misc. Pubi. 200, 1051 pp., illus. Hubbard, C. E. 1934. Gramineae. In J. Hutchinson, Families of Flowering Plants. Vol. 2, Monocotyledons. Macmillan Co., London. 1948. Gramineae. In J. Hutchinson, British Flowering Plants, Evolution and Classification. P. R. Gawthorn Ltd., London. 1959. Gramineae. In J. Hutchinson, Families of Flowering Plants. 2nd ed. Clarendon Press, Oxford. 2 vols.

344

GRASSES OF H A W A I I

Mason, H. L. 1957. A Flora of the Marshes of California. University of California Press, Berkeley. 848 pp., illus. McClelland, C. K. 1915. Grasses and Forage Plants of Hawaii. Hawaii Agr. Exp. Sta. Bull. 36, 42 pp., illus. Meredith, D., ed. 1955. The Grasses and Pastures of South Africa. Cape Times Ltd., C.P. Parrow, Union of South Africa. 771 pp., illus. Phillips, E. P. 1951. The Genera of South African Flowering Plants. 2nd ed. Union of South Africa Dept. of Agr. Div. Bot. PI. Path. Bot. Survey Memoir no. 25. Praetoria, South Africa. 923 pp. Reeder, J. R. 1957. The embryo in grass systematics. American Journ. Bot. 44:756-768. Ripperton, J. C., R. A. Goff, D. W. Edwards, and W. C. Davis. 1933. Range Grasses of Hawaii. Hawaii Agr. Exp. Sta. Bull. 65, 38 pp., illus. Silveus, W. A. 1933. Texas Grasses. Classification and Description of Grasses, Descriptive Systematic Agrostology. The Clegg Co., San Antonio, Texas. 782 pp., illus. Snowdon, G. D. 1936. The Cultivated Races of Sorghum. Adlard & Son, Ltd., London. 273 pp. 1955. The wild fodder sorghums of the section Eu-Sorghum. Journ. Linn. Soc. (Bot.) 55:191-260. Stebbins, G. L., and B. Crampton. 1959. A suggested revision of the grass genera of temperate North America. In Recent Advances in Botany Whitney, L. D., E. Y. Hosaka, and J. C. Ripperton. 1939. Grasses of the Hawaiian Ranges. Hawaii Agr. Exp. Sta. Bull. 82, 148 pp., illus.

INDEX Agropogon Fourn., 51, 53, 313 littoralis (Smith) C. E. Hubb., 53, 7 i . 313 Agropyron Gaertn., 124, 125, 313 cristatum (L.) Gaertn., 125, 313 dasystachyum (Hook.) Scribn., 125, 313 desertorum (Fisch.) Schult., 125, 313 elongatum (Host) Beauv., 125, 313 inerme (Scribn. et Smith) Rydb., «5,31.3 intermedium (Host) Beauv., 126, 3'3 pauciflorum (Schwein.) Hitchc., 126 repens (L.) Beauv., 126, 127, 313 smithii Rydb., 126, 313 spicatum (Pursch.) Scribn. et Smith, « 6 , 313 tenerum Vasey, 126 trachycaulum (Link) Malte 126, 313 trichophorum (Link) Rieht., 126, 313 Agrostideae Dumort., 28, 51 Agrostis L., 51, 52, 53, 313 africana Poir., 179 alba L., 54, 55, 313 alba auctt. non L., 54 avenacea Gmel., 54, 313 borealis Hartm., 54, 313 canina L., 54, 313 capensis (L.) Lamk., 179 capillaris L., 54 diandra Retz., 179 exarata Trin., 54, 56 exarata microphylla (Steud.) Hitchc., 54 fallax Hillebr., 54, 57, 3 I 3 fertilis Steud., 179 interrupta L., 62 kauaiense Hillebr., 54 littoralis With., 71 matrella L., 186 microphylla Steud., 54, 313 microsperma Lag., 68 palustris Huds., 54, 313 spicata Vahl, 179 spicatus Thunb., 179 radiata L., 142 retrofracta Willd., 54 rockii Hack., 54 rubra L., 54 sandwicensis Hillebr., 54, 58, 313 scabra Willd., 54, 313

semiverticillata (Forsk.) C. Christ., 54. 59, 313 stolonifera L., 53, 54, 313 tenuis Sibth., 54, 313 ventricosa Gouan, 66 verticillata Vili., 54 vulgaris With., 54 Aira L., 73, 74, 314 australis Nees, 82 caryophyllea L., 75, 76, 314 flexuosa L., 84 hawaiiensis (Hillebr.) Skottsb., 82 hawaiiensis f. depauperata Skottsb., 82 hawaiiensis f. haleakalensis Skottsb., 83 indica L., 302 nubigena (Hillebr.) Hitchc., 82, 83 obtusata Michx., 87 pallida (Hillebr.) Skottsb., 82 pallida var. tenuissima Skottsb., 82 praecox L., 75 pumila Pursh., 105 spicata L., 302 Alloteropsis J. S. Presi ex C. B. Presi, 231, 234, 235, 314 cimicina (L.) Stapf, 235, 314 Alopecurus L., 51, 52, 60, 314 monspelions (Sphalm) L., 71 monspeliensis L., 71 pratensis L., 60, 61, 314 Ammophila Host, 51, 52, 60, 314 arenaria (L.) Link, 60, 314 Andropogoneae Dumort., 24, 188 Andropogon L., 188, 192, 193, 314 aciculatus Retz., 199 annulatus Forsk., 202 argenteus DC., 197 barbatum L., 142 barbatus L., 138 barbinodis Lag., 197 barbinedis var. perforatus (Trin. ex Fourn.) Gould., 197 bicornis L., 193, 314 caricosus subsp. mollicomus (Hack.) DC. var. mollicomus (Kunth) Hack., 202 caucasicus Trin., 197 citratus DC., 201 contortus L., 206 furcatus Muhl., 193 gerardi Vit., 193, 194, 314 glomeratus (Walt.) B. S. P. Presi, 195, 314

INDEX

346

halepensis Brot., 218 halepensis ( L . ) Brot., var. typicus Aschers, et G r a e b n . , 218 halepensis (L.) Brot. var. genuinus (Hack.) Stapf, 218 hallii Hack., 195, 314 insularis L . , 308 intermedins R. Br., 197 intermedins var. caucasico ( T r i n . ) Hack., 197 ischaemum L . , 197 ischaemum Roxb. ex W i g h t , 202 littoralis Nash, 195, 314 nodosus (Willem.) N a s h , 202 perforatus T r i n . , 197 pertusus (L.) Willd., 197 polydactylon L . , 142 rufus (Nees) K u n t h , 208 saccharoides S w . , 197 scoparius M i c h x . , 214 sericeus R. Br., 202 sorghum (L). Brot, s u b s p . halepensis Hack., 218 sorghum sudanensis Piper, 218 subulatus Presi, 199 virginicus L., 195, 196, 314 Anthistiria L i n n , f., 220 A n t h o x a n t h u m L., 73, 75, 314 o d o r a t u m L., 77, 78, 314 A p e r a Adans., 51, 52, 60, 314 i n t e r r u p t a (L.) Beauv., 62, 314 Aristida L., 38, 314 adscensionis L., 39, 40, 314 divaricata J a c q . , 39 Aristideae C. E. Hubb., 29, 38 A r r h e n a t h e r u m Beauv., 73, 77, 314 aveaceum (Scop.) Beauv., 77 elatius (L.) J. S. et C. B. Presi, 77, 79i 314 A r u n d i n e a e Dumort., 27, 4 1 Arundinella Raddi, 44, 314 agrostoides T r i n . , 45 holcoides (Kunth) Trin., 45, 314 A r u n d i n e l l e a e Stapf, 28, 44 A r u n d o L., 4 1 , 314 donax L., 42, 43, 314 d o n a x var. versicolor ( Miller ) Stokes, 42 Aspris Adans., 74, 75 carophyllea (L.) N a s h , 75 Astrebla F. Muell., 46, 314 elymoides Bailey et Muell., 47, 314 lappacea (Lindl.) Domin., 47, 314 p e d i n a t a (Lindl.) F. Muell., 47, 3*4

Avena L., 73, 74, 77, 314 b a r b a t a Pott ex Link, 80, 314 byzantina C. Koch, 80, 314 elatior L . , 77 fatua L., 80, 81, 315 flavescens L . , 87 sativa L., 8o, 315 Aveneae Dumort., 26, 73 A x o n o p u s Beauv., 231, 234, 2 3 5 , 315 affinis Chase, 236, 315 Beauv.,236,237, c o m p r e s s u s (Sw.) 315 B a m b u s e a e Aschers et Graebn., 25 B e c k m a n n i a Host, 95, 96, 315 erucae/ormis var. uniflora Scribn., 97 syzigachne (Steud.) Fern., 97, 98, 315 Bothriochloa O. Ktze., 188, 191, 195, 31S a m b i g u a S. T. Blake, 195, 3 1 5 b a r b i n o d i s (Lag.) Herter, 197, 198, 315 caucasica (Trin.) C. E. Hubb., 1 9 7 , 31S insculpta (Höchst.) A. Camus, 1 9 7 , 31S i n t e r m e d i a (R. Br.) A. Camus, 1 9 7 , 315 i s c h a e m u m (L.) Keng, 197, 315 p e r f o r a t a (Trin. ex Fourn.) Herter, 197. 31S pertusa (L.) A. Camus, 1 9 7 , 315 saccharoides (Sw.) Rydb., 197, 315 Bouteloua Lagasca, 134, 1 3 5 , 315 chondriosioides (H. B. K.) Benth. ex Wats., 1 3 5 , 315 c u r t i p e n d u i a (Michx.) Torr., 1 3 5 , .136, 315 eriopoda (Torr.) Torr., 1 3 7 , 315 filiformis (Fourn.) Griffiths, 137, 315 gracilis (H. B. K.) Lag. ex Steud., 137» 315 heterostega (Trin.) Griffiths, 1 3 7 , 315 hirsuta Lag., 1 3 7 , 315 oligostachya, ( N u t t . ) T o r r , ex A. Gray, 137 parryi (Fourn.) Griffiths, 1 3 7 , 315 rothrockii Vasey, X37, 316 Brachiaria Griesb., 231, 234, 236, 316 b r i z a n t h a (Höchst, ex A. Rieh.) Stapf, 238, 239, 316 eiliatissima (Buck.) Chase, 238, 316 dietyoneura (Fig. et De Not.) Stapf, 238, 316

INDEX mutica (Forsk.) Stapf, 238, 240, 316 plantaginea (Link) Hitchc., 238, 316 reptans (L.) Gard. et C. E. Hubb., 238, 316 stolonifera Goossens, 3 1 1 Brachypodieae Harz, 25, 90 Brachypodium Beauv., go, 316 distachyon (L.) Beauv., 90, 3 1 6 Briza ¿ . , 9 5 , 96, 97, 3 1 6 maxima L., 97, 3 1 6 minor L., 97, 3 1 6 Bromeae Dumort., 25, 91 Bromus L., 91, 3 1 6 breviaristatus Buckl., 92, 3 1 6 carinatus Hook, et Arn., 92, 3 1 6 catharticus Vahl, 94 commutai us Sehr ad., 92, 316 diandrus Roth, 92 erectus Huds., 92, 3 1 6 hordeaceus L., 92, 3 1 6 inermis Ley ss., 92, 3 1 6 marginatus Nees, 92, 3 1 6 maximus Desf., 92 maximus var. gussonei (Pari.) Pari., 92 molliformis Lloyd, 92, 3 1 6 mollis L., 92, 3 1 6 polyanthus Scribn., 92, 316 racemosus L., 92, 3 1 6 rigidus Roth, 92, 93, 3 1 6 rigidus var. gussonei (Pari.) Coss. et Dur., 92 rubens L., 92, 3 1 6 squarrosus L . , 94, 3 1 6 squarrosus var. villosus (Gmel.) Reich., 94 sterilis L., 94, 316 tectorum L., 94, 316 unioloides (Willd.) H. B. K., 94, 316 villosus Forsk., 92 villosus var. gussonei (Pari.) Aschers. et Graebn., 92 vulgaris (Hook.) Shear, 94, 3 1 7 Buchloè Engelm., 134, 137, 3 1 7 dactyloides (Nutt.) Engelm., 138, 139, 3 1 7 Calamagrostis Adans., 5 1 , 52, 62, 3 1 7 canadensis var. scabra (Presi) Hitchc., 62, 3 1 7 expansa (Munro) Hitchc., 62, 63, 317 forsteri Steud., 54 hillebrandii (Munro) Hitchc., 62, 64, 3 1 7

347 langsdorfi (Link) Trin., 62, 3 1 7 retrofracta (Willd.) Link, 54 Capriola Adans., 142 dactylon (L.) O. Ktze., 146 dactylon var. maritimus Hitchc., 146 Cenchrus L., 2 3 1 , 235, 238, 3 1 7 agrimonioides Hitchc., 241 agrimonioides Trin., 241, 242, 3 1 7 agrimonioides var. laysanensis F. B. H. Brown, 241, 3 1 7 brevisetus Fourn., 241 biflorus Roxb., 241, 3 1 7 calyculatus var. uniflorus Hillebr., 241 ciliaris L., 241, 243, 3 1 7 echinatus L., 241, 244, 3 1 7 echinatus var. hillebrandianus (Hitchc.) F. B. H. Brown, 241, hillebrandianus Hitchc., 241 montanus Nees, 241 pedunculata Degener et Whitney, 34*. 245. 3 1 7 pungens H. B. K . , 241 setigerus Vahl, 241, 3 1 7 setosus Sw., 295 Centrophorum Trin., 197 Chaetochloa Scribn., 302 barbata (Lamk.) Hitchc. et Chase, 303 geniculata (Lamk.) Millsp. et Chase, 3°3 glauca (L.) Scribn., 303 lutescens (Weig.) Stünz, 303 palmifolia (Willd.) Hitchc. et Chase, 306 verticillata (L.) Scribn., 306 Chlorideae Agardh, 26, 29, 134 Chloris Sw., 134, 135, 138, 3 1 7 barbata (L.) Nash, 142 barbata (L.) Sw., 138, 140, 3 1 7 berroi Arech., 138, 3 1 7 ciliata Sw., 138, 3 1 7 culcullata Bisch., 138, 3 1 7 curtipendula Michx., 135 distichophylla Lag., 138, 3 1 7 divaricata R. Br., 138, 1 4 1 , 3 1 7 elegans H. B. K . , 142 gayana Kunth, 142, 143, 3 1 7 inflata Link, 138 paraguayensis Steud., 138 polydaetyla (L.) Sw., 142, 3 1 7 radiata (L.) Sw., 142, 144, 3 1 8 truncata R. Br., 142, 3 1 8 virgata Sw., 142, 145, 3 1 8 Chrysopogon Trin., 188, 1 9 1 , 197, 3 1 8 aciculatus (Retz.) Trin., 199, 200, 318

34« fulvus (Spreng.) Chiov., 199, 318 montanus Trin. ex Spreng., 199 montícola (Roem. et Schult.) Haines, 199 subulatus Miq., 199 Coix L., 223, 224, 318 lacryma-jobi L., 225, 226, 318 Coridochloa cimicina (L.) Nees, 235 Cortaderia Stapf, 41, 42, 318 argentea (Nees) Stapf, 42 selloana (Schult.) Aschers, et Graebn., 42, 318 Cymbopogon Spreng., 188, 192, 199, 318 citratus (DC.) Stapf, 201, 318 nardus (L.) Rendle, 201, 318 refractus (R. Br.) A. Camus, 201, 318 schoenanthus (L.) Spreng., 201, 318 Cynodon Rieh., 134, 135, 142, 318 arcuatus J. S. Presi ex C. B. Presi, 142, 318 barberi Rang, et Tad., 142, 318 daetylon Hitchc., 146 daetylon (L.) Pers., 146, 147, 318 daetylon var. maritimus (H. B. K.) Hack., 146 indicus (Gaertn.) Rasp., 155 magennisii Hur combe, 146, 318 maritimus H. B. K., 146 plectostachyus (K. Schum.) Pilger, 146, 148, 318 transvaalensis Burtt.-Davy, 146, 318 Cynosurus L., 95, 97, 318 aegyptius L., 153 cristatus L., 99, 318 indicus L . , 155 virgatus L . , 169 Dactylis L., 95, 96, 99, 318 glomerata L., 99, 100, 318 Dactyloctenium Willd., 152, 318 aegyptium (L.) Beauv., 153, 154, 318 Danthonia DC., 46, 47, 318 pilosa R. Br., 48, 318 racemosa R. Br., 48, 318 semiannularis (Labili.) R. Br., 48, 318 Danthonieae Nevski, 26, 46 Deschampsia Beauv., 73, 74, 75, 80, 319 australis Nees ex Steud., 82, 83, 319 australis f. haleakalensis Skottsb., 83, 319

INDEX australis subsp. nubigena (Hillebr.) Skottsb., 83, 319 australis subsp. nubigena var. gracilis Skottsb., 83, 319 australis subsp. nubigena var. gracilis f. major Skottsb., 83, 319 australis subsp. nubigena var. gracilis f. media Skottsb., 83, 319 australis subsp. nubigena var. gracilis f. minor Skottsb., 83, 319 australis subsp. nubigena var. pumila Skottsb., 83 australis subsp. nubigena var. tenuissima Skottsb. ; 83 australis var. pumila Rock, 83 caespitosa (L.) Beauv., 84, 85, 319 elongata (Hook.) Munro ex Benth., 84. 319 flexuosa (L.) Trin., 84, 319 hawaiiensis (Skottsb.) St. John, 83 hawaiiensis f. depauperata (Skottsb.) St. John, 84 hawaiiensis f. haleakalensis (Skottsb.) St. John, 84 nubigena Hillebr., 82, 83 pallens Hillebr., 82 pallens var. beta Hillebr., 82, 83 Deyeuxia expansa Munro, 62 forsten Kunth, 54 hillebrandii Munro, 62 retrofracta (Willd.) Kunth, 54 Dichanthium Willemet, 188, 191, 201, 319 annulatum (Forsk.) Stapf, 202, 319 aristatum (Poir.) C. E. Hubb., 202, 319 nodosum Willm., 202 sericeum (R. Br.) A. Camus, 202, 319 Dichelachne Endl., 51, 52, 62, 319 micrantha (Cav.) Domin., 65 montana Endl., 65, 319 sciurea (R. Br.) Hook, f., 65, 319 Digitaria Heister ex Fabricus, 231, 234, 246, 319 adscendens (H. B. K.) Henr., 246, 247. 319 argyrograpta (Nees) Stapf, 246, 319 bicornis (Lamk.) Roem. et Schult., 246,319 brownii (Roem. et Schult.) Hughes, 248, 319 chinensis (Nees) A. Camus, 253 consanguinea Gaud., 250 daetylon Scop., 146 debilis (Desf.) Willd., 248, 319 decumbens Stent, 248, 249, 319 diagonalis (Nees) Stapf, 248, 319

INDEX didactyla Willd., 248, 319 divaricatissima (R. Br.J Hughes, 248, 319 diversinervis (NeesJ Stapf, 248, 319 eriantha Steud., 248, 319 eriostachya Mez., 248, 319 exilis (Kippist) Stapf, 248, 319 filiformis (L.) Koeler, 248, 319 fuscescens (Presi) Henr., 248, 319 glauca A. Camus, 248, 319 henryi Rendle, 246 herpoclados Pilger, 248, 320 horizontals Willd., 248, 320 iburua Stapf, 248, 320 ischaemum (Schreb.) Schreb. ex Muhl., 248, 320 kilimandscharica Mez., 248, 320 longiflora (Retz.) Pers., 248, 320 macroglossa Henr., 248, 320 microbachne (Presi) Henr., 250, 320 milanjiana (Rendle) Stapf, 250, 320 milanjianassp. eylesiana Henr., 250, 320 monodactyla (Nees) Stapf, 250, 320 natalensis Stent, 250, 320 pentzii Stent, 250, 320 phaeothrix (Triti.) Parodi, 250, 320 pruriens (Trin.) Biise, 250, 251, 320 pseudo-ischaemum Biise, 248 sanguinalis (L.) Scop., 250, 252, 320 seriata Stapf, 250, 320 smutsii Stent, 250, 320 stolonifera Schrad., 146 swazilandensis Stent, 250, 320 timorensis (Kunth) Bai., 250, 320 tricholaenoides Stapf, 250, 320 uniglumis (Rieh.) Stapf, 250, 320 valida Stent, 250, 320 violascens Link, 253, 320 Dissochondrus (Hillebr.) Kuntze, 231, 232, 253, 320 bifidus (Hillebr.) Kuntze, 253 biflorus (Hillebr.) Kuntze ex Hack., 253. 254. 320 Echinochloa Beauv., 231, 233, 255, 320 colonum (L.) Link, 255, 256, 320 colonum var. frumentacea (Roxb.) Ridley, 258 crusgalli (L.) Beauv., 255, 320 crusgalli f. longiseta (Trin.) Farwell, 255

349 crusgalli var. cruspavonis (H. B. K.) Hitchc., 258 cruspavonis (H. B. K.) Schult., 255. 320 frumentacea (Roxb.) Link, 258, 320 polystachya (H. B. K.) Hitchc., 258, 320 polystachya var. spectabilis (Nees) M . Crovetlo, 258 stagnina (Retz.) Beauv., 258, 259, 321 walteri (Putsch) Heller, 258, 321

Ehrharta Thunb., 30, 321 calycina J. E. Sm., 31, 32, 321 erecta Lamk., 31, 321 stipoides Labili., 33 Ehrharteae Nevski, 27, 30 Eleusine Gaertn., 151, 152, 153, 321 aegyptia (L.) Desi., 153 coracana (L.) Gaertn., 155, 321 gracilis Salisb., 155 indica (L.) Gaertn., 155, 156, 321 polydactyla Steud., 155 Elymus L., 124, 125, 126, 321 canadensis L., 126, 321 condensatus J. et C. Presi, 126, 321 glaucus Buckl., 126, 321 triticoides Buckl., 126, 128, 321 virginicus L., 126, 321 Enneapogon Desi>. ex Beauv., 175, 321 brachystachyus (Jaub. et Spach) Stapf, 176, 177, 321 Enteropogon Nees, 134, 146, 321 macrostachyus (Höchst.) Munro ex Benth., 146, 321 Eragrosteae Stapf, 26, 151 Eragrostis Beauv., 151, 152, 155, 321 abyssinica (Jacq.) Link, 165 amabilis (Linn.) Wight et Arn., 165 atropioides Hillebr., 157, 158, 321 brownei (Kunth) Nees, 157, 321 caroliniana (Spreng.) Scribn., 165 chloromelas Steud., 157, 321 cilianensis (Ali.) Vignolo-Lutati, 157. 159, 321 curvula (Schrad.) Nees, 157, 321 deflexa Hitchc., 157, 160, 321 elongata (Willd.) Jacq., 157, 321 equitans Trin., 169 falcata (Gaud.) Gaud, ex Steud., 157. 321 fosbergii Whitney, 157, I6I, 321 grandis Hillebr., 157, 162, 321 grandis var. oligantha Hillebr., 157, 321 grandis var. polyantha Hillebr., 157, 321

35°

INDEX

hawaiiensis Hillebr., 169 hosakai Degener, 1 5 7 , 163, 321 intermedia Hitchc., 1 5 7 , 3 2 1 lehmanniana Nees, 1 5 7 , 322 leptophylla Hitchc., 1 5 7 , 164, 322 leptostachya (R. Br.) Steud., 165, 322 major (L.) Host, 1 5 7 mauiensis Hitchc., 165, 166, 322 megastachya (Koel.) Link, 157 mexicana (Hörnern.) Link, 1 6 5 , 322 monticola (Gaud.) Hillebr., 165, 167, 322 neomexicana Vasey, 1 6 5 , 322 niihauensis Whitney, 1 6 5 , 168, 322 paupera Jedw., 1 6 5 , 3 2 2 pectinacea (Michx.) Nees, 1 6 5 , 322 phleoides Hillebr., 169 pilosa (L.) Beauv., 1 6 5 , 322 plana Nees, 1 6 5 , 322 piumosa (Retz.) Link, 165 poaeoides Beauv., 1 5 7 , 1 6 5 , 322 superba Peyr., 1 6 5 , 322 tef (Zucc.) Trotter, 1 6 5 , 322 tenella (Link) Beauv., 1 6 5 , 322 thyrosoidea Hillebr., 169 trichodes (Nutt.) Wood, 1 6 5 , 322 unioloides (Retz.) Nees ex Steud., 1 6 5 , 322 variabilis (Gaud.) Steud., 169, 170, 322 variabilis var. beta ciliata Hillebr., 169 viahowensis T r i n . , 169 whitneyi Fosb., 169, 322 whitneyi var. caumii Fosb., 169, 322 Eragrostis Host, 1 5 5 Eremochloa Biise, 188, 190, 202, 322 ophiuroides (Munro) Hack., 202, 322 Erianthus Michx., 188, 190, 203, 322 arundinaceus (Retz.) Jeswiet, 2 1 3 maximum Brongn., 2 1 3 ravennae (L.) Beauv., 203, 322 Eriochloa Kunth, 2 3 1 , 234, 258, 322 polystachya H. B. K., 260, 322 procera (Retz.) C. E. Hubb., 260, 322 punctata (L.) Desv., 260, 322 subglabra (Nash) Hitchc., 260 Eriochrysis porphyrocoma Hance, 2 1 2 Etonia obtusata A. Gray, 87 Euchlaena Schrad., 223, 224, 225, 322 mexicana Schrad., 225, 227, 3 2 2 perennis Hitchc., 225, 322 Eulalia Kunth, 188, 190, 203 fulva (R. Br.) Kunth, 203

Festuca L . , 95, 96, 99, 323 arundinacea Schreb., 1 0 1 , 323 bromoides L . , 109 dertonensis (All.) Aschers. et Graebn., 109 drymeja Mert. et Koch, 1 0 1 dumitorium L . , 1 0 1 duriuscula Duthie, 1 0 1 elatior L., 1 0 1 , 323 fluitans L . , 1 1 2 hawaiiensis Hitchc., 1 0 1 , 102, 323 idahoensis Elmer., 1 0 1 , 323 kashmiriana Stapf, 1 0 1 , 323 megalura Nutt., 109 myuros L . , 109 octiflora Walt., 109 ovina L., 1 0 1 , 323 pratensis Huds., 1 0 1 , 323 rubra L., 1 0 1 , 323 sandvicensis Reich., 109 tenuifolia Sibth., 101, 323 viridula Vasey, 1 0 1 , 323 Festuceae Dumort., 26, 95 Garnotia Brongn., 49, 323 sandwicensis Hillebr.,

49, 50, 323

Garnotieae Tateoka, 28, 49 Gastridium Beauv., 5 1 , 65, 323 australe Beauv., 66 phleoides (Nees et Meyen) C. Hubb., 65, 323 ventricosum (Gouan) Schintz Thell., 66, 323 lendigerum (L.) Desv., 66 Ginannia

E. et

lanata (L.) F . T . Hubb., 84

Glyceria R. Br., Iii, 323 fluitans (L.) R. Br., 1 1 2 , 323 grandis S. Wats. ex A. Gray, 1 1 2 , " 3 , 323 Glycerieae Endl., 26,

III

Gramineae A. L. de Jussieu,

23

Hackelochloa O. Ktze., 188, 190, 203, 323 granulans (L.) O. Ktze., 2 0 4 , 2 0 5 , 323 Heteropogon Pers., 188, 192, 204, 323 contortus (L.) Beauv., 206, 207, 323 Hilaria H. B. K., 182, 323 belangeri (Steud.) Nash, 183, 323 cenchroides H. B. K., 1 8 3 , 323 jamesii (Torr.) Benth., 1 8 3 , 1 8 4 , 323 mutica (Buckl.) Benth., 183, 323

35 1

INDEX Holcus L., 73, 74, 84, 324 halepensis L . , 218 lanatus L., 84, 86, 324 sorghum sudanensis (Piper) Hitchc., 218 Hordeeae Spenner, 124 Hordeum L., 124, 125, 126, 324 brachyantherum Nevski, 129, 324 bulbosum L., 129, 324 murinum L., 129, 324 nodosum L., 129, 324 vulgare L., 129, 130, 324 Hyparrhenia Anderss., 188, 192, 206, 324 hirta (L.) Stapf, 208, 324 rufa (Nees) Stapf, 208, 209, 324 Imperata Cyr., 188, 189, 208, 324 cylindrica (L.) Beauv., 208, 324 Isachne R. Br., 1 1 4 , 324 distichophylla Munto, 1 1 4 , 1 1 5 , 324 distichophylla var. beta Hillebr., 1 1 4 myosotis Nees, 1 1 4 , 324 pallens Hillebr., 1 1 4 , 324 Isachneae Benth., 28, 1 1 4 Ischaemum L., 188, 190, 208, 324 byrone (Triti.) Hitchc., 210, 2 1 1 , 324 byronis Sprague, 2 1 0 ciliare Retz., 2 1 0 digitatum Btongn., 210, 324 indicum (Houtt.) Merr., 210, 324 laxum R. Br., 2 1 5 lutescens Hack., 2 1 0 secundatum Walt., 308 Ixophorus Schlecht., 2 3 1 , 232, 260, 324 unisetus (Presi) Schlecht., 260, 261, 324 Koleria glomerata Kunth, 87 Lachnagrostis phleoides Nees et Meyen, 65 Lamarkia Moench, 95, 1 0 1 , 324 aurea (L.) Moench, 103, 324 Leptochloa Beauv., 1 5 1 , 152, 169, 324 virgata (L.) Beauv., 169, 324 Leptureae Holmb., 29, 1 1 6 Lepturus R. Br., 1 1 6 , 324 cinereus Burcham, 1 1 6 , 324 repens (G. Forst.) R. Br., 1 1 7 , 324 repens var. cinereus (Burcham) Fosberg, 1 1 7 Lolium L., 95, 96, 103, 324 italicum R. Br., 103

multiflorum Lamk., 103, 104, 324 perenne L., 103, 324 rigidum Gaud., 103, 324 subulatum Vis., 103, 324 temulentum L., 103, 324 westernwoldicum Breakw., 103 Lycurus H. B. K., 5 1 , 52, 66, 325 phleoides H. B. K., 66, 67, 325 Maydeae Dumort., 24, 223 Megastachya Beauv., 155 Melica L., 1 1 8 , 325 californica Scribn., 1 1 8 , 325 Meliceae Reichb., 26, 1 1 8 Melinis Beauv., 2 3 1 , 232, 260, 325 minutiflora Beauv., 262, 325 Microlaena R. Br., 30, 3 1 , 325 stipoides (Labili.) R. Br., 33, 34, 325 Milum compressum Sw., 236 lendigerum L . , 66 Muhlenbergia Schreb., 5 1 , 52, 66, 325 fascicuìata Trin., 68 microsperma (DC.) Kunth, 68, 69, 325 porteri Scribn., 68, 325 racemosa (Michx.) B. S. Presi, 68, 325 repens (Presi) Hitchc., 68, 325 Narenga Bor, 188, 189, 210, 325 porphyrocoma (Hance) Bor, 325 Nazia Adans., 183 Neurachne montanum Gaud., 281

212,

Oplismenus Beauv., 2 3 1 , 233, 262, 325 burmannii (Retz.) Beauv., 262, 325 compositus (L.) Beauv., 263, 325 hirtellus (L.) Beauv., 263, 264, 325 polystachyus H. B. K . , 258 Oryza L., 35, 325 sativa L., 36, 325 Oryzeae Dumort., 27, 35 Oryzopsis Michx., 122, 325 coerulescens (Des}.) Hack., 123, 325 hymenoides (Roem. et Schult.) Ricker, 123, 325 miliacea (L.) Benth. et Hook, ex Aschers, et Schwein/., 123, 325 Osterdamia Neck., 186 matrella (L.) O. Ktze., 186

352

INDEX

Paniceae R. Br., 24, 231 Panicularia fluitans (L.) K u n t z e , 112 Panicum L., 231, 232, 263, 326 affine Hook, et Arn., 265 alakaiense Skottsb., 265, 326 antidotale Retz., 365, 326 aquaticum Muhl., 302 barbatum Lamk., 303 barbinode T r i n . , 238 beecheyi Hook, et Arn., 265, 266, 326 bulbosum H. B. K., 265, 326 caespitosum Sw., 238 capillare L., 265, 326 carteri Hosaka, 265, 267, 326 cinereum Hillebr., 281 colliei Endl., 265, 268, 326 colonum L . , 255 coloratum L., 265, 326 conchoideum Hillebr., 273 conjugens Skottsb., 265, 326 crusgalli L . , 255 contractum Wight et Arn. ex Nees, 302 crusgalli var. colonum Coss., 255 crusgalli var. longisetum T r i n . , 255 cynodon Reich., 265, 326 dactylon L . , 146 debile Desf., 248 degeneri Potztal, 265, 269, 326 dichotomiflorum Michx., 265, 326 fauriei Hitchc., 265, 270, 326 filiforme L . , 248 forbesii Hitchc., 265, 271, 326 frumentaceum Roxb., 258 geniculatum Lamk., 303 glaucum L . , 303 gossypinum Hook, et Arn., 281 guadaloupense Steud., 238 havaiense Reich., 273 heupueo St. John, 265, 326 hillebrandianum Hitchc., 265, 272, 326 hillebrandianum var. gracilius Skottsb., 265, 326 imberbe Poir., 303 imbricatum Hillebr., 273, 274, 326 imbricatum/. minus Skottsb., 273. 326 imbricatum var. molokaiense Skottsb., 273, 326 imbricatum var. oreoboloides Whitney, 281 isachnoides Munro, 273, 275, 326 isachnoides var. kilohanae Skottsb., 273. 326 jubiflorum T r i n . , 285 kaalaense Hitchc., 273, 276, 326 kauaiense Hitchc., 265

konaense Whitney et Hosaka, 273, 277, 326 koolauense St. John et Hosaka, 273, 326 lachnanthum Torr., 308 laevifolium Hack., 273, 326 lanaiense Hitchc., 265 lanatum Rottb., 308 leucophaeum H. B. K . , 308 leutescens Weig., 303 maximum J acq., 273, 278, 326 maximum var. trichoglume Eyles, 273, 326 mtcrobachne Presi, 250 miliaceum L., 273, 326 molokaiense Degener et Whitney, 273. 279, 326 montanum Gaud., 281 monticola Hillebr., 265 muticum Forsk., 238 nephelophilum Gaud., 273, 280, 326 nephelophilum beta var. rhyacophilum Hillebr., 273 nephelophilum beta var. tenuifolium (Hock, et Arn.) Hillebr., 273 nephelophilum beta var. xerophilum Hillebr., 285 nervosum Roxb., 306 neurodes Schult., 306 niihauense St. John, 273, 326 nodosum Willd., 265 nubigenum Kunth, 281, 282, 326 numidianum Hack., 311 oahuense Steud., 281 obtusum H. B. K., 281, 327 oreoboloides (Whitney) Skottsb., 281, 327 oreoboloides var. subimbricatum Skottsb., 281, 327 orientale (Rich.) Willd., 295 pellitoides Brown et St. John, 281, 327 pellitum Trin., 281, 327 pellitum var. pseudagrostis (Trin.) Nees, 281 pilosum Svi., 281, 327 prolutum F. Muell., 281, 327 pruriens Fisch, ex Trin., 250 pseudagrostis T r i n . , 281 purpurascens Raddi, 238 radiatius (Vickery) St. John, 281, 327 ramosius Hitchc., 265 repens L., 281, 327 reptans L . , 238 sanguinale L . , 250 scribnerianum Nash, 281, 327 spectabile Nees, 258

INDEX

stagninum Retz., 258 tenuifolium H00A. et Arn., 281, 283, 327 texanum Buchi., 281, 327 torridum Gaud., 281, 284, 327 verticillatum L . , 306 violascens K u n t h , 253 virgatum L., 281, 327 xerophilum (Hillebr.) Hitchc., 285, 286, 327 Pappophoreae Kunth, 26, 175 Pappophorum brachystachyum Jaub. et Spach., 176 Paspalidium Stapf, 231, 233, 285, 327 jubiflorum (Tritt.) Hughes, 285, 3?7 radiatum Vickery, 281 Paspalum L., 231, 234, 285, 327 almum Chase, 327 bertonii Hack., 287, 327 chinensis Nees, 253 commersonii Lamk., 287, 288, 327 compressum (Sw.) Rasp., 236 conjugatum Berg., 287, 289, 327 dilatatum Poir., 287, 290, 327 dilatatum var. decumbens Vasey, 287 distichum L., 287, 291, 327 fimbriatum H. B. K., 287, 292, 327 floridanum Michx., 287, 327 hartwegianum Fourn., 287, 327 larranagai Arech., 293 lividum Triti., 287, 328 longiflorum Retz., 248 malacophyllum Tritt., 287, 328 minutiflorum Steud., 253 notatum Flügge, 287, 328 orbiculare Forst., 293, 294, 328 ovatum Nees, 287 platense Spreng., 287 plicatulum Michx., 293, 328 plicatum Pers., 293 racemosum Lamk., 293, 328 sanguinale Lamk., 250 sanguinale var. pruriens Hook, f., 250

scrobiculatum var. commersonii (Lamk.) Stapf, 287 stoloniferum Bosc., 293 stramineum Nash, 293, 328 urvillei Steud., 293, 328 vaginatum Sw., 293, 328 vaseyanum Scrib., 293 virgatum L., 293, 328 Pennisetum L. C. Rich., 231, 234, 293, 328 asperifolium (Desv.) K u n t h , 295 cenchroides Rich., 241 ciliare (L.) Link, 241

353 clandestinum Höchst, ex Chiov., 295. 296, 328 complanatum (Nees) Hemsl., 295, 328 glaucum (L.) R. Br., 303 macrostachyum (Brongn.) Triti., 295, 328 orientale (Willd.) L. C. Rich., 295, 328 merkeri Leeke, 295 purpureum Schumach., 295, 297, 328 ruppelii Steud., 295 schimperi Steud. ex A. Rich., 295, 328 setaceum (Forsk.) Chiov., 295, 298, 328 setosum (Sw.) L. C. Rich., 295, 299. 328 typhoides (Burm.) Stapf et C. E. Hubb., 295, 328 typhoideum L . C . Rich., 295 verticillatum R. Br., 306 villosum R. Br., 295, 328

Phalarideae Kunth, 27, 119 Phalaris L., 119, 328 arundinacea L., 120, 121, 328 californica Hook, et Arn., 120, 328 caroliniana Walt., 120, 328 coerulescens Desf., 120, 328 commutata Roem. et Schult., 120, 328 minor Retz., 120, 328 paradoxa L., 120, 328 stenoptera Hack., 120 tuberosa L., 120, 328 tuberosa var. stenoptera (Hack.) Hitchc., 120 Phleum L., 51, 52, 68, 329 crinitum Schreb., 71 pratense L . , 68, 70, 329 subulatum (Savi) Aschers, et Graebn., 68, 329 Poa L., 95, 96, 105, 329 abyssinica Jacq., 165 amabilis L . , 165 ampia Merr., 105, 329 annua L., 105, 106, 329 aquatica var. americana Torr., 112 arachnifera Torr., 105, 329 brownei K u n t h , 157 bulbosa L., 105, 329 caroliniana Spreng., 165 cilianensis Ali., 157 compressa L., 105, 329 falcata Gaud., 157 flabellata (Lamk.) Rasp., 105, 329 longeradiata Hillebr., 109

INDEX

354 m a n n i i Munro, 1 0 5 , 1 0 7 , 3 2 9 monticola G a u d . , 1 6 5 nemoralis L., 1 0 5 , 3 2 9 nervosa (Hook.) Vasey, 1 0 5 , 3 2 9 pratensis L., 1 0 5 , 3 2 9 sandvicensis (Reich.) Hitchc., 1 0 5 , 108, 329 scabrella (Thurb.) Benth., 109,329 senegalensis D e s v . , 1 6 5 siphonoglossa Hack., 1 0 9 , n o , 3 2 9 tef Z u c c . , 1 6 5 tenella L . , 1 6 5 trivialis L . , 1 0 9 , 329 variabilis G a u d . , 1 6 9 Pollinia S p r e n g . , 1 9 7 fulva ( R . B r . ) B e n t h . , 1 9 9 , 203 P o l y p o g o n Desf., 5 1 , 68, 3 2 9 interruptus H. B. K., 7 1 , 3 2 9 lutosus (Poir.) H i t c h c . , 7 1 monspeliensis (L.) Desf., 5 3 , 72, 329

71,

Polytoca R. Br., 2 2 3 , 2 2 4 , 2 2 5 , 3 2 9 m a c r o p h y l l a Benth., 2 2 5 , 3 2 9 Polytrias Hack., 1 8 8 , 190, 2 1 2 , 3 2 9 a m a u r a (Buse) O. Ktze., 2 1 2 , 3 2 9 praemorsa (Nees) H a c k . , 2 1 2 Rhaphis aciculatus (Retz.) D e s v . , 199 trivialis L o u r . , 1 9 9 R h y n c h e l y t r u m Nees, 2 3 1 , 2 3 2 , 2 9 5 , 3 2 9 repens (Willd.) C. E. Hubb., 3 0 0 , 3 0 1 , 329 roseum ( N e e s ) S t a p f , 3 0 0 Rottboellia repens G . F o r s t . , 1 1 7 stolonifera Poir., 308 S a c c h a r u m L., 1 8 8 , 1 8 9 , 2 1 2 , 3 2 9 a r u n d i n a c e u m Retz., 2 1 3 , 3 2 9 barberi Jeswiet, 2 1 3 , 3 2 9 bengalense Retz., 2 1 3 , 3 2 9 edule H a s s k . , 2 1 3 , 3 2 9 fuscum R o x b . , 2 1 4 o f f i c i n a r u m L., 2 1 3 , 3 2 9 porphyrocomum (Hance) H a c k . , 2 1 2 repens W i l l d . , 300 r o b u s t u m Brandes et Jeswiet ex Grassi., 2 1 3 , 3 2 9 sinense Roxb., 2 1 3 , 3 2 9 s p o n t a n e u m L., 2 1 3 , 3 2 9 Sacciolepis Nash, 2 3 1 , contrada (Wight 302 indica (L.) Chase, Striata (L.) Nash,

232, 300, 330 et A r n . ) H i t c h c . , 302, 330 302

S c h e d o n n a r d u s Steud., 134, 135, 330 paniculatus (Nutt.) Trel., 1 4 9 , 33° S c h i z a c h y r i u m Nees, 1 8 8 , 1 9 2 , 2 1 3 , s c o p a r i u m (Michx.) Nash, 2 1 4 , S c l e r o s t a c h y a (Anderss.) A. Camus, 189, 2 1 4 , 330 f u s c a (Roxb.) A. Camus, 2 1 4 , Sclerostachyum

149, 150, 330 330 188, 330

Stapf, 214

Secale L., 1 2 4 , 1 2 9 , 3 3 0 cereale L., 1 3 1 , 1 3 2 , 3 3 0 S e h i m a Forsk., 1 8 8 , 1 9 0 , 1 9 1 , 2 1 4 , 3 3 0 n e r v o s u m (Rotti.) Stapf, 2 1 5 , 3 3 0 Setaria Beauv., 2 3 1 , 2 3 2 , 2 3 5 , 3 0 2 , 3 3 0 barbata (Lamh.) Kunth, 3 0 3 , 3 3 0 biflora H i l l e b r . , 2 5 3 geniculata (Lamk.) Beauv., 303, 304. 3 3 ° glauca (L.) Beauv., 303, 305, 330 italica (L.) Beauv., 3 0 6 , 3 3 0 longiseta Beauv., 3 0 6 , 3 3 0 lutescens (Weig.) F . T . H u b b . , 303 palmifolia (Koen.) Stapf, 3 0 6 , 3 3 0 purpurascens H . B . K . , 303 sphacelata (Schumach.) Stapfet C. E. Hubb., 3 0 6 , 3 3 0 verticillata (L.) Beauv., 306, 307, 33° S o r g h a s t r u m Nash, 1 8 8 , 1 9 1 , 2 x 5 , 3 3 0 nutans (L.) Nash, 2 1 6 , 2 1 7 , 3 3 0 S o r g h u m Moench, 1 8 8 , 1 9 1 , 2 1 6 , 3 3 0 X a l m u m Parodi, 2 1 8 , 3 3 0 c a f f r o r u m Beauv., 218 dochna (Forsk.) Snowdon, 218 dochna var. t e c h n i c u m Snowdon, 218 durra Stapf, 2 1 8 halepense (L.) Pers., 2 1 8 , 2 1 9 , 3 3 0 r o x b u r g h i i Stapf, 2 1 8 sudanense (Piper) Stapf, 2 1 8 , 3 3 0 vulgare P e r s . , 2 1 8 , 3 3 0 vulgare var. caffrorum (Retz.) H u b b . et R e h d r . , 2 1 8 vulgare var. durra ( F o r s k . ) H u b b . et R e h d r . , 2 1 8 vulgare var. roxburghii (Stapf) Haines, 2 1 8 vulgare var. saccharatum ( L . ) B o e r i . , 218 vulgare var. technicum ( K o e r n . ) J a v . , 218 Sorgum

Adans., 2 1 6

S p a r t i n a Schreber, 1 3 4 , 1 4 9 , 3 3 0 X townsendii H. et J . Groves, 33°

149,

INDEX

355

Sphenopholis Scribn., 73, 74, 87, 331 obtusata (Michx.J Scribn., 87, 88, 331 Spodiopogon Triti., 188, 189, 320, 331 aureus Hook, et Arn., 220, 331 Sporoboleae Stapf, 28, 1 7 8 Sporobolus R. Br., 178, 331 africanus (Poh.) Robyns et Tourn., 179, 180, 331 airoides (Torr.) Torr., 179, 331 berteroanus (Trin.) Hitchc. et Chase, 179 capensis (Willd.) Kunth, 179 cryptandrus (Torr.) A. Gray, 179, 331 diander (Retz.) Beauv., 179, 331 elongatus R. Br., 179, 331 fertilis, 1 7 9 fimbriatus Nees, 179, 331 flexuosus (Thurb.) Rydb., 179, 331 indicus (L.) R. Br., 179, 1 8 1 , 331 indicus, 179 poirettii (Roem. et Schult.) Hitchc., 179 virginicus (L.) Kunth, 179, 331 wrightii Munro ex Scribn., 179, 331 Stenotaphrum Trin., 2 3 1 , 233, 306, 331 americanum Schrank, 308 glabrum Trin., 308 secundatum (Walt.) Kuntze, 308, 309, 331 secundum (Osphalm.) Kuntze, 308 Stipa L., 122, 1 2 3 , 331 cernua Stebbins et Love, 1 2 3 cornata Trin. et Rupr., 1 2 3 , 331 leucotricha Trin. et Rupr., 1 2 3 , 331 papposa Nees, 1 2 3 , 331 viridula Trin., 1 2 3 , 331 Stipeae Dumort., 29, 1 2 2 Syntherisma Walt., 246 chinensis (Nees) Hitchc., 253 consanguinea (Gaud.) Skeels, 250 debilis (Desf.) Skeels, 248 helleri Nash, 253 longiflora (Retz.) Skeels, 248 microbachne (Presi) Hitchc., 250 pruriens (Trin.) Arthur, 250 sanguinalis (L.) Dulac, 250

Trichachne Nees, 2 3 1 , 234, 308, 332 californica (Benth.) Chase, 308, 332 insularis (L.) Nees, 308, 310, 332 Trichochloa microsperma D C . , 68 Tricholaena repens (Willd.) Hitchc., 300 rosea Nees, 300 Trichoneura Anderss., 1 5 1 , 152, 169, 332 elegans Suallen, 1 7 1 , 172, 332 Tridens Roem. et Schult., 1 5 1 , 152, 1 7 1 , 332 flavus (L.) Hitchc., 1 7 1 , 173, 332 Triodia flava Smyth, 1 7 1 Tripsacum L., 223, 224, 225, 332 dactyloides (L.) L., 228, 229, 332 laxum Nash, 228, 332 Triraphis R. Br., 1 5 1 , 152, 1 7 4 , 332 mollis R. Br., 174, 332 Trisetum Pers., 73, 74, 87, 332 flavescens (L.) Beauv., 87, 332 glomeratum (Kunth) Trin., 87, 89, 332 inacquale Whitney, 87, 332 Triticeae Dumort., 29, 1 2 4 Triticum L., 124, 1 3 1 , 332 aestivum L., 1 3 1 , 1 3 3 , 332 sativum Lamk., 1 3 1 vulgare Vili., 1 3 1 Urochloa Beauv., 2 3 1 , 233, 3 1 1 , 332 bolbodes (Steud.) Stapf, 3 1 1 , 332 panicoides var. panicoides Bor, 3 1 1 , 332 pullulans Stapf, 3 1 1 , 332 stolonifera (Goossens) Chippindall, 3 " Valota Adans., 308 insularis (L.) Chase, 308 Vulpia Gmel., 95, 96, 109, 332 bromoides Dum., 109 megalura (Nutt.) Rydb., 109, 332 myuros (L.) Gmel., 109, 332 octaflora (Walt.) Rydb., 109, 332

Themeda Forsk., 188, 192, 220, 331 arguens (L.) Hack., 2.2,1, 331 australis (R. Br.) Stapf, 2 2 1 , 331 gigantea (Cav.) Hack., 2 2 1 , 331 quadrivalvis (L.) Kuntze, 2 2 1 , 3 3 1 triandra Forsk., 2 2 1 , 222, 331 Trachypogon ruf us Nees, 208 Tragus Scop., 182, 183, 332 berteronianus Schult., 183, 185, 332 occidentalis Nees, 183

Zea L., 223, 228, 333 mays L., 228, 230, 333 Zizania L., 35, 333 latifolia (Griesb.) Turoz, 35, 333 Zoysia Willd., 182, 186, 323 japonica Steud., 186, 333 matrella (L.) Merr., 186, 187, 333 pungens Willd., 186 tenuifolia Willd., 186, 333 Zoysieae Miq., 29, 1 8 2