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Notes on Gramineae. 




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THE JOURNAL 




OP 



THE LINNEAN SOCIETY. 



Vol. XIX. BOTANY. Fos. 115-116. 



CONTENTS. 

I /" Page 

I /. On Central- African Plants collected by Major Serpa 
' I / Pinto. By Prof. Count Ficajlho and W. P. HiEEif, 
' * M.A., P.L.S. (Abstract reprinted.) 13 




rr Notes on Gramineas. By Gteoege Bentham, F.E.S.. . 14 — 

III. E^ort on the Arctic Drift Woods collected by Capt. 
)^ /'Fealden and Mr. Hart in 1875 and 1876. By "W". E. 
■ ' f M'Nab, M.D., P.L.S., Professor of Botany, Eoy. CoU. 
u of Science, Dublin. (With, a woodcut.) 135 



''\ LONDON: 

SOLD AT THE SOC^JETY'S APAETMBNTS, BUELINQ-TON HOUSE, 
■??" A2fl) BT 

LONGMANS, G ^EN, EEADEE, AND DTEE, 

i AND 

WILLIAMS (AND NOEGATE. 

1881. 




&7aM 



D LINNEAN SOCIETY OE LONDON. 

iJNNIVEESAEY MEETING, TUESDAY, MAY 24ih, 1881. 



Prof. Allman, LL.D., P.E.S., President, in the Ohfiir.— There was a numerous 
attendance of the FelloTPs. — The Treasurer presented his Annual Account, 
see page 3 of wrapper. — Afterwards the Secretary (Mr. B. Daydon Jackson) 
read hia Eeport. Since the last AnniTersary 11 Fellows of the Society 
had died and i had withdrawn. Against this 37 new Fellows had been 
elected, besides 1 Foreign Member and 1 Associate. During the past year 
there had been received as Donations to the Library 106 Tolurnes and 125 
pamphlets and separate memoirs. From Scientific Societies in exchange 
tliere had been received 96 volumes and 248 detached parts of publications, 
besides 23 vols, of Donations from Editors of independent periodicals. Some 
90 vols, had been purchased, viz. 80 separate and 63 serials equal to 10 vols. 
The total additions to tlie Library being 315 vols, and 373 separate parts. 
Framed water-colour sketches of Dr. Eob. Brown's birthplace, his London resi- 
dence, and of Sir Joseph Banks's Library had been presented by Mr. R. Kippist. 
Tlie Society's CoUeetions and Herbaria had been duly examined!~»id reported 
on to the Council as in good condition. After 50 years' service Sji"- Eppist 
liad resigned his position aa Librarian to the Society, and the Council in ac- 
knowledgment thereof, had gi-anted him a retiring pension. — Thereant^^ pre- 
sentation of portraits of the late Mr. John Miers and of Prof. St. George BlQvart 
were made. — Prof. Allnian then delivered his Anniversary Address, " EecSj^* 
Advances in our Knowledge of the Development of the Ctenophora." — Ti'^^ 
Secretary afterwards read Obituary Notices of the several Fellows, making*' 
special mention of Mr. E. B. Alston (late Zool. Secretary), Mr. John Gould * 
(Ornithologist), Mr. Gerrard Krefft (of Sydney), Dr. Lauder Lindsay, and K. 
A. Pryor, of Herts. — The Scrutineers having examined the ballot, then re- 1 
ported that Mr. A. W. Bennett, Mr. F. Darwin, Prof. E. E. Lankester, Sir J. 
Lubbock, and Mr. G. J. Eomanes had been elected into the Council in the room 
of E. E. Alston (deceased). Dr. T. Boycott, Prof. M. Foster, Dr. J. G. Jeffreys, 
and Prof. Mivart, who retired ; and for Officers, Sir J. Lubbock as President, 
F. Currey as Treasurer, B. D. Jackson and G. J. Eomanes as Secretaries. 



5i 



LIST OE THE OFFICEES AND COUNCIL. 



PRESIDENT. 
Sir John Lubbock, Bart., M.P., D.C.L., LL.D., F.R.S. 
VICE-PRESIDENTS. 
Prof. G. J. Allman, LL.D., P.E.S. I Charles Baron Clarke, M.A.,P.as 
George Busk, F.E.S.,F.G.S. | Frank Crisp, B.A., LL.B. 

SECRETARIES. 
B. Daydon Jackson. | George J. Eomanes, F.R.S. 

TREASURER. 
Frank Crisp, M.A., LL.B. 



COUNCIL. 



Prof. G. J. Allman, LL.D., F.E.S. 
Alfred W. Bennett, M.A., B.Sc. 
George Busk, F.E.S., F.G.S. 
Charles Baron Clarke, M.A., F.G.S. 
Frank Crisp, B.A., LL.B. 
Eev. James M. Crombie, M.A., F.G.S. 
William Sweetland Dallas. 
Francis Darwin, M.B. 



Arthur Grote, P.G.S., E.A.S. 

B. Daydon Jackson. 

Sir John Kirk, K.C.M.G. 

Prof B. Eay Lankester, P.E.S. 

Sir J. Lubbo'ek, Bart., LL.D,, P.E.i 

Eobert M-- cLachlan, F.E.S. 

George J. Eomanes, F.E.S. 



LIBRARIA'-f. 
James Murie, M.D., LL.D. 

ASSISTANT IN THE LIBRARY. 
James Vi''e8t. 



ON AFRICAN PLANTS COLLEOTBD Et MAJOA BEBPA PINTO. 13 



On Central-African Plants collected by Major Serpa Pinto. 
By Prof. Count Ficalho and "W. P. Hiben, M.A., F.L.S. 

[Abstract, read June 16, 1881.] 

The specimens herein discussed were collected by Major Serpa 
Pinto, during the month of August 1878, along the upper course of 
the river Ninda, an affluent of the Zambesi, on the west side of 
the high plateau. As regards the climate of this locality, the 
temperature is described as variable, the weather as very dry 
during seven or eight months of the year, and very wet during 
two or three months. The nature of the soil is metamorphic 
argillaceous schist; the latitude is 14° 46' S., the longitude 
20° 56' E., and the elevation 1143 metres above the ocean. 

The present little collection consists of seventy-two numbers, 
comprising sixty-five species in thirty-nine genera ; more than 
a quarter of these species are new or not previously described 
and published, and at least one new genus appears amongst 
them. Some of the specimens are imperfect and have been diffi- 
cult of final determination, especially tlio grasses and sedges ; the 
greater part have had their approximate position ascertained ; 
five specimens are hopelessly defective, and accordingly have 
been excluded from the examination. 

As in the case of the previously known species, the affinities of 
many of those of the present collection are not only with the flora 
of Huilla, in South Angola, but also, in several instances, with 
that of extratropical South Africa ; only a few of the species are 
widely distributed in the tropics of this and other continents. 

This paper, with illustrations, will appear in full in the Society's 
' Transactions.' 



LINN. JOtJEN.— botany, VOL. XIX. 



l4i ME. G. BENTHAM ON GEAMIltfi^. 

fotes on G-ramineEe. By GtEOege Bentham, P.E.S. 
[Read November 3, 1881.] 
GEAsimEiE, SO long believed to be the largest Order amongst 
Monocotyledons, must now yield the palm to Orchidefe in respect 
of number of species ; but they must still be acknowledged as 
immensely predominant, as well in individual numbers as in the 
part they take in the vegetation of the globe. The great ma,iority 
of Orchidese are very local, and amongst the few that are spread 
over wider areas it is frequently only in a few individuals dotted 
here and there; whilst a considerable proportion of Gramineas are 
almost cosmopolitan in their geographical distribution within or 
without the tropics, often covering the ground with innumerable 
individuals. OrchidesB are difficult to preserve ; collectors bring 
home but few specimens from their chief stations in tropical 
lands, aud those few often imperfect. Their study is therefore 
surrounded by many impediments, and, with the exception of the 
few European ones, is in the hands of very few botanists; whilst 
G-rasses, easily dried, abound in herbaria in specimens readily ex- 
hibiting their most essential characters ; and every local botanist 
considers himself perfectly competent to describe as new species or 
genera siiggested only by comparison with the few forms known 
to him from the same limited locality. The consequence is that 
amongst the large number of new species of Orchidete described 
of late years the great majority (always excepting garden hybrids 
or varieties) appear to be really distinct ; whilst the number of 
bad species and genera of G-raminese with which science has been 
overwhelmed is truly appalling. Looking to the future, it is only 
probable that the preponderance in number of species of Orchidese 
over Gramineae is likely to be greatly increased as well by new 
discoveries among the former, as by a critical revision of old 
species of the latter. On the other hand, although the interest 
in Orchideas has been so much intensified of late years, as well by 
the extent to which they are cultivated as by the singularities 
observed in their fertilizing-apparatus, yet their importance in 
the study of the history and development of vegetation, and in 
their application to the uses of man, remains as nothing compared 
to that of Gramineffi. 

This paramount importance of the latter Order in an economical 
point of view has called forth innumerable treatises, memoirs, and 
essays on cereals, on forage and other cultivated grasses, on 



ME. 6. BENTHAM ON GRAMI1IEJ3. 15 

meadows and pastures, on ornamental grasses, on the physiology 
and properties of the Order, &c., to which I need not further 
allude, my present object being merely to consider Graminefe with 
reference to their classification and affinities. In a systematic 
point of view, the great mistake of Linnaeus and the earlieir syste- 
matists was the attempt to regard the whole spikelet as a single 
flower, with a calyx and corolla to be compared with those of the 
more perfect Monocotyledons. Robert Brown, with his usual 
sagacity, pointed out this and other errors, and fiyst laid down 
the true principles upon which the Order could best be divided into 
tribes and genera ; but he unfortunately took up the idea that 
the so-called lower and upper palese represented three outer seg- 
ments of a perianth ; and although this theory lias long since been 
proved to be groundless, especially by Hugo Mohl, whose views 
have been fully confirmed by all subsequent careful observers, 
yet so great is the authority so deservedly attached to every 
thing that has issued from the pen of Brown, that his explana- 
tion of the structure of the spikelet is still allowed to influence 
the terminology adopted in generic and specific descriptions. 

Shortly after the publication of Brown's ' Prodromus,' Gra- 
minese were taken up by several French botanists who had 
acquired materials, rich for the time, chiefly from North America 
and the "West Indies. Some of these had already been published 
by Michaux or by Persoon, with more or less of assistance from 
Louis Claude Eichard, to whom the credit of all that is good in 
Persoon's ' Synopsis ' as well as in Mijchaux's ' Plora ' has been 
attributed by several subsequent writers. The greatest value is 
justly attached to all of the elder Eichard's observations in every 
Order that he worked up ; and there is no doubt that such assist- 
ance as he gave to tbose two works added much to their import- 
ance ; but we know that he declined to attach his name to Persoon's 
Synopsis, chiefly from an unwillingness to sanction tbe arrange- 
ment under the Linnean system, and we are by no means assured 
tbat there may not have been other details in both works which 
he did not concur in. We therefore are not justified in fixing on 
him a responsibility wbich he refused to undertake; and the 
genera and species first published by Michaux or by Persoon 
should be quoted as theirs and not Eichard's, except where 
Eichard's name is expressly attached to them. Michaux's ' Plora ' 
was published in 1803, the first volume of Persoon's ' Synopsis ' 
in 1805, both of them therefore antecedent to Brown ; but two 

d2 



16 Mte. &. fiEin^HiH Oit GBAMlNEiE. 

other special agrostologists, Desvaux and Palisot de Beauvois, bad 
ample time to avail themselves of Brown's work. Desvaux pub- 
lished his new genera in a m«moir which first appeared in abstract 
in the ' Nouveau Bulletin de la Societe Philomathique ' for 1810, 
and afterwards in full in the first volume of his second ' Journal 
de Botanique ' in 1813. Between these two periods Palisot de 
Beauvois published his 'Agrostographie,' in which he undertook 
a general arrangement of the whole Order, with definitions as well 
of the old-established genera as of a large number of new ones, 
including those of his contemporary Desvaux. The majority of 
these genera have since been adopted ; but his arrangement of 
them was far too technical and his characters often so vague, that 
they could in most instances scarcely have been identified, were it 
not for the names of the species which he refers to them and for 
the really good analytical drawings accompanying his work. As 
it is, several of his names have been misapplied by subsequent 
botanists, who have not paid sufficient attention to, or have not 
seen, those drawings. 

A few years later, three eminent botanists undertook the 
general study of Graminese. Kunth at Paris and afterwards at 
Berlin, Trinius in Germany and afterwards at St. Petersburg, 
and Nees von Esenbeck at Bonn, afterwards at Breslau, worked 
more or less contemporaneously, but with little or no communi- 
cation with each other. Kunth's 'Eevisio Graminum,' published 
in 1829 and following years, is a work not only splendidly illus- 
trated, but remarkable alike for the accuracy of detail in the descrip- 
tions of species, as for several of the views given of their structure 
and arrangement. This work, however, is so costly as to be acces- 
sible to few botanists, and the more generally known first two 
volumes of his ' Enumeratio Plantarum,' containing the Grasses, 
were unfortunately a far too hasty compilation. He had entered 
into an agreement with old Cotta for the preparation of a com- 
pact Synopsis Plantarum on the plan of Persoon's, and had 
received a considerable sum of money on account of the work ; 
but wlien it came to the actual drawing it up, Cotta insisted upon 
its being arranged according to the Linn88an system, which Kunth 
would no more agree to than did the elder Eichard in the case of 
Persoon. The Synopsis or Enumeratio was therefore still in 
abeyance when old Cotta died ; and his successors, not caring for 
the special plan adopted, insisted on an immediate return for the 
money advanced ; and I several times heard Kunth himself much 



MB. G. BENTHAM OK GEAMINEiE. 17 

bewail the necessity he was under of getting up these volumes 
without the care and study he could have wished to bestow on 
them, and which he did apply to his next volume on Cyperacese. 
Kunth also in all his works fully adopted Brown's theory as to 
the homology of the parts of the spikelet, carrying it out in detail 
to a degree which sometimes amounts almost to a reductio ad 
absurdum; as, for instance, in PiptatTiemmi and Milivm, two 
genera so closely connected in structure that they are still regarded 
by many experienced botanists as slightly diiferent sections of 
one genus. In both genera we see the whole spikelet consist of 
two similar outer glumes without the slightest rudiment of a 
flower in their axis, and of a third glume enclosing a flower and 
its palea; and yet we are told that whilst in PiptatTiermn we 
have two glumes and one flower, we must in Milium consider 
them as one glume and two flowers. 

Trinius published his ' Pundamenta AgrostographisB ' in 1820, 
something on the plan of Beauvois's ' Agrostographie,' but evi- 
dently founded on insufficient materials and bibliographical re- 
sources, and with some neglect of the already well-established rules 
of nomenclature. Prom that time, however, he devoted himself 
with the greatest zeal and increasing success to the study of the 
Order. I heard him say, apropos of some rather costly collection 
of specimens, that he would willingly sell his last coat for a new 
grass; and aU his later works, down to his last papers worked up 
in conjunction with Euprecht, and published in the Memoirs 
of the Petersburg Academy, are of the greatest value to agros- 
tologists, though he never followed them up by any general synop- 
tical view of the Order. In respect of terminology, he so far 
modifled that of Kunth, that where a glume is theoretically 
supposed to have a flower in its axil, but really has not even the 
slightest rudiment, he does not, like Kunth, call it a whole 
(neutral) flower, but only half a flower. 

Nees von Esenbeek never confined himself so exclusively to 
Q-raminese as did Trinius ; he never published any general con- 
spectus of the Order, and entered but little into general consi- 
derations of their structure and terminology ; but he described 
with great care the grasses of various tropical and other extra- 
European regions; hehad ample materials placed at his disposal, 
from the collections of Martins, Drege, Preiss, and other German 
travellers, and from the herbaria of Hooker, Arnott, and Lindley 
in this country, and he came to be regarded as the great autho- 



18 ME. G. BENTHAM ON GEAMINEiE. 

rity for the determination of exotic GraminesB. His 'Agrosto- 
graphia Brasiliensis ' is perhaps the best of all his works ; and 
his GraminesB for the ' Plora Africse australis ' is also very good. 
His generic and subgeueric groups appear to me to be often 
better, or at least more natural, than those of Kunth or Trinius ; 
although they show in some degree that tendency to multiply 
genera as well as species, which he afterwards carried to so great 
an extent ia Cyperaceae, Laurineee, and Acanthaceae. Moreover, 
he worked up the grasses of each country separately, without 
paying sufficient attention to the cosmopolitan nature of so many 
species, which thus appear under different names in his diflferent 
works. Brown's Australian Panicum semialatum, for instance, 
is raised by Nees to the rank of a genus under the name of Cori- 
dochloa in India, and that of Sluffia in South Africa, without any 
attempt at a comparison of the three plants. 

The last general Enumeration of Graminese was thatof Steudel, 
who published in 1855 the first volume of his ' Synopsis Plan- 
tarum Glumacearum,' the worst production of its kind I have 
ever met with. He was an excellent mechanical compiler ; his 
' Nomenclator Botauicus ' was a most useful work ; and if in the 
Grasses he had confined himself to collecting all the published 
species with references to or copies of their author's characters or 
descriptions, he would have rendered good service to the students 
of the Order ; but beyond that, as he was no botanist, he was 
thoroughly incompetent for the task he had undertaken. When- 
ever he met with a grass which he could not readily make out, he 
set it down as new, with a new name, and a character so carelessly 
drawn up as to render its identification hopeless without recourse 
to the specimens themselves. Several of his new genera are well- 
known species repeated in the ' Synopsis ' under their published 
names without recognition. A few, indeed, may have to be re- 
tained ; but others, again, are founded upon the grossest errors, 
as, for instance, where he describes as a caryopsis the larva 
which had eaten up the ovary and taken its place in the enlarged 
pericarp. Having, moreover, no idea of methodical arrangement, 
his work is a perfect chaos. 

Much has been done, however, for the elucidation of the Order 
in local Ploras. Already at the close of last century and the 
commencement of the present one, several continental botanists 
proposed new genera for anomalous European grasses ; but these 
were published in works which entered but little into general cir- 



ME. G. BENTHAM ON QEAMIH-EiE, IE 

culation, and were overlooked by Beauvois, Persoou, Willdenow, 
and other general systematists. Several of the same genera 
have since been reestablished, but under other names which have 
now been so long and so universally adopted, that they must be 
considered as having acquired a right of prescription to overrule 
the strict laws of priority. It would indeed be mere pedantry, 
highly inconvenient to botanists, and so far detrimental to science, 
now to substitute Blumenbachia for Sorghum, FihicMa for Oyno- 
don, Santia for Poli/^Offon, or Singlingia for Triodia. Since the 
days of Kunth, Trinius, and Wees, the most important local re- 
visions of Graminese are : Andersson's ' G-ramineae ScandinavisB,' 
Parlatore's first volume of his ' Flora Italiana,' Cosson and 
Durieu's G-lumaceous volume of the great unfinished 'More 
d'Algerie,' Doell's Grraminese for the great Brazilian Plora 
founded by Martins, and Pournier's Graminese for the Mexican 
Flora he has undertaken ; besides more partial revisions by 
Grisebach in his ' Spicilegium Flor» Eumelicse et BithynicsB,' in the 
fourth volume of Ledebour's ' Plora Eossica,' and in various con- 
tributions to the Floras of extratropical South America, the West 
Indies, the Himalayas, &c., and by Emile Desvaux in Claude 
Gay's ' Chilian Flora,' supplemented by new genera and species 
published by Philippi in various papers on Chilian plants. 
Andersson was a most acute observer, and had studied well the 
northern grasses of the old world ; but from want of access to 
a sufficiently extensive library, his synonyms, especially when 
treating of extra-Scandinavian species, are often very inaccurate. 
Parlatore's detailed monograph of Italian grasses is thoroughly 
to be relied upon when the result of his own observations; but 
unfortunately neither he nor Andersson sufficiently distinguished 
the characters they had taken from other works from those they 
had themselves verified. Old errors, for instance, in the de- 
scriptions of the style or of the ripe fruit, which it is often very 
difficult to ascertain from dried specimens, have been in several 
instances repeated by both authors, sometimes in identical terms. 
Both of them also, especially Andersson, show a great tendency to 
the multiplication of genera and siDecies. Cosson and Durieu's 
' Monograph of Algerian Grasses,' comprising the chief portion 
of those of the rich West-Mediterranean Flora, is a most valuable 
treatise, both for methodical arrangement and specific distinc- 
tions. Grisebach has also done much for the elucidation of oriental 
Graminese. In Doell's work I have been disappointed. In many 



20 ME. G. BENTHAM ON GEAMIKEJE. 

instances I cannot approve of his distinctions or combinations of 
genera or species. That may, however, be a matter of opinion 
only ; but in regard to several of the exceptional characters he 
gives, such as the five lodicules of Pariana or the three oiAris- 
tida, they have not been verified on reexamination ; in his spe- 
cific names he has not unfrequently departed from the established 
rules of nomenclature without giving any special reasons for so 
doing ; and there is a general carelessness in redaction showing, 
for instance, on several occasions that when he had found reason 
to modify his first ideas as to the limits of species, he had neglected 
to revise his manuscript accordingly. He also makes frequent 
use of the expression "partis nomine," the meaning of which 
neither Munro nor myself, nor any of our classical friends to whom 
we have applied, can make out. Eugene Fournier's ' Enumeration 
of Mexican Graminese ' is not yet published ; but being already 
printed off, and M. Fournier having obligingly supplied me with 
a copy, I feel bound, in so far as I am concerned, to treat it as 
having already taken date. He has had at his disposal rich col- 
lections of the grasses of a country where they are perhaps more 
local and varied even than in South Africa ; and he has made good 
iise of these materials, although there is still much to be learnt 
with regard to Mexican forms. "We have at Eew several, not 
only species but genera, which are not included in his work ; and 
there are not a few of his which I cannot recognize in our gene- 
rally rich Kew collections. A further comparison is also required 
with extra-Mexican genera and species, and especially with those 
of extrabropical South America. His genus Lesourdia, for in- 
stance, had already been published for a southern species by 
Philippi under the name of Soleropogon. His Trichloris is re- 
presented in the south by two species separately recognized by 
Munro and by Jean Gay as constituting a distinct genus, but 
under names hitherto unpublished, which must therefore give way 
to Fournier's. In a systematic point of view also his work would 
have been much more useful if he had more frequently given the 
characters of the tribes, genera, or other groups which he has 
modified, instead of limiting himself to dichotomous keys. These 
dichotomous keys, when carefully drawn up, are of the greatest 
use as guides or indexes to direct the botanist where to look for 
his plant, but are wholly insufiicient for its identification either 
generic or specific. For above sixty years I have had great 
experience both in using and in making them. It was with the 



ME. G. BENTnAM ON GBAMINEJE. 21 

aid of the admirable "Analyses " ia DeCandoUe's 'Flore Fran- 
§aise ' that I was enabled in 1817 and 1818 to learn botany without 
any extraneous teaching ; their principle was developed in the 
' Essay on Nomenclature and Classification ' which I published in 
1823 as a French edition of Jeremy Bentham's ' Chrestomathia,' 
and I have introduced them more or less into all my local floras. 
I am thus well aware of the great difficulties in the way of draw- 
ing them up satisfactorily, requiring much testing before their 
final revision. They are chiefly useful where all, or nearly all, 
the plants of a country or of a group are well known ; and even 
then they frequently require the repetition of the same plant 
under different branches of the key. The best genera and other 
groups are usually distinguished by a combination of characters, 
to each one of which there may be occasional exceptions, and 
these cannot be provided for in any key that presupposes limits 
definitely marked out by single characters. As a result, there 
are some of Fournier's groups which are evidently good, but to 
which we have no clue but that supplied by the species he includes 
in each. The two genera or subgenera, for instance, into which 
he divides Souteloua, Lag. {Hutriana, Trin.), are natural and well 
limited ; but the only character he gives, the prolongation of the 
rhachis of the spike beyond the last spikelet in the one and not 
in the other, is in fact variable in both groups. Of others, again, 
I can form no idea of the limits he proposes to assign them. In 
Uniola, for instance, he admits species (unknown to me) which do 
not appear from his description to have what we have been accus- 
tomed to consider as an essential character of the genus, the four 
to six empty glumes at the base of the spikelet. Where, there- 
fore, I feel obliged to differ from him in the genus to which I 
woidd refer a species, it may as often be from the inability to 
ascertain what are his views as to the limits of a genus, as from 
that difference of opinion which so frequently prevails atpongst the 
best of botanists. 

In recent days, however, we had all been led to look up to my 
much lamented friend the late General Munro as the one who 
was to unravel the intricate web into which the order had become 
involved. His ' Monograph of Bambusese ' and various detached 
papers and communications were instalments of great promise ; 
he was known to have a thorough acquaintance with species, and 
to have already formed a well-digested framework for genera and 
tribes, an important sample of which he had given in the second 



22 ME. &. BENTHAM ON GEAMINEa!. 

edition of Harvey's ' G-enera of South-African Hants.' He had 
also amassed an immense number of notes on synonyms he had 
verified, on points of structure he had ascertained, &c., as mate- 
rials for the general work he was preparing for DeCandoUe's 
Monographs. His death has extinguished all such hopes as we 
had entertained; and although liis notes, mostly dispersed in his 
herbarium or in the gramineous boots of his library, are now left 
at our disposal at Kew, yet he had unfortunately not committed 
to paper his ideas on the limits and distinctive characters of tribes, 
genera, and subgenera not included in the South-African Flora ; 
and these I could only gather from his conversation and corre- 
spondence. My own preparation for the work I have now under- 
taken was chiefly the study of European grasses for my ' Hand- 
book of the British Mora,' and of Old- World Graminese generally 
for the ' J?lora Hongkongensis ' and ' Mora Australiensis,' whenl 
was in constant correspondence relating to them with Greneral 
Munro. Having now had to work also upon American forms and 
to examine with more detail the South-European, Oriental, and 
African ones, I have had to modify in some respects the views I 
had expressed as to the relative importance and constancy of some 
of the characters, and partially to rearrange some of the tribes 
and subtribes, although the general principles of classifi.cation 
■which had been suggested by General Munro have only been con- 
firmed by further experience. 

I have already, in mypajjer on the classification of Monocoty- 
ledons (Journ. Linn. Soc. (Bot.) xv. p. 513), entered so fully into my 
reasons for adopting as to G-ramineas a terminology in accordance 
with the observations of Mohl and in harmony with that followed 
as to Cyperacess, that I need not repeat them on the present occa- 
sion. I would only add a few words in further reply to the objec- 
tion repeatedly made to me that the falling off together of the 
flowering glume and palea (commonly called the two paless) en- 
closijig the fruit, is a strong evidence of their being really homo- 
logous. But this is a mistake. A careful observation will show 
that they never do both together fall away from the rbachilla or 
axis of the spikelet ; it is the rhachilla itself that breaks up, a por- 
tion of which always remains attached to the glume and palea and 
keeps them together round the fruit. In most Panicacese, espe- 
cially in AndropogonesB, the whole spikelet with the empty glumes 
as well as the flowering one falls off with the fruit. In the majo- 
rity of Poacese the disarticulation takes place between each two 



ME. Q. BENTHAM ON GEAMINEiE. 23 

flowering glumes, leaving the intervening portion either attached 
to the glume next above it, when it is usually described as a callus 
proceeding from the glume, or to the glume next below it, when 
it is often half concealed between the keels of the palea and taken 
no notice of; or if it be a continuation of the rhachilla above the 
last glume, it is often termed a neuter or abortive flower. The 
cases where the flowering glume really detaches itself ultimately 
from the inarticulate and persistent rhachOla are very few, chiefly 
in several species of JEragrostis, where the glume and caryopsis 
fall away, leaving the palea and floral axis persistent on the rha- 
chilla. In some cases the apparently terminal fruiting glume 
enclosing the palea and caryopsis falls away without any percep- 
tible portion of the rhachilla above or below it ; but that arises 
from the disarticulation taking place so close under it that the 
fragment carried ofi' is only that minute portion actually em- 
braced by the base of the glume. 

The homology of the glumes of Gramineffi, whether empty 
or flowering, with those of Cyperacese may now be considered 
as generally admitted ; and a total absence of perianth in the 
former order might not be regarded as improbable when we have 
traced in Cyperaceee its gradual reduction from the regular hexa- 
merous perianth of Oreobolus to its absolute deficiency in Cyperus 
and others. But we have in Graminese a new element on the 
floral axis below the stamens and pistil or actual flower, in the 
palea and lodicules, for which we cannot at once find any parallel 
in other orders, and which have been very variously accounted 
for. They have very recently been the subject of a very able 
paper in Engler's ' Botanische Jahrbiieher' (i. p. 336) by Pro- 
fessor Hackel of Vienna. He comes to the conclusion that the 
palea and the pair of lodicules (when two only) are each of them 
single, more or less bifid, organs, and that they and the third lodicule, 
when present, must be regarded as two or three bracteoles inserted 
alternately fore and aft on the floral axis below the flower. And 
he has made out a good case in favour of his view, but perhaps 
not an unanswerable one. The first objection that strikes one is 
that the difficulty of finding any homologues in other orders is 
by no means diminished. In other orders where bracteoles do 
exist below the flower, they are usually lateral with reference to 
the main axis, not fore and aft, never more than two, unless when 
representing a continuation, as it were, of the sepals, and never 
developed, to my knowledge, when the perianth is suppressed ; 



24 ME. G. BENTHAM ON GBAMINE^. 

the bracts performing the functions of the deficient perianth 
are always, I believe, on the maia axis, like the glumes of Gra- 
minese. Then, again, the perfect union of the two lobes of the 
palea or of the two lodicules, or even the occasional development 
of a single central nerve or central lobe, is no absolute proof that 
they are not in fact double organs ; for where the segments of a 
perianth are united in a tube or cup, the lateral nerves of two 
adjoining segments (sepals or petals) often coalesce into a single 
one which may protrude at the top into an intermediate tooth or 
lobe. Hackel has w^ell shown that the unity or duplicity is the 
same in the case of the palea and of the two lodicules ; but it is 
only conjecturally that he continues the parallel through the third 
lodiciile, which, when present, never shows any tendency to divi- 
sion, and whose insertion is not perceptibly higher up than that 
of the two others. It is quite true that it is often much smaller 
than the other two, sometimes very minute ; but in several spe- 
cies of Stipa, in the majority of Bambuseae &c., I have seen the 
three quite equal and perfectly similar. The only instances I know 
of more than three lodicules are those of Ochlandra, where they 
are exceedingly irregular, and of Reynaudia, where I find fonr in 
two pairs, as described and figured by Kunth ; but then the outer 
pair, although closely contiguous (on the opposite side of the 
floral axis) to the upper ones, appear to me to represent the palea 
which is otherwise deficient. The minute bodies above the lodi- 
cules in the female flowers of Pariana, which Doell mistook for 
additional lodicules, a,ppear to me to be rudimentary staminodia ; 
they are very mipute and irregular, and not always to be found. 
I have observed that the search for homologues to the palea and 
lodicules in the Orders nearly allied to Graminese has met with 
but little success. The only representation of the palea that I 
can find is that mentioned in my above-quoted paper ( Journ. Linn. 
Soc. (Bot.)xv. p. 516), where it is compared with the hypogynous 
scales of Hypoelytrum pungens and Platylepis ; and I find that in 
some species of Eriocaulon (Flora A.ustraliensis, vii. p. 190) the 
perianth is composed of two outer segments inserted near the base 
of the floral axis and two or three inner ones close under the 
androecium, or these inner ones occasionally deficient, the arrange- 
ment passing gradually through other species to the normal two 
contiguous series of two or three each. It might therefore be 
suggested that the palea and lodicules of Graminese represent 
perianth-segments of an outer and inner series, although I by no 



ME. G. BEKTHAJU: ON GEAMlNEiE. 25 

means pretend to assert it as a proved fact. If the suggestion be 
confirmed, we might be justified in designating as a neutral flower 
that in which the palea alone, or the palea and lodicules without 
stamens or pistil, are developed ; but we must not include in the 
flower the bract or glume which subtends it. 

In all cases the palea, whatever its origin, is called upon in con- 
junction with the subtending glume to perform more or less of the 
functions of the deficient or absent perianth, and thus acquires a 
certain fixity of character, and requires mention in all full generic 
characters. The lodicules, on the other hand, are generally rudi- 
mentary representatives of suppressed organs having lost^all func- 
tional powers, and their slight variations in form or consistency 
are generally not even of specific importance, and they only re- 
quire mention in generic characters in the few cases where they 
have retained a greater and more constant development. 

There is much of interest in the question of the geographical 
distribution of Grasses as compared with that of Orchidese, and in 
the consideration of the causes which have produced the diifer- 
ences observed in the two Orders, amongst which perhaps the very 
different agencies through which cross-fertilization is effected may 
be most infiuential ; these questions may have also more or less 
bearing on tribual and generic arrangement ; and there are nume- 
rous observations which I should have been desirous of recording. 
This, however, would lead to speculations which it would not be 
safe to indulge in without a far more detailed and closer study of 
ascertained facts than I have time to carry out ; and I feel obliged 
to confine myself on the present occasion to the purely systematic 
consideration of real or supposed affinities and diversities. 

The division of the Order into tribes and subtril)es is a matter 
of exceptional difficulty. Whatever tribes have been proposed, 
whatever characters have been assigned to them, there have 
always been more or less ambiguous forms uniting them and 
preventing the restricting them within absolutely definite limits. 
We are obliged in Graminese, more perhaps than in any other 
Order, to rely upon combinations of characters, allowing for 
occasional exceptions in every one of our groups, preferring 
those which experience has shown to present the fewest aberra- 
tions. Tollowing up these views, none of the general divisions 
of the Order hitherto proposed have proved to be more natural 
or more definite than Brown's original primary one into two 
great groups or suborders — Fanicacees, in which the tendency to 



26 ME. 6. BEHTHAM ON GEAMINE^. 

imperfection is in the lower flowers of the spikelet ; and Poacete, 
in which the tendency is in the opposite direction. This indica- 
tion of the principle kept in view is too indefinite to serve as a 
practical character; but, combining it with that proposed by Munro 
of tlie articulation in the axis of the spikelet being below the 
spikelet itself (in the pedicel) in Panicacese, and above the lowest 
glume or none in Poacese, the exceptional forms are reduced 
to the lowest possible figure. This primary division, although 
tacitly approved of by many partial agrostologists, has not been 
generally adopted in systematic works, and many attempts have 
been made to divide the Order according to more positive cha- 
racters, but as yet with but little success. 

Kunth entirely gave up Brown's primaiy groups and divided 
the Order into thirteen tribes, many of which were natural, 
fairly deiined by a combination of characters, and bare been very 
generally adopted. Others have been objected to on various 
grounds. He attached too much importance to such characters 
as the separation of the sexes or the increase in the number of 
stamens, which are exceptional in different groups rather than 
tribual distinctions ; in the general arrangement, his removal of 
the Andropogoneae to a distance from the Panicese is disapproved 
of; and his describing flowers as actually existing when only theo- 
retically imagined is sometimes misleading. Nees generally 
adopted Kunth' s tribes, but improved the circumscription of 
some of them, and added two or three small ones. Trinius never 
completed his revised arrangement of the Order. Since the time, 
however, of these great agrostologists, systems have been sketched 
out which require a few words of notice. 

Pries, followed by Andersson, proposed for a primary division 
of Graminese that into CliscmtJiece, with the flower («. e. the 
flowering glume and palea) closed and the elongated styles pro- 
truding at the apex, and JEuryantTiece, with the glume and palea 
open at the time of flowering and the short styles protruding 
laterally. This division is, however, practically useless, except 
perhaps for the limited number of species that can be observed 
in a living state. The flowers of most species open only for a 
very short time, and in dried specimens are almost always closed. 
The styles, again, are in many cases so exceedingly slender and 
fugacious as to be very difficult to observe in dried specimens, 
except in the bud, when they have not yet attained their full 
development, or after fertilization, when they are withering away. 



MB. G. BENTHAM ON GEAMINEiE. 27 

The long styles, moreover, would place the majority of the sub- 
tribe Seslerie£B, for instance, among Panicacess, when all their 
other characters are those of Poacese ; and the species are very 
numerous in which, from the intermediate length of the styles, 
or from both the lower smooth part and the stigmatic portion, 
or the lower part alone, being described as styles, they are 
differently characterized as long or short by different writers. 

Pournier rejects both Brown's and Pries's primary divisions, 
but proposes a new one founded on the position of the lowest 
glume of the spikelet, next to the main axis in Chlorideas and 
Hordeacece, and averted from it or external in other tribes. But, 
in the first place, this relative position cannot well be ascertained 
in loosely paniculate G-raminesB, where there is so frequently a 
slight, almost imperceptible torsion of the pedicel, and, in the 
next place, in one-flowered spikelets it is often uncertain which 
is to be regarded as the lowest glume. The total number of 
glumes in the tribe Panicese, for instance, is variable, according 
to the genus or section, two, three, or four; the lowest in 
Eeimaria, and in a few species of Paspaliim, corresponds to the 
second in the majority of Paspala and a few allied genera, and to 
the third in Panioum. All these genera are included by Pournier, 
as by all others, in one and the same tribe ; and if so, are we to 
regard as the outer glume the small outer one of Panicum, called 
by some an extra bract, and an imaginary one in Paspalum and 
its allies, or the outer one of Paspalum, which is second in Pani- 
cum ? Again, in one and the same genus, the relative position of 
the outer glume and the main axis is not always constant, as, for 
instance, in Paspalitm, in Nees's section BigitaricB {JSmprosthion, 
Doell, Anastrophus, Schlecht.), the outer glume and the flowering 
one above it are external, whilst in the majority of the genus 
they are turned towards the central rib of the main axis, and 
yet the two groups are not distinguished by Pournier even as 
sections. 

Another character much insisted on of late years for tribual 
distinction is still more uncertain, the adherence of the ripe grain 
or caryopsis to the palea, as in Festuca, Bromus, &c. This is 
usually very 'conspicuous in the dry state, although even then 
the grain is often only closely embraced by the palea, and when 
moistened the adherence very generally disappears. The union 
of the two is perhaps never truly organic, and in hot water I 
have always found them readily separable without any tearing. 



28 ME. G. BENTHAM ON GBAMINEjE. 

The consequence is that there are a considerable number of 
sjiecies in which the grain has been described by some as ad- 
herent and by others as free, and which have consequently been 
transferred from one geniis to another. Yet, if not taken too 
absolutely, the character is sometimes a useful one, assisting, for 
instance, in the arrangement of the genera of some of the sub- 
tribes of the difficult tribe Pestucea?. 

Considerable importance was attached by the earlier agrosto- 
logists to the presence or absence of the awn on the back or 
apex of the flowering glume ; but this has subsequently been 
found to be subject to great variations. The spiral twist, how- 
ever, in the lower part of the awn in some genera is more con- 
stant, and in the ' Flora Australiensis ' I had taken it as an essential 
character of some tribes or subtribes ; but there are more ex- 
ceptions than I was then aware of. The awn, when present, 
is generally twisted in Audropogoneee, Tristegiueae, Agrostideae, 
and Aveuacese, and not in Pauiceae, Chloridea;, Festuceffi, or 
HordesB ; but it is sometimes very slightly so in a few species of 
the latter group, and in the former tribes, where the awn is much 
reduced, if there be any twist it is scarcely perceptible. In all 
the tribes, also, the awn is occasionally, and in the straight- 
awned ones frequently, altogether deficient ; and in some genera, 
as in Sfipa for instance, where it is usually twisted, there are 
exceptional species in which it is straight or curved only. The 
character must therefore generally be used with more or less 
of reservation. 

The partial or absolute separation of the sexes or the increase 
in the number of stamens observed in a few genera have been 
occasionally introduced amongst tribual characters ; but further 
observation has shown that they occur amongst Q-ramine£e of 
very different affinities, and have thus proved to be often of no 
more than generic value, although in one tribe, the Maydese, the 
absolute unisexuality of the spikelets may be constant, 

Differences in the size of the embryo, in the form of the so- 
called scutellum on the caryopsis (indicative, apparently, of the 
hilum of the seed), or in the longitudinal groove or cavity fre- 
quently observable on the caryopsis, have been sometimes brought 
forward as absolute generic, if not tribual, characters, and they 
may often be really important; but we know, as yet, too little 
about them to test their value fairly. Herbarium specimens 
rarely supply ripe fruits, and they have been carefully observed 



ME. Q. BENTHAM Olf flEAMtNEjl. 29 

and accurately described in comparatively few species. The cha- 
racters thus ascertained in a single one have been supposed to 
belong necessarily to the whole genus ; and when differences have 
afterwards been found in some other species, it has at once been 
generically separated, without ascertaining whether these dif- 
ferences might not be reconciled or connected through other 
species. Before, therefore, we can ascertain the real generic 
value of characters which cannot be tested in herbarium speci- 
mens, it is necessary that we should have them well and authen- 
tically described in a much greater number of species from 
actual observation. I have on several occasions had reason to 
believe that, in long-detailed descriptions drawn up by accurate 
botanists from dried specimens, the seminal characters have been 
rather guessed at on theoretical grounds, than actually verified on 
really ripe seeds. 

Following out the views of General Munro as to the general 
arrangement of the Order in as far as I have been able to ascer- 
tain them, we have divided it into tribes and subtribes, of which 
the following are the most prominent characters, omitting for 
the present exceptional forms, which occur in almost all of 
them : — 

A. Panicaceje. Spiculas cum pedicello infra glumas articulatse, 
flore fertili unico terminali, addito interdum inferiore masculo 
V. sterili. 

Tribus i. Pajiiceae. Spiculse hermaphroditse, rarius abortu unisexuales, 
spicata; v. paniculatoe, rhachi infiorescentiae inarticulata. Gluma 
florens exaristata, fruetifera indurata v. saltern exterioribus rigidior. 

Tribus ii. Maydeae. Spiculse unisexuales, masculBe terminales spicatse 
V. paniculatse v. (in Pariand) foemineam circumdantes, foeminese in- 
feriores spicatse, cum rhacheos internodio (excepta Zea) articulatim 
secedentes. 

Tribus iii. Oryzeae. Spicula: hermaphroditae v. rarius unisexuales, pani- 
culatse V. spicatse, rhachi inflorescentise inarticulata. Gluma sub flore 
summa (palea ?) uninervis v. carinata. 

Tribus iv. Tristeginese. Spiculse hermaphroditse, secus paniculse ra- 
mulos inarticulatos solitarise v. rarius geminse v. fasciculatee, cum 
pedicello articulatse. Glumae vacuse aristatse v. muticae, florens hyalina 
V. tenuiter membranacea, arista geniculata terrainata v. mutica. 

Tribus v. Zoysiese. Spiculae hermaphroditse v. nonnuUse imperfectas, 
cum rhachi inarticulata spicse simplicis sigillatira v. fasciculatim ar- 
ticulates. Gluma florens membranacea, saepius vacuis minor hyali- 
uaque. 

LINlf. JOXJEN. — BOTANY, TOL. XIX. E 



30 Me. g. bentham on geamine^. 

Subtribus 1. Anthephoreffi. Spiculm in pedicello 3-oo, in fasciculum 
deciduum conferta. Gluma florens nunc vacuis sublongior, nunc 
brevior hyalina. 
Subtribus 2. Euzoysiese. Spiculeein pedicello soUtarice, rarius gemina. 

Glmnajlorens vaouis brevior, hyalina. 
Tribus vi. Andropogonese. Spiculse secus spicse rhachin v. panicula: 
ramulos, ssepissirae geminse v. terminales ternse, in quoque pari homo- 
gamse v. heterogamae. Gluma florens vacuis minor, hyalina, ssepe 
aristata. 
B. PoAOBiB. Pedicellus infra glumas continuus. Ehacbilla 
supra glumas inferiores persistentes saepe articulata, ultra Acres 
fertiles jwoducta, stipitiformis v. glumas vacuas v. flores imper- 
fectos ferens, v. interdum flos fertilis more Pauicacearum unicus 
termiualis, sed cum gluma sua a vacuis persistentibus articulatim 
secedens. 

Tribus vii. Phalarideae. Flos hermaphroditus unicus, terminalis. 

Glumse 6 (v. 5 et palea) uninerves v. carinatse. 
Tribus viii. Agrosteae. Spiculse 1-florse, rhachilla ultra florem uuda v. 

in setam v. stipitem producta. 
Subtribus 1. Stipese. Panicula laxa v. irregulariterspiciformis. Gluma 
florens arista scepius terminata, fructifera caryopsin arete involvens. 
Rhachilla ultra florem non producta. 
Subtribus 2. Phleoidese. Panicula spiciformis densa, cylindracea v. sub- 
globosa. Gluma florens mutica v. aristis 1-3 terminata fructifera 
caryopsin laxe includens. Rhachilla interdum producta. 
Subtribus 3. Sporobolese. Panicula laxa v. ad racemum reducta, raris- 
sime spiciformis. Gluma florens mutica. Caryopsis demum saipius 
glumis apertis subdenudata. Rhachilla non producta. 
Sithtvihus 4. 'EiuagrosteiB. Panicula varia, scepius laxa. Gluma florens 
scBpius arista dorsali instructa, rarissime mutica. Caryopsis gluma 
laxe inclusa. Rhachilla stepe producta. 
Tribus ix. Isachneae. Spiculse sequaliter biflorae. GlumiE ssepius mu- 

ticae. Rhachilla ultra flores non producta. 
Tribus X. Avenese. Spiculse bi- v. pluriflorae, ssepius paniculate. Gluma; 
florentes arista dorsali v. interdum terminali sepissime instructa. 
Rhachilla ultra idores sjepius producta. 
Tribus xi. Chlorideae. Spicula; uni- v. pluriflora, secus rhachin spica- 

rum urilateraliura biseriatim sessiles, secunds. 
Tribus xii. Festuceae. Spicula? bi- v. pluriflorje, varie paniculate v. 

rarius racemosse. Gluma; florentes muticse v. aristis terminata;. 
Subtribus 1. Pappophoreje. Glumes florentes plurinerves tri- pluri- 

aristatcB, v. absque aristis quadriloba. 
Subtribus 2. Triodiese. Gluma florentes uni- v. trinerves, tridentatm, tri- 
fid(E V. triaristata. 



Me. G. BENTHAM on OEAMINEiE. 31 

Subtribus 3. Arundineae. Rhachilla sub glumis florentibus longe pilosa. 
Subtribus 4. Seslerieae. Inflorescentia spiciformis v, capituliformis, basi 

glumis vacuis v. spicis sterilibus seepius stipata. Stylus v. rami sapius 

longi tenues. 
Subtribus 5. Eragrostese. Glumte florentes trinerves. Ccetera normalia. 
Subtribus 6. Melicese. Glumai florentes tri- v. plurinerves, superiores 

dual V. plures vacua, semet involventes. 
Subtribus 7. Centothecese. Folia plana, lanceolata v. ovata, inter venas 

transverse venulosa. Glumm florentes quinque- v. plurinerves. 
Subtribus 8. Eufestuceae. Glumes florentes quinque- v. plurinerves. 

CcBtera normalia. 
Tribus xiii. Hordeese. Spiculse uni- v. pluriflorse, ad dentes seu excava- 

tiones rhacheos spicse simplicis sessiles. 
Subtribus 1. Triticese. Spicules ad nodos solitaria, tri- v. pluriflorcB, 

rarius biflorce. 
Subtribus 2. Leptureee. Spicules ad nodos solitaries, uni- v. bifiores. Spica 

tenuis. 
Subtribus 3. Elymeae. Spicules ad nodos geminee v. plures collaierales. 
Tribus xiv. Bambuse . Gramina elata, ssepius basi saltern lignosa. 

Folia plana, ssepissime cum vagina articulata. Spiculae uni- v. pluri- 

florje. Lodiculse ssepius 3. Stamina 3, 4, v. plura. 
Subtribus 1. Arundinariese. Stamina 3. Paleabicarinaia. Pericarpium 

tenne, semini adnatum. 
Subtribus 2. Eubambusese. S^amiraaC. Paleabicarinaia. Pericarpium 

tenue, semini adnatum. 
Subtribus 3. Dendrocalameae. Stamina 6. Palea bicarinata. Peri- 
carpium crustaceum v. carnosum, a semine liberum. 
Subtribus 4. Melocannere. Stamina 6 v. plura. Palea nisi glumis 

simillima. Pericarpium crustaceum v. carnosum, a semine liberum. 

T now proceed to a more detailed revision of the several tribes, 
subtribes, and genera, in the order in which I have worked theni 
up for the forthcoming part of our ' G-enera Plantarum,' to which 
I must refer for the technical characters and references, as well 
as for the synoptical clavis of the genera. 

Series A. PANICACE^. 
This .first main division of Gramiaeas is very fairly defined by 
the combination of two characters — the articulation of the pedicel 
below the spikelet or cluster of spikelets, and the single fertile 
flower apparently terminal, with or without a single male or 
sterile one below it. "Where either of these two characters fails, 
the plant should be referred to Poaceee. 

The articulation of the pedicel is usually immediately below 

e2 



32 ilE. a. BENTHAM ON SEAMINBiE. 

the lowest glume, leaving, as the spikelet falls away, a slight 
dilatation or callosity at the apex of the persistent portion. 
Sometimes it is not easily observed at the time of flowering, hut 
becomes more marked as the fruit ripens. A similar marked 
articulation has not hitherto been observed in PoaceiB, except in 
FingerhutUa. There are also a very few cases where the lowest 
glumes are reduced to slight callosities, or are so rudimentary as 
to render it difficult to say whether the articulation is in the 
pedicel or in the rhachilla. In the Getichrus group of the tribe 
Panicese, in the subtribe AnthephoresB of Zoysieae, and in some 
AndropogonesB, the articulation is not under each spikelet, but 
under a little cluster of two or more spikelets ; and in Maydese it 
is the rhachis of the spike which disarticulates under each female 
spikelet. In GramiuesB generally, however, the articulation, 
whether of the rhachis, of the pedicel, or of the rhachilla, is 
usually under the fertile spikelets or flowers only ; under the 
males it is apt to be very obscure or quite obsolete. 

The fertile flower is above spoken of as only apparently ter- 
minal, because the presence of the palea and a slight obliquity 
tend to show that the floral axis is not really the continuation of 
the rhachilla, but, as in Poacese, a secondary or axiUary branch. 
Doell says, indeed, that a continuation of the rhachilla behind 
the palea has been observed in a species of Panicum ; but I have 
never succeeded in meeting with it in any PanicacesB. In the 
tribe Oryzeae, where there is no two-nerved palea, it may still re- 
main a matter of doubt whether the floral axis is or is not distinct 
from the rhachilla — whether the uppermost scale is a glume on 
the rhachilla or a palea at the base of the floral axis. The pre- 
sence or absence of a central nerve is not an absolute test ; for it 
is occasionally, though very rarely, absent in the lower glumes. 

Panicacese have never more than four glumes, the uppermost 
one usually enclosing or subtending the fertile flower, though in 
some Andropogonese it is excessively reduced or even quite 
obsolete or rudimentary. The next under it may be empty like 
the lower ones, or may enclose a palea, a rudimentary flower, or 
a perfecb male flower, and in Beckmannia, and a very few species 
or individuals of Setaria and Panicum, this lower flower may be 
hermaphrodite, but usually, if not always, sterile. The two lower 
glumes when present are always empty. "Where the spikelets 
are unisexual, the females have only the single terminal flower, 
the males most frequently two flowers, both witii perfect 
stamens. 



ME. 0. BENTHAM ON GRAMINEJi. 33 

The tribes composing the series of Panicacese run much into 
each other, and have been very variously extended or reduced. 
We have adopted the following six, as having appeared to us to 
be rather better defined than the smaller or larger ones that have 
been proposed. 

Tribe I. PAiiTiCEiE. 

The principal character of the Paniceae, considered as a tribe 
of Panicacese, consists in the hardening of the fruiting glume. 
In several of the smaller genera, however, and even in some 
species of Panicum itself, it is membranous, but usually larger 
than the outer ones, and forming the chief covering of the fruit, 
never hyaline or much reduced as in Andropogonese. Orymopsis, 
Milium, and their allies, which were formerly included in Panicese, 
have been transferred to Agrostidese on account of the persistent 
lower glumes below the articulation. Among the other general 
characters of the tribe, the inarticulate rhachis of inflorescence is 
constant except in Stenotaphrum jwherejlaowever, the articulation 
is very tardy and not constant, so that it has often been denied. 
The flowering glume never bears the twisted awn, so general in 
Andropogoneee and Tristeginess, although in UriocMoa and a very 
few species of Panicum its obtuse apex has a short, erect, almost 
dorsal point ; the awns of Oplismenus, Ghcetaria, the section 
Ijchinochloa of Panioum, &c. are straight and terminate one or 
more of the empty glumes only. The fertile flower terminating 
the spikelet is, in the normal genera, either perfectly hermaphro- 
dite, or, at any rate, as far as I have observed, has staminodia 
round the pistil. It is only in a few of the abnormal genera 
added to the tribe that there are strictly female spikelets. 

The normal genera of the tribe may be distributed in four 
rather distinct groups, though scarcely marked enough to be 
raised to the rank of subtribes ; and to these we would add a few 
more or less abnormal genera, but little connected with each 
other, but all apparently more nearly allied to Paniceje than to 
any other tribe. 

In the first group, or Paniaece proper, we have distinguished 
eleven genera — a number somewhat arbitrary ; for much might be 
said in favour either of uniting the whole into one vast genus 
Panicum, or of dividing them still further, as some have proposed, 
into about twice as many as those here adopted, the distinctive 
characters being often either very uncertain, or such as are not 
universally recognized as generic in the Order. 



34 ME. Q. BENTHAM ON GEAMINE^. 

1. Eeimabia, Fliigge.— This old-established and universally 
acknowledged genus has generally been limited to two tropical 
and subtropical American species, with a peculiar slender habit 
and inflorescence, and characterized by having only one empty 
glume below the flowering one, and by the constant reduction of 
the number of stamens to two. It has since, however, been as- 
certained that several species which cannot well be separated from 
Faspalum have only a single lower empty glume ; and Doell has 
distinguished Beimaria chiefly by the reduction of the stamens, 
together with the form of the spikelets more acuminate and 
more closely appressed to therhachis than in any Paspalum. He 
has added, under the name of B. aberrans, a third species, which, 
with a more vigorous habit, rather invalidates the natural di- 
stinction from Faspalum, but has all the characters of Beimaria ; 
and Munro recognizes a fourth species, allied to B. aberrans, but 
with only two, or at most three, spikes to the panicle and a 
much thicker rhachis, in the Florida plant distributed by Curtis 
with the number 3566 as Paspalum vaginatmn, but probably not 
the one entered under that name in Chapman's ' Mora of the 
Southern United States.' It occurs also in Wright's Cuban 
collection under n. 3854, and may be characterized as B. oli- 
gostaohya, Munro, spicis in pedunculo 2 rarius 3 (nee 6-15), 
rhachi dilatata spiculis sublatiore. The true Paspalum vagina- 
tmn, Sw., is a synonym of P. disticJiwn, Linn. 

2. Paspalum, Linn., ranks among the large genera of tropical 
Grarainese, and in respect of the greater number of species is a 
natural one, readily distinguished from Panicimi by the inflo- 
rescence and by the technical character of the deficiency of the 
small lowest glume. It is now, however, ascertained that neither 
character is quite constant. A few Panica of the section Bra- 
cJiiaria have the inflorescence of Paspalum ; and the lowest glume 
is frequently reduced to a small callus, or is entirely deficient in 
the section Bigitaria ; and the consequence has been, that several 
species have been referred by some botanists to the one genus 
and by others to the other. These ambiguous species appear, 
however, to be best placed in Panicmi ; and all true Paspala have 
the spikelets sessile or nearly so, in two or four rows along the 
lower or outer side of the rhachis of the spikes or simple branches 
of the panicle, and they show no trace of the small lowest glume 
of Panicum. Thus defined, the number of species may be esti- 
mated at about 160, by far the greater proportion of them tro- 



ME. a. BENTHAM OS GBAMINEyE. 35 

pical Americau, a few of which are also generally spread over 
the warmer regions of the Old "World, especially P. distichwm, Linn. 
(P. vaginatvm, Sw.), which reaches southern Europe as an intro- 
duced weed. Scarcely five species can be regarded as belonging 
exclusively to the Old "World. The above estimate of the total 
number is founded chiefly on the investigations of Munro, who 
had nearly completed the working-up of the genus, and has left 
full descriptions with diagnoses and synonymy of 138 species, 
besides a few that he had left for further inquiry. Steudel 
enumerates 262 species, but nearly half of them have proved to 
be mere synonyms or very slight varieties. Doell describes in 
detail 105 Brazilian species ; but some of them are what I cannot 
consider as really distinct ; and his own views of them were any 
thing but stable, as there are several which he at one time re- 
ferred to one species and later transferred to another, forgetting 
to eliminate them from their former place, thus : — 

G-ardner, u. 2354, is repeated under P. malacophyllitm and 

P. subsesquiglvme. 
Hostmann, n. 658, under P. densiflorum and P. ccsspitoswn. 
P. distachyum, Salzmann, n. 667, under P. pumilum and P. 

divergens. 
G-ardner, n. 3496 and 3497, under P. maculosum and P. notatum. 
Q-ardner, n. 2975, under P. vaginatum, P. tropicum, and P. 

_filifoUum. 
P. ccBspitosum, Hochst., n. 1543, | ^^^^^ p n^^^^i^ ^nd P. 
P. amaxomcmn,J^m., and \ dissectim. 

P. numile, bteud., J 

Bigitaria uniflora, Salzm., n. 659, ] under P. platycaulon and 

and Spruce, n. 679, J P. furcatmn. 

P. surincmense, Hochst., n. 1283, under P. furcatmn and P. 



Spruce, n. 30, under P. chrysodactylon and P. chrysollephare. 

Tournier enumerates 40 Mexican species, of which thirteen 
are described as new ; but he is, in G-raminese, generally disposed 
to admit as distinct species forms which I perfectly agree with 
Munro in regarding as slight varieties, corresponding to what so 
many local European botanists describe as critical species. 

"With regard to the subdivision of the genus, Trinius, in his 
several revisions, distributed the species chiefly according to the 
size of the spikelets, which, however much it may afi'ect the 
general aspect of the species, is in many cases far too uncertain 
a character to be practically useful. Nees, in his ' Agrostologia 
Brasiliensis,' proposed six sections, which Doell reduced to four, 



36 ME. G. BENTHAM ON GEAMINE^. 

Munro, though he had so nearly completed his descriptions of 
species, and often indicated the sections to which he referred 
them, had not yet definitively grouped them, leaving his manu- 
scripts, for convenience of reference, in alphabetical order. We 
have adopted three sections, founded on Nees's, which appear to us 
well defined by positive characters — Uupaspalum, Gahrera, and 
Anastrophm, subdividing the first, and largest, into four groups 
or subsections, Anachyris, Opisthion, Pseudoceresia, and Oeresia, 
much less marked in their outlines, but generally speaking fairly 
natural. 

Etipaspalum comprises the great majority of the species, and is 
distinguished by the spikelets strictly secund along the rhachis of 
the spikes, with the back of the flowering glume and of the lower 
empty one (when present) turned outwards — that is, away from 
the rhachis or from its midrib ; whilst in Anastrophm, which ia- 
cludes the remainder of the genus except the monotypic Cabrera, 
the spikelets are almost distichous, and the back of the flowering 
glume and of the lower empty one turned towards the midrib of 
the rhachis. This distinction was specially relied upon by JN'ees 
under the terms spicule aAversce and spioulce inverses, and followed 
up by Doell. It is not alluded to byPournier with regard to the 
Mexican Paspala ; but, if I understand correctly his words (Graiti. 
Mex. p. vii), it nearly corresponds to the character he proposes 
for the primary division of GrramineEe. 

Anachyris, the first subsection of lEupaspalum , is a purely arti- 
ficial one, characterized solely by the having only a single empty 
glume below the flowering one. It was first proposed as a genus 
by Nees for the Paspalmn malacopliyllwm, Trin., which has all the 
habit and floral and other characters of Paspalmn except this 
single one ; and Pournier, apparently on this account, transfers 
it to the tribe Oryeem. DoeU, however, reduces it to a section 
of Paspalum under the name of HremacJiyrion, associating with it 
a few other species, some of them evidently more nearly allied to 
corresponding species of the section Opisthion than to each other. 
And even the technical character is not always constant ; for in 
P. {Eremachyrion) sesquiglume, Doell, a species closely allied to 
P. {Opisthion) maritimum, Trin., I frequently find a minute outer 
glume ; and, again, P. pallidum and P. eandidum, H. B. K., both 
of which Doell places in Eremachyrion, are scarcely to be distin- 
guished from each other except by the lowest empty glume absent 
in the one, present in the other, as originally pointed out by 



MB. G. BENTHAM ON aEAMINI!^. 37 

Kunth. Nees describes the palea (upper palea) of the typical 
Anachyris paspalodes or Paspalum malaccyphyllum as 3-iierved ; 
Pournier says it is 1-nerved. The species is very variable as to 
the size of the spikelets, the hairs or setae on the rhachis of the 
spike, &c. ; but in all the specimens I have examined I have 
uniformly found the palea normally 2-nerved. 

Opisthion, proposed by Doell as a section of Paspalum, is our 
second subsection of Eupaspalum. It includes all the typical 
Paspala with two lower empty glumes, and the rhachis of the 
spikes not dilated. The species are numerous and varied, but 
scarcely reducible to distinct groups. 

Pseudoceresia is a subsectional name I should propose for the 
genus Ceresia as understood by Elliott and other North- American 
botanists. In it the rhachis of the spikes is more or less dilated 
and concave, but green and herbaceous throughout, and the spike- 
lets are small and glabrous or nearly so. The species are few, 
including P. stoloniferuin, Bosc, P. repens, Berg., and their allies. 
Oeresia is the name we would reserve for our fourth subsection, 
being the genus Oeresia as originally established by Persoon, in 
which the dilated rhachis of the spikes is bordered by a coloured 
or smooth membranous margin, and the half-enclosed spikelets 
are larger than in Pseudoceresia and densely ciliate. Besides 
several Brazilian and other tropical species, it includes the Mexi- 
can P. cymliforme, Pourn. 

Cabrera, our second section of Paspalum, is limited to the single 
P. aureum, H. B. K. (not of Trinius), forming Lagasca's genus 
Cabrera, in which the direction of the spikelets is nearly that of 
Anastrophus ; but instead of being marginal on each side of the 
rhachis, they are deeply embedded in alternate cavities on each 
side of the midrib, on the outer or lower side of that rhachis. This 
remarkable arrangement is very well described by Lagasca, who 
was a most accurate botanist. His ' Nova Genera et Species 
Plantarum,' forming part of the ' Elenchus Horti Matritensis,' is 
a model for the clearness and conciseness of the characters given, 
which are most thoroughly to be depended upon. The work is 
quoted by Nees and by Doell, but evidently at second hand ; had 
they really read it, and had they studied Kunth's good figure and 
description, they could never have given to the P. aureum the 
new name of P. immersum, or have transferred the synonym of 
Cabrera chrysoblepharis. Lag., to the P. exasperatum, Nees, or to 
the supposed distinct P. chrysoblepliaris, Doell, both of them at 



38 ME. a. BENTHA.M ON GBAMINEa!. 

complete variance with Lagasca's description. The genus Axo- 
nopus, Beauv., sometimes given as a synonym of Ga&rfiJ-a, because 
Beauvois had suggested that P. auremn might possibly be a con- 
gener, was founded on various heterogeneous species of Paspalwn 
and Panicum ; and the name has to be wholly expunged. 

Anastrophws, our third section, was proposed as a genus under 
that name by Schlechtendahl, and inchides Nees's section Bigi- 
tariecB or Doell's MmprostUon. It is characterized by the posi- 
tion of the spikelets on the alternate margins of the narrow, 
somewhat flexuose rhachis of the spike, so as to be rather di- 
stichous than secund, and by their direction, the back of the flower- 
ing glume and of the lower empty one being turned outwards or 
away from the rhachis. The spikes are also generally several 
close together at the end of the peduncle, as in the section Bigi- 
taria of Panicum, suggesting to Nees his sectional name, which, 
however, is inconvenient as being adjective in form, and too liable 
to be confounded with the true Bigitaria. Some of the species 
have, like Odbrera, long cilia on the spikes, but have otherwise all 
the characters of Anastrophus, of which they might form a sub- 
section under the name of Lappagopsis, given by Steudel to the 
P. dissitifloricm, Trin., which he proposed as a distinct genus. The 
several species which we would include in the subsection show a 
curious diversity in the position of the cilia : in P. fastigiatwm, 
Nees, they are long on the empty glumes, none on the rhachis ; 
in P. senescens, Nees, short on the empty glumes, long on the 
rhachis ; in P. dissitiflorum, Trin., long both on the rhachis and 
on the empty glumes ; and in a few other species, referred by 
Nees and by Doell to Cabrera, although without the peculiar 
characters of Lagasca's genus, the rhachis alone is fringed with 
long cilia, the glumes having none. 

Paspalum saceharoides, Trin., referred by Kunth to Panicum, 
is one of those small-flowered species which seem to connect Pas- 
palum with Panicum {Bigitaria), whilst the long silky hairs of the 
spikes and the consistence of the glumes show an approach to the 
AndropogonecB (Sacehareis). The arrangement of the spikelets 
along the rhachis, the number of glumes, <fec. show a nearer 
affinity to Paspalum than to any other genus. 

3. AifTHiENANTiA, Beauv. {Aulaxanthus, BIL), was founded upon 
two North-American species, with the hairy inflorescence and 
membranous glumes of the section Triahaohne of Panicum, but 
without the small lowest glume of that genus j and the second 



ME. G. BENTHAM ON GEAMINEiE. 39 

glume (corresponding to the third of Panicum) usually encloses 
a palea or a male flower — a circumstance unusual in the Order, 
where the exposed glumes are almost always empty. Prom these 
I cannot separate g^nerically the South- American Leptocoryphium, 
Nees, which, besides some slight specific characters, only differs 
from the North-American species in the second glume being con- 
stantly, instead of occasionally only, empty. The genus Anthce- 
nantia thus constituted includes three species — A. villosa, Beauv. 
{Awlaxanthus ciliatus, Ell., Panicum ignoratum, Kunth), A. rufa, 
Benth. (Aulaxanthits rufus, Ell., Panicum rufum, Kunth), and 
A. lanata, Benth. {Paspdlum lanatum, H. B. K., Milium lanatum, 
Kunth, Leptocoryphiiim lanatum and L. molle, Nees). 

4. Amphicahpum, Kunth, with spikelets unisexual by abortion 
and a peculiar inflorescence, remains limited to the single North- 
American species on which the genus was founded. 

5. Eeiochloa, H. B. K. (a name having the right of priority 
over CEdipachne, Link, and Helopus, Trin.), has the habit rather of 
the section BracTiiaria oi Panicum than of Paspalum, but wants the 
small lower glume of the former genus, and difiers generally from 
both in a peculiar callous thickening of the pedicel at the articu- 
tion. There are, however, a very few species with more or less 
of this callosity, which on other accounts cannot well be separated 
from Panicum. The flowering glume has also the peculiar point 
on the obtuse apex observable in Panictim helopus, Trin., and in a 
few others, and supposed to characterize a section or genus Uro- 
chloa. It is, however, an uncertain character, both in Eriocliloa 
and in Panicum. Nearly twenty supposed species of P]riocTiloa 
have been described ; but the greater number of them are scarcely 
even varieties of the H. polystachya, H. B. K., which is widely 
spread over the warmer regions of the Old as well as the New 
World, and known under the various names of JSJ. punctata, H. 
annulata, &c. There appear also to be at least four really distinct 
species — E. distachya, H. B. K., and PI. grandiflora {Helopus, 
Trin.) from tropical America, H. trichopus, Hochst., from tropical 
Africa, and PJ. villosa, Kunth, from eastern Asia. 

6. Beckmanhia, Host, is a single species, ranging from eastern 
Europe across Eussian Asia to North America. It has been 
usually placed in Phalaridese, a tribe with which it appears to me 
to have but little connection. The habit and inflorescence are 
those of Panicum colonwm ; but it is exceptional in PanicesB as 
having both the flowers hermaphrodite; the lowest flower is. 



40 MH. a. BENTHAM ON GBAMINEiE. 

however, as far as I have observed, usually sterile ; and a similar 
character is to be found in some species or varieties of Setaria, 
and very rarely in Panioum itself, nest to which the genus appears, 
to be best placed. The synonym JoacJiimia, Ten., given by Kunth, 
was a name intended for it by Tenore, but I believe never actually 
published. Tenore figures the plant as a BecTcmannia. 

7. Panicum:, Linn., after deducting Ichnanthus, Oplismenus, 
Setaria, and several smaller genera, remains one of the larger, and 
probably the largest, among tropical Grrasses, and is still in many 
respects polymorphous. In habit and inflorescence it may be 
confounded sometimes with Paspalum, sometimes with Arundi- 
nella, or even with some Agrostese. Generally speaking, it may 
be easily recognized by technical characters ; but the most marked, 
the yery small size of the lowest empty glume, is not quite con- 
stant ; for in a few species this glume is wholly deficient as in 
Paspalum, whilst in a few others it is of the size of the second 
glume ; the hardening also of the fruiting glume and palea is in 
some species very slight. There is nothing, however, sufficiently 
definite or constant in these exceptional species to mark them out 
as intermediate genera ; and here, as in so many other cases of 
large genera of Cyperaceae and G-raminesB, we must admit the 
existence of forms which must be placed in one or the other of 
allied genera from considerations of convenience rather than of 
strict character. Taking the genus Panicum within the limits we 
have ascribed to it, nearly 800 supposed species have been pub- 
lished : Steudel enumerates 716 ; Doell has 134 Brazilian ones, 
Fournier 97 Mexican, Nees 44 South-African ; I described 54 
Australian ones ; and they are rather numerous in tropical Africa 
and Asia ; but a considerable number are repeated in several or 
even in all of these Moras, and a large proportion of Steudel's 
species are mere synonyms or blunders. The total number of 
fairly distinct species can therefore scarcely be estimated at much 
above 250. These have been variously grouped, chiefly according 
to their inflorescence ; and no less than eighteen supposed genera 
have been at different times separated from it, but are now re- 
united, either as being founded on insufficient, uncertain, or even 
mistaken characters, or as being, in our opinion, more conveniently 
regarded as sections than as genera. But, even as sections their 
limits are often as far from being absolutely definite as are those 
of the whole genus. The following eleven are those which have 
appeared to be the most distinct ; but they are aU more or less 



ItE. G. BliSTTHAM ON dRAMDiTEJJi 41 

connected by intermediate forms, and several of them would pro- 
bably be modified, and may hereafter be much improved, by a 
closer study of species than I have at present been able to bestow 
upon them. 

(1) Digitaria. Spikelets usually small and in alternate pairs or 
clusters along one side of the simple spike-like branches of the 
panicle ; those of each pair or cluster unequally pedicellate, or 
one of them almost sessile, and the lowest glume often very 
minute or sometimes quite deficient. This section was proposed 
as a distinct genus in "Walter's ' Flora Carolinensis ' under the 
name of Syntherisma, and by Eichard, in Persoon's ' Synopsis,' 
under that of Digitaria, and is still maintained as such by many 
botanists. It was founded originally on the cosmopolitan weed 
Paniov/n sanguinale, Linn., in which the spike-like branches of 
the panicle are clustered at the end of the peduncle like those of 
Oynodon and some other ChloridesB. There are now, however, 
nearly forty species to be included in the group, in many of which 
the spikes or branches are distant along the peduncle, as in 
Schedonnardus, Gynmopogon, Leptochloa, &c., among ChloridesB. 
From this tribe the structure of the pedicellate spikelets and their 
articulation always keep tliem perfectly distinct ; but there is a 
series of small-flowered species, including the Australian and 
Asiatic P. parviflorum, Br., P. tenuiflorum, Br. (Paspalum hrem- 
foliv/m, Miigge), and Paspalum minutiflorum, Steud., and two or 
three from South Africa, wbich have been almost equally well 
placed by some in Paspaltmi, by others in Panicum. As in some 
species allied to P. sanguinale, and even in some varieties of 
P. sanguinale itself, the minute outer glume is frequently abso- 
lutely deficient. The more pedicellate spikelets and the occa- 
sional, however rare, appearance of the outer glume may justify 
the placing these species rather in Panicum than in Paspalum, to 
which I referred them in the ' Elora Australiensis.' P. platy- 
carplhum, Trin., from Benin Island, with all the characters of true 
Digitaria, is remarkable for the dilated membranous rhachis of 
the spike-like branches as in the section Geresia of Paspalum. 

(2) TrichacJine. In this section, distinguished as a genus under 
that name by Nees and others, the branches of the panicle are 
simple as in Digitaria, but usually few, loose, scattered along the 
peduncle, and erect. The glumes are all, or the second ones 
alone cUiate or clothed with soft hairs as in the section Tricho- 
Icsna ; and the fruiting glume is not much hardened. The species 



42 MB. G. BENTHAM ON GBAMINEiE. 

are few : P. semialatum, Br., is widely spread over the Old World, 
for I am unable to distinguisli tlie Asiatic Ooridochloa, Nees, and 
the South- African Bluffia, Ness, from Brown's Australian species ; 
P. Qayanimii Kunth, is confined to tropical Africa; P. leueo- 
pJicBvm, H. B. K., is frequent under various names in the tropical 
and subtropical regions of the Wew and the Old World. It is a 
very variable species ; and specimens gathered at difierent stages 
of development look very different from each other, but are not 
separable into marked varieties. It was included by Beauvois in 
his genus Urochloa, and appears to have been the type of the 
proposed genera Aciearpha, Eaddi, JUriacIme, Philippi, and Holo- 
setum and Mesosetum, Steud., and is probably the principal ele- 
ment of Presl's supposed genus Alloteropsis. 

(3) Diplaria. This section is proposed for a few American species 
with a simple terminal spike-like inflorescence. The spikelets 
are sessile along the rhachis in two rows and distichous, as in the 
section AnastropJius of Paspalum, from which Diplaria differs 
technically in the presence of the small outer glume characteristic 
of Panicum. It comprises P. rottloellioides, H. B. K., P. exaratwni 
and P.ferruginemi, Trin., P.pappopTiorum, Nees, and a few others. 

(4) Thrasya, distinguished as a genus by Kunth, has a simple 
terminal spike-like inflorescence as in Diplaria ; but the rhachis is 
more or less dilated as in the section Ceresia of Paspalmm, and 
the spikelets, sessile along the midrib, although really alternate 
and biseriate, have all the appearance of being in a single row. 
The species are few, all American, and include, besides the ori- 
ginal Thrasya paspaloides, Kunth, the P. ansafmn, Trin., which 
is scarcely specifically distinct from it, P. tTirasyoides, Trin., P. 
petrcBum, Trin., and perhaps two or three others. The P. petrmmi 
forms the genus Tylothrasya of Doell, which he characterizes by 
a callous thickening of the pedicel like that of Mrioohloa ; but the 
plant is in all other respects too closely allied to the typical 
Thrasya to be generically separated, and the callosities are slightly 
prominent in various species of Panicum. 

(5) Harpostachys. The inflorescence is again simple and spike- 
like ; but the spike is more or less falcate, with the spikelets 
crowded in two or four rows along one side of the slender rhachis 
as in Chlorides, and the common peduncles are usually long and 
often clustered two or three together in the upper axils. To 
this section belong P. monostachymi, H. B. E., P. decumhens, E. 
et Schult., and P. subfalcatum, Doell. 



UR. &. tENTnAM ON GEAMINB^. 43 

The genus DimorphostaoJiys of Pournier is founded upon the 
above P. monostachyum and some other American species, which we 
should refer to the sections Digitaria or Srachiaria, but which 
he connects generically by the small lowest glume being more 
developed or differently shaped in one spikelet of each pair than 
in the others ; but the difference is often exceedingly slight, 
and the character so little connected with any other or with 
habit, that it seems difficult to attach any more than specific 
importance to it. 

(6) Bracliiaria. This section, sometimes referred to as Paspaloid 
Panica, comprises a large number of species both from the New 
and the Old World, in which the inflorescence is that which is 
regarded as specially characteristic of Paspalum : the panicle con- 
sists of a number of spike-like simple branches, distributed along 
a simple common peduncle ; but the small lowest glume of Pani- 
cum is always present. If we regard only such typical species as 
P. flavidum or P. fiuiians of Eetz, or P. paspaloides of Persoon, 
the section appears a most distinct one ; but, on the other hand, 
several such species as P. adspersum, Trin., P. argenteum, Br., 
P. Petiveri, Trin., P. poh/phyllum, Br., &c. so closely connect it 
with some of the sparingly-flowered species of JEwpanicum, as to 
make it impossible to draw a precise line of demarcation between 
the two. Amongst these intermediate forms, Paracteenmn, pro- 
posed as a genus by Beauvois, appears to be only a starved state 
of P. gracile, Br. 

P. helopus, Trin., bears on the obtuse apex of the flowering 
glume a short point, like that of most species of Eriochloa, and 
was therefore joined by Beauvois to the P. {Trichachne) semi- 
alatum, Br., to form his genus Urochloa ; but the two are in other 
respects too dissimilar to be united in one section, and P. helopus 
appears to be altogether a true Bracliiaria. 

(7) PicMiiooMoa, was regarded by Beauvois as a distinct genus, 
founded chiefly on two very widely-spread and most variable 
species, P. colonum, Linn., and P. crus-gaUi, Linn., the former 
often cultivated, the latter a most abundant tropical and sub- 
tropical weed. Both have nearly the inflorescence of the section 
PracMaria but they are coarser plants, with the spikelets densely 
crowded on the partial spikes or branches of the panicle, and the 
second and third empty glumes, in the one rarely, in the other 
very generally, terminating in long awns. It was probably on 
this account that Kunth united Beauvois's JCcJiinochloa with his 



44 ME. G. BENTHAM ON GBAMIIfE^. 

Oplismenus ; but the development of the awn has now been shown 
to be so frequently uncertain in one and the same species of 
Graminese, that the character has quite lost the absolute import- 
ance once attributed to it by Beauvois and others, and JEchinocJiloa 
is generally admitted only as a slightly distinct section of Panicmn. 
The true Oplismenus may, however, be well maintained as a sepa- 
rate genus, to which I shall presently refer. 

(8) Ptychophyllum has been well worked up as a very distinct 
section of Panicum by A. Braun. It comprises P. plicatum, Lam., 
from the Old World, P. sulcatum, Aubl., from America, and a 
few others, which, with a peculiar foliage, have more or less of 
setsB in the panicle, which seem to connect them with Setaria. 
On examination, however, these setae will be found in PtycJiopTiyl- 
lum to be merely the setiform tips of the ultimate spikelet-beariiig 
branches of the panicle, whilst the bristles or setae of Setaria are 
abortive branchlets, forming a kind of involucre below the spike- 
lets. The remaining floral characters o{ PiycJiophyllwn are entirely 
those of the loosely-panicled species of Uupanicmn. 

(9) Symenacline of Beauvois, often retained wholly or partially 
as a genus, comprises a small number of species both from the 
New and the Old "World, in which the small, very numerous 
spikelets are usually crowded in a long narrow cylindrical spike- 
like panicle. In the typical species, P. myurus, Linn., the spike- 
lets are rather acuminate and the fruiting glume scarcely hardens ; 
in P. indicum, Linn., and others the spikelets are small, and 
quite those of a large number of true Panica. 

(10) Eupaniomn. After deducting the nine preceding sections 
and the succeeding Tricholoena, which have all some distinguishing 
peculiarity, there remain a large number of sjjecies strictly normal 
in the structure of their awnless spikelets, and connected together 
by their more or less spreading panicle, the Bj)ikelets, on short 
or on slender pedicels, clustered or scattered along its simple or 
divided branches. These species, in number not far from two 
hundred, may vary much in the size of the spikelets, in the degree 
of development of the panicle, and in other minor points, but 
seem little capable of being classed in distinct subsections. They 
form Trinius's two sections Virgaria and Miliaria, characterized 
by the branches of the panicle being angular in the one, terete in 
the other — a distinction which I have been quite unable to follow 
out, at least in the dried specimens. All I have been able to 
suggest has been their distribution into seven groups or series 



ME. G. BENTHAM ON GEAMINBJl. 45 

vaguely dlstinguialied cliiefly by their inflorescence and general 
habit. Amongst the somewhat exceptional species are P. unci- 
natum, Trin. (Echinolana polystacTiya, H. B. K.), in which the 
three empty glumes are nearly equal to each other, though shorter 
than the flowering ones, and P. •pterygodiwm, Trin. (forming the 
genus Otachyriwm, Nees), in which the two lower empty glumes 
are about equal, but shorter thau the third. In all the others the 
lowest empty glume is much the shortest. Goleatxnia, from 
extratropical South America, is proposed as a genus by Griaebach 
as having dioecious flowers. I have not seen any specimen ; but 
from his description it seems to be in all other respects a true 
Panicum {Hupanicuni) ; and as he has only seen the male, evidently 
with the flowers still young, he may have overlooked the pistil, or 
its abortion may not be constant. At any rate that character 
standing alone can scarcely be sufficient to separate it generically. 
Several of the cultivated Millets are species of JSupanicum with 
large, loose, often nodding panicles. 

(11) Tricholana (including Nees's genus Bhynchelytrum), raised 
by Parlatore and some others to the rank of a genus, has the loose 
panicle of Mupanicum ; but the fruiting glumes are not much 
hardened, and the whole inflorescence is ciliate with long hairs 
as in Trichachne, on which account the oldest known species, the 
widely-spread P. Teneriffw, was originally published as a Saccha- 
rum. There are now about fifteen species known, chiefly South- 
African; but one, the above-mentioned P. TeneriffcB, extends to 
the Mediterranean region, two are East-Indian, and two or three 
South-American. Shynchelytrum, Steud., is a different genus 
from Nees's, and belongs to the TristeginesB. 

8. IcHNANTHtrs, Beauv., is so closely allied in habit and general 
character to some species of Panicum {Hupanicum) that it is 
perhaps rather in deference to the authority of all the principal 
recent agrostologists, than from any conviction of our own, that 
we retain it as a distinct genus. The character is' a purely technical 
one — a thin hyaline auricle or wing to the rhachilla on each side 
close under the flowering glume, as is observed in some species 
of Oyperus. In the species forming the section Macropteris of 
Doell these auricles are often more than half as long as the glume 
itself; in I. longiflora (Panicum longiflorum, Trin.) they are very 
small, but prominent ; in 1. pallens and its allies, forming Doell's 
section Micropteris, they are often scarcely perceptible, and 
Fournier has restored these species to Panicum, though Muuro 

LINN. JOrnN. — BOTANY, TOl. XIX. P 



46 ME. G. BENTHAM ON GEAMINE^. 

keeps them up as Ichnanthus. Two species, I. Sqffinimseggii, 
DoeU, and I. oplismenioides, Munro, are remarkable for the long 
spreading hairs, which give them a very peculiar aspect. There 
are altogether about twenty species, all tropical American. 

9. OpLisMEiors, Beauv. (OriAqpoyow, Br.), though very near the 
section Bracliiaria of Panicum, appears to be a natural genus, and 
is well characterized by the greater development of the lowest 
empty glume, which is, moreover, always awned, whilst in Panicv/m 
it is much smaller than the others and always unawned. Kunth 
adopted the genus, but, relying on the awns alone, united with it 
JEchinocUoa, in which the proportions of the glumes are the ordi- 
nary ones of Panicum, and which I have referred to above as a 
section of Panicum. Fournier adopts Kunth's view. Steudel 
and DoeU both reduce the whole to Panicum. The true Oplis- 
meni are widely spread over the warmer regions both of the New 
and the Old World, and are variable as to the number and length 
of the spikes or panicle-branches, &c. Some botanists adopt 
above thirty species, others reduce the whole to varieties of a 
single one ; it is probable that some three or four may be fairly 
distinguished as species. Hekaterosachne of Steudel is one of 
the common forms of Oplismenus. 

10. Ch^titjm, Nees, to which Doell has properly referred 
Berchtoldia of Presl as a second species, has nearly the spikelets 
of Oplismenus, to which Kunth reduces it, the outer glumes 
being much more developed and awned than the flowering ones ; 
but, besides some minor points, the inflorescence appears quite 
different enough to justify the maintaining it as a distinct genus. 
Doell considers it as a section of Panicum, with two species, one 
Brazilian, the other Mexican. Fournier retains the genus Serch- 
ioldia for the Mexican one, without comparing it with Chmtium, 
and adds two supposed new Mexican species : the one, B. Jiolci- 

formis, judging from the specimens he quotes, is one of the large 
coarse forms of Panicum {EchinocJiloa), very nearly allied to, if 
not varieties of, P. crus-galli; the other, B. opUsmenoides, is 
unknown to me, but must from his description be referable also 
to JEchinocMoa. 

11. Setaeia, Beauv., was included by the older authors in 
Panicum, and has been restored to that genus as a section by 
Steudel and by Doell, but is retained by most modern botanists 
as a well-marked natural genus, easily recognized by the dense 
spike-like panicle usually bristling with numerous setse issuing 



MB. G. BENTHAM ON GtEAMINE^. 47 

from the pedicela below the spikelets. These setse are not 
epidermal like the rigid hairs of many G-raminese, but, as ia 
Pennisetvm, are supposed to be abortive branchlets of the 
panicle, differing, however, from those of the latter genus by 
being inserted below the articulation of the pedicel, so as to 
remain persistent after the fall of the spikelet. The species are 
very variable, and a large number have been described as distinct ; 
they appear, however, to be reducible to about ten, three of which 
are common weeds over a great part of the civilized world, and a 
fourth ((S. italica) has been much cultivated as one of the Millets 
of the Mediterranean region and the Levant. The genus was 
first fully characterized by Beauvois in his ' Agrostographie,' 
chiefly from the above-mentioned common weeds ; but he had pre- 
viously published and figured, in his Elora of Oware and Benin, 
under the name of Setaria longiseta, a plant which, as far as I can 
judge without seeing the specimen, proves to be no Setaria at all, 
but the Pennisetiwi {Becheropsis) unisetum, to which I shall pre- 
sently refer. A few species or varieties of Setaria — one, for 
instance, gathered by Hildebrandt in the Sandwich Islands, allied 
to S. viridis, another, not uncommon in the Mexicano-Texan region, 
allied to S. italica — have, like the variety of 8. glauca figured by 
Trinius, 1. 195, the lower flower hermaphrodite as well as the upper 
one, which is quite exceptional throughout all genera of Paniceae 
except Bechmannia. laophorus, Schleoht. in LinnsBa, xxxi. 420, 
was founded as a genus on TTrochloa uniseia, Presl, a Mexican 
grass which we do not identify in our collections ; but Trinius 
refers it to Panicum and Fournier to Setaria, with which Schlech- 
tendal's description agrees very fairly. 

In our second or Gendhrus group of Panicese we include four 
genera, chiefly tropical or subtropical, characterized by the 
so-called involucre of bristles surrounding each spikelet or 
sometimes each cluster of two or three spikelets ; this involucre, 
supposed to represent abortive branchlets of the inflorescence, 
being placed above the articulation of the pedicel, always falls 
away with the spikelets ; the spikelets themselves are quite those 
of Panicum, the inflorescence usually a simple spike raceme or 
spikelike panicle, rarely a loose panicle of two or more pedunculate 
spikes. 12. CEifCHErs itself, as reduced from the original Linnean 
genus, consists of about a dozen species, both from the New and 
the Old "World, two or three of them of very wide geographical 
range, all characterized by the numerous bristles of the involucres 

i'2 



4» ME. G. BENTHAM ON aBAMINEJi. 

hardened and frequently more or less united at the base, the 
inner ones often broad and scale-like. In some specimens, how- 
ever, of O. cdlyculatus, Oav., and its allies the hardening appears 
so slight as to bring the genus into very close connexion with 
I'ennisetum. 

13. Pennisetum, Pers., the principal genus of the group, would 
now contain about forty species, chiefly African, amongst which 
two or three extend to the Mediterranean region, tropical or central 
Asia, or tropical America, and a very few may be endemic in Asia, 
Australia, or America. It has been at various times proposed to 
separate several genera from it, and two or three of these have been 
pretty generally adopted ; but they pass so gradually one into the 
other, and their chief characters, derived from the hairiness or 
numbers of the involucral bristles, are so little in accord with any 
other characters or habit, that the several following groups can 
scarcely be considered even as definite sections. Pennisetwm 
itself has been restricted to those species in which the bristles are 
numerous and some or all of them more or less hairy ; whilst those 
in which the whole of the bristles are perfectly glabrous form the 
genus Qymnotrix, Beauv. But however easy this distinction may 
appear at first sight, it is neither natural nor always definite. In 
a few African species proposed by Figari and De Notaris as their 
genus EriocJicBte, the whole of the setse are densely woolly-plumose; 
in some of the commoner species numerous outer setse of each 
involucre are glabrous, and as many or more or fewer of the inner 
ones are hairy. In F. flaccidum, Munro, from East India, and 
P. BentJiamianum, Steud., from tropical Africa, amongst very 
numerous glabrous ones there are generally only two or three 
hairy ones, or sometimes none at all, thus forming a gradual con- 
nexion with the true species of Qymnotrix, where the setsB are 
always quite glabrous ; and there is nothing else whatever to 
distinguish the two series even as marked sections. F. lanatum, 
Klotzsch, is a remarkable Himalayan species, in which the 
involucral bristles are few, sometimes reduced to a single long 
rigid branched one, either plumose or glabrous, showing well the 
true nature of the involucre of the genus. Penicillaria, Willd., 
often still retained as a genus, was founded upon a plant frequently 
cultivated in the Indo-African regions, which may at first sight 
appear to be abundantly distinct. The long dense cylindrical 
spilve or spike-like panicle is often above a foot long and an inch 
m. diameter, although in other cultivated specimens not above 



ME. G. -bektham: oh gbamife^. 49 

half that size. The involucres sometimes remain persistent after 
the spikelets have fallen away, and the filiform styles are remark- 
ably long; but many cultivated specimens and some East- African 
ones, possibly wild, oifer so much variety in these respects, some 
passing quite into normal Penniseta, that it seems probable that 
the peculiarities of habit have arisen from long cultivation. The 
long styles united at the base occur in other species, amongst 
which P. ( Oymnotrix) macrostachyum, Brongn., has on that account 
been proposed by Hasskarl as a genus, under the name oiSericura. 
AmpJiocJKSta of Andersson is a Gralapagos species of the Gymnotrix 
group, with small spikelets in slender pedunculate spikes, forming 
a loosely paniculate inflorescence, very different from that which 
characterizes the greater number of Penniseta, but closely con- 
nected with them through the several varieties of P. ( Grymnotrioo) 
tristachyum, Kunth. In P. {Gymnotrix) unisetum, Nees, an 
African species proposed as a genus by Eigari and De Notaris 
under the name of BecTceropsis, this peculiar inflorescence is 
carried still further, and the involucre is sometimes reduced to a 
single bristle (always above the articulation and falling away with 
the spikelet), though I usually find 2, 3, or even more bristles. 
It is probable that the plant figured by Beauvois as Setaria lov^gi- 
seta is this same species of Pennisetum. Steudel's proposed genera 
CatatTieropJiora and Oxyanthe are normal species of Pennisetum 
{Gymnotrix). 

li. PLA&iosETrM, Benth.,is a single Australian species, which 
I characterized as a genus chiefly from its peculiar inflorescence 
and habit, which prevented my retaining it in Pennisetum without 
an extension of the generic character beyond what I felt justified 
in proposing. 

15. Paeatheeia, Grriseb., is a single "West-Indian species, 
which proves to be identical with the Brazilian plant since pub- 
lished by Doell as a section of Leptachyrium of Panicum, but 
which is evidently more nearly related to Pennisetum. The 
inflorescence is a simple spike-like panicle, of which the numerous 
short articulate branchlets or pedicels are continued beyond the 
single spikelets into long awns or bristles, which fall away with 
the spikelet like the involucres of Fennisetum, thus forming in 
some sort a connexion between the Genchrus group of genera and 
the following one. 

Our third or ChamcBraphis group of Panicese consists of seven 
small genera, loosely connected by a character which may be con- 



50 ME. G-. BEJTTHAM ON GEAMINEiE. 

sidered as rather artificial than natural, but which I believe to be 
constant. The spiiielets are nearly those of Fanicum, but with 
the fruiting glume usually less hardened; the inflorescence is 
nearly that of the paspaloid Fanica or of the Chloridese, but dis- 
tinguished from the former by the rhachis of the partial spikes or 
fascicles or branches of the panicle being produced beyond the 
spikelets into a more or less rigid point. From Chloridese the 
articulation of the pedicel below the spikelet always separates the 
present group. The genera are : — 16. Echinol^na, Desr., a 
single tropical American species (H. scabra), which has quite the 
rigid single spike of some Chloridese, but the spikelets of Paniceae 
intermixed with barren ones, on which account Eudge originally 
figured the plant as a Cenchrus. The loosely paniculate species 
added to EcMnolcena by Kunth have been rightly restored to 
Fanicum by Trinius. 17. Chamjeeaphis, Br., four Australian or 
tropical Asiatic species, fully described in my ' Flora Australiensis.' 
18. Spaetina, Schreb. {Trachynotia, Mich., Limnetis, Pers., Fon- 
celetia, Thou., SolenacJine, Steud.), five or six European, African, 
or American species, chiefly maritime, has been usually placed 
amongst Chloridese ; but the spikelets themselves containing a 
single terminal flower, and the articulation of their pedicels, are 
quite those of Pauicese, fiot of Chloridese. 19. Xeeoohloa, Br., 
three Australian species, 20. Stenotaphettm, Trin. (Biastem- 
anthe, Steud.), two or three tropical maritime species, 21. Phtl- 
LOEHACHis, Trimen, a single one from Angola, and 22. Thuaeea, 
Pers. ( OrnitJiocep'halochloa, Kurz), also a single maritime species 
from the shores of the Indian and South- Pacific oceans, are all 
perfectly isolated genera whose peculiarities have been well 
pointed out. StenotapTirum is the only genus I know in the 
tribe Paniceae which has the rhachis of the inflorescence articu- 
late ; but this can usually not be perceived except in an advanced 
state, and has been denied by some botanists. I have already 
alluded (Journ. Linn. Soc, Bot. xvii. 196) to Kunth's mistake, 
which induced him to alter Persoon's name Thuarea (abridged 
from Thouars's then MS. name of Microihuarea) to Thouarea. 

There remain seven very anomalous genera, but little connected 
with each other, and still less with any other genera of Gramineae, 
but which have all more of the general character of Panicese than 
of any other tribe. They have all been well defined and illus- 
trated, and require no more than a bare enumeration on the 
present occasion. They are : — 23. Spinifex, Linn., three Austra- 



ME. G. BENTHAM ON GEAMIlfEJE. 51 

lian species, of which one extends to New Zealand and New- 
Caledonia, with a fourth from the coasts of tropical Asia closely 
allied to one of the Australian ones ; 24. Oltba, Linn., about 
twenty species, of which one is tropical African, the remainder 
tropical American, including as a section Lithachne, Beauv. 
{Strephium, Schrad., Baddia, Bertol.) ; 25. Phaetjs, Linn., five 
American species; 26. Leptaspis, Br., three or four tropical species 
from Africa, Asia, or Australia, a genus nearly allied to, but per- 
fectly distinct from, Pharus ; 27. Lygeum, Linn., a single maritime 
species from the Mediterranean region ; 28. Steeptoch.sta, 
Schrad. {Lepideilema, Trin.), and 29. Anomochloa, Brongn., 
both single Brazilian species. 

Tribe II. Matdejb. 

The grasses composing this tribe are usually erect and taU, with 
flat, long or broad leaves, the spikelets always unisexual, the 
males, in all except Pariana, in the upper part of the plant or of 
the inflorescences, the females at the base or in the lower axils, 
the grain, in all except Zea, enclosed in a hard stony case, formed 
Tariously of an outer glume or of a subtending bract. Where 
there are several fruiting spikelets in one inflorescence they are 
superposed, and each one falls away separately with the internode 
to which it is attached, the rhacbis of the spike disarticulating at 
each node. The male spikelets either wither away or remain 
persistent above at the end of the stem or on the top of the 
uppermost fruiting spikelet. The tribe is thus perfectly well 
defined and quite distinct from any other ; and the eight following 
genera of which it is composed, all tropical or American, and 
mostly small or monotypic, are likewise marked by positive cha- 
racters. 

1. Paeiana, Aubl., an American genus of about ten species, is 
in many respects anomalous. The females, as in the other genera, 
are single at each node of the articulate inflorescence ; but the 
male spikelets, instead of forming a terminal panicle, surround 
the female at each node and fall away with it. The stamens 
are also indefinite in number, ten to twenty in the spikelets 
examined, but Nees found as many as forty ; whilst in all the 
other genera of the tribe there are only the normal three. 
Doell describes the female flower as having five lodicules ; but 
here there is probably a mistake. I have never been able to 
see more than three, which are rather large; but there are 



52 ME. G. BENTHAM ON GEAMINE*. 

sometimes within them two or three very minute scales, which 
may possibly be rudimentary staminodia. Doell has also pro- 
posed to separate generically, under the name of Eremites, a 
Brazilian plant -which, from the single spike I have seen as well 
as from his description and figure, appears to be no more than 
a starved state of some true I'ariana. 

2. Coix, Linn. {Lithagrostis, Gaertn.), contains three or four 
East-Indian species closely allied to each other, one of which, 
the common " Job's tears," is widely spread over the warmer 
regions both of the New and the Old World, but in many places 
of comparatively modern introduction. The hard covering of the 
fruit here consists of the sheath of a subtending bract, the 
withered glumes as well as the internode of the rhachis remaining 
entirely enclosed within it. 

3. PoLTTOCA, Br. {Gyaihorhachis, Nees), three or four tropical 
Asiatic species in which the stony ease of the fruit is formed by 
the outer empty glume, which is completely closed over the 
remainder of the spikelet as well as the internode to which it is 
attached. The species are : — (1) P. hracteata, Br. (Coix heteroelita, 
Eoxb.), spicis masculis terminalibus ramosis, inferioribus an- 
drogynis v. fcemineis plerisque simplicibus, glumis exaristatia : 
(2) P. WalUchiana {OyathorJiachis Wallichiana, Nees), spicis 
masculis terminalibus ramosis, inferioribus androgynis v. fcemineis 
plerisque simplicibus, spicularum mascularum gluma exteriore 
longe tenuiterque aristata: (3) P. maorophylla, sp. n., spicis 
longis (omnibus ?) androgynis simplicibus, glumis aeuminatis 
exaristatis ; folia adsunt 2-pedalia, 2 poll, lata, spicae 4-6-polli- 
cares : from the Louisiade Archipelago {MacQillivray). 

4. Chionachne, Br., contains three species from tropical Asia 
or Australia, in which the hardened fruit-case is formed, as in 
Polytoca, of the outer empty glume, but the internode of the 
rhachis, instead of being completely enclosed within it, is em- 
braced only by its thickened margins, and is seen lying as it 
were in a groove of the fruit-case. 

5. ScLEEACHNE, Br., is a single Javan species, with the 
fruit nearly of Chionachne, but with a different habit, and the 
hardened outer glume is produced beyond the fruit into an open 
membranous appendage. 

6. Teipsacum, Linn., consists of two or three American species 
with the terminal male inflorescence usually more branched than 
in the preceding Asiatic genera, approaching that of EucUcena 



MB. G. BENTHAM ON GHAMINEJ3. 53 

and Zea ; and the hardened fruit-case is formed partly only by 
the outer glume, and partly also by the broad thickened and 
hardened internode of the rhachis. 

7. EuCHLiEifA, Schrad. {Beana, Brign.), has, like Zea, the ter- 
minal male inflorescence paniculate with numerous spikelets, and 
the female spikes in the lower axils wrapped up in broad bracts, 
from which are protruded the long filiform styles ; but, as in the 
preceding genera, the female spikelets are within each bract 
superposed in a single row on the articulate rhachis of the single 
spike. The affinity to Zea appears to be recognized in the 
country ; for specimens have been received from Schaffner pur- 
porting to be known as " wild maize." 

8. ZiA, Linn. {Mays, Gsertn.). — This most important, widely 
diffused, and most striking grass is only known in a cultivated 
state, or perhaps as an escape from cultivation. With most of the 
general characters of the tribe to which it gives its name, it 
is exceptional not only in that tribe, but in the whole Order, 
by the manner in which its numerous female spikelets are densely 
packed in several vertical rows round a central spongy or corky 
axis. How far this arrangement may have gradually arisen after 
so many centuries of cultivation can only be a matter of conjec- 
ture. Its gradual progress cannot be traced through the nume- 
rous cultivated varieties, many of them described as species in 
Eonafous's splendidly illustrated monograph ; and the idea that 
some of them are wild indigenous forms must be traced to the 
insufficiency of the observations recorded by travellers. 

Tribe III. OEYZE.ffi. 

This tribe, as originally constituted, was loosely characterized, 
chiefly by uniflorous spikelets and stamens more than three — a 
character more or less dispersed through various different tribes ; 
and several of the genera included in it by Kunth have since been 
rejected. The close affinity of Oryzese and Phalarea; has also been 
recognized, though the limits of the latter tribe also have been very 
unsettled. In the ' Flora Australiensis ' I had united the two as 
an intermediate tribe, connecting, as it were, the two great primary 
series of Panicacese and Poacese ; but upon the whole it seems better 
to separate them as tribes technically distinct, but representative 
of each other in the two great series. The essential character of 
both resides in having the scale immediately under the single ter- 
minal perfect flower keeled or 1-nerved like the glumes, so as to 



54 ME. G. BENTHAM ON QEAMINE^. 

leave it uncertain wiietlier it is a glume or a palea — ttat is, whether 
it be attached to the end of the rhachilla or primary axis of the 
spitelet, or to a secondary or floral axis reduced to a mere point. 
There are theoretical reasons in favour of both explanations, and 
actual observation is insufficient for determining the point. The 
first of these views has appeared to me the most plausible ; and I 
have accordingly in my diagnoses and descriptions treated the 
scale in question as the flowering glume, and considered the 
palea as deficient, as it certainly is in some Andropogonese and 
Agrosteae. In this view the technical distinction between the 
two tribes would be, that the Oryzese have 2, 4, or rarely 3 glumes, 
all above the articulation of the pedicel, and the Phalarese 4, 6, or 
rarely 5 glumes, the lowest pair persistent below the articulation 
of the rhachilla. Oryzese thus characterized may be thought as a 
whole to be a rather artificial tribe ; but they are divisible into 
two much more natural groups or subtribes — Zizaniea, tropical or 
American genera, often semiaquatic plants, with a loose in- 
florescence and stamens often, but not always, more than three ; 
and AlopeourecB, European or temperate Asiatic or African genera, 
with a dense spike-like inflorescence and stamens never more than 
three. 

ZizaniesB includes the following eight genera : — 

1. Htdeochloa, Beauv., a single species from Carolina, and 
there apparently rare, diifering from Zizania chiefly in the inflo- 
rescence reduced to few-flowered spikes, of which the terminal 
one male and pedunculate, the lower ones female and sessile in 
the axils. 

2. ZizANiA, Linn., comprises two species, or according to others 
two genera, each with two or more species. As a whole, the genus 
is a natural one, well characterized by the unisexual spikelets in 
an androgynous panicle, each one with only two glumes and the 
males with six stamens. The typical Z. aquatiea, Linn. {Sydro- 
fyrwm. Link), has the lower part of the panicle more spreading 
and male, and the upper part narrow and female ; it is widely 
spread over North America, and includes the East-Eussian and 
Japanese Z. latifolia, which is absolutely identical with some 
North-American specimens. The other species, Z. miliacea, 
Kunth {Zizaniopsis, Doell), has the male and female spikelets 
more mixed in the panicle, the awns shorter, the styles more 
connate, and the grain broader — characters which appear to me 
quite insufficient for generic distinction. It is a North-American 



ME. a. BENTHAM ON GEAMINE^. 55 

plant, and possibly also South- American, if Sello's single specimen 
described by Trinius and figured by Doell as Z. microstacyJis, 
Nees, was really from Soutb Brazil. I see nothing in the figure 
or description to distinguish it from Z. miliacea. 

3. LuzioiiA, Juss., has, like Zizania, unisexual spikelets with 
only two glumes ; but the spikelets are smaller, not awned, the 
styles short and quite distinct, and there are usually more than 
six stamens in the males. Six species are known from tropical 
America or the southern States of North America. The relative 
arrangement of the males and females varies as in Zizania. In 
the typical L. peruviana, Juss. (i. hrasiliensis, Moric), in L. ala- 
hamensis, Ohapm., and ia an apparently unpublished G-uiana 
species, both sexes are in terminal panicles, but on distinct stems. 
In L. Spruceana, Benth., described by Doell (figured by G-. P. W. 
Meyer as L. peruviana, but not Jussieu's plant), the males are in 
^terminal panicle, whilst the females are in the lower axils of 
the same stem, as they are also said to be in L. longivalviSjUoeW, 
a Brazilian plant which I have not seen. In the proposed genus 
Caryochloa, Trin. {Arrozia, Schrad.), also Brazilian, the males and 
females are in the same panicle, the former in the upper, the latter 
in the lower part. The stamens in this species appear also to be 
always six only, which only occasionally occurs in the others ; but 
the other characters are entirely those of Luziola, to which I 
should unite the Caryochloa as L. mierantha (Arrozia micraniha, 
Schrad.). 

4. PoTAMOPHiLA, Br., if we include in it Maltelrunia, Kunth, 
is a natural genus of three species, connecting in some measure 
Zizania, of which it has the habit, with Oryza, of which it has the 
small setaceous or acuminate outer glumes. In the typical 
P. parmfiora, Br., from Australia, the spikelets are more or less 
polygamous, though the greater number appear to be her- 
maphrodite ; in P. leersioides {Maltelrunia leersioides, Kunth) 
from Madagascar, and in P. preJiensilis (Maltehrunia prehensilis, 
Nees) from South Africa, they are usually all, or nearly all, her- 
maphrodite. Kunth also distinguishes Potamophila from Malte- 
hrunia as having two flowers to the spikelet, a character not 
mentioned by Brown and which I liave been unable to verify. 
The spikelets figured by Kunth, Eev. Gram. t. 5. figs. 1, 2, & 5, 
must be very rare and probably abnormal ; I have searched in 
vain for them both in Brown's and in Beckler's specimens. 

5. Htgeoehiza, Ifees {Fotamochloa, G-rifi'.), is a single East- 



56 ME. G. BBNTHAM ON GEAMINE^. 

Indian semiaquatic species nearly allied to Zizania, but quite 
distinct in its hermaphrodite flowers and other characters. 

6. Oktza, Linn. {Padia, Zoll. and Mor.), an Asiatic genus, of 
which the typical species, the well-known Eice, appears to be 
really indigenous in Australia as well as in East India ; but it 
has been so much cultivated from time immemorial, that it is now 
found apparently wild in various parts of Africa and America. 
It has produced a large number of difi'erent forms, nearly twenty 
of which have been published as substantive species, all of which, 
or nearly all, are reduced by others to varieties of O. sativa. The 
Himalayan 0. coarctata, Grifi"., appears, however, to have more 
positive characters 5 and possibly two or three others may be 
maintained as fairly established species. 

7. Leeesia, Swartz (HomalocencArus, Mieg., Ehrartia, Wigg., 
Asprella, Schreb., Blepharochloa, Endl.), is essentially American; 
but the two commonest species — L. hexandra in tropical, L. orf- 
zoides in more temperate regions — are widely spread also over the 
Old "World, and had probably long been so before the civilized 
communication between the two continents. The genus is closely 
connected with the Asiatic Oryza ; but, besides the apparent 
diversity in geographical origin, the smaller spikelets with thinner 
glumes and the general inflorescence give to Leersia a difi'erent 
aspect, and, in technical character, the want of the two small outer 
glumes may justify its retention as a distinct genus. It is true 
that those who unite it with Oryza maintain that these outer 
glumes are represented by a cartilaginous ring at the base of the 
spikelet ; but this ring is often so slight as to be rather imaginary 
than real, and never more than what is observable in Eriochloa 
and some other G-raminesB, where no such theoretical explanation 
is wanted or attempted. 

8. AcHLiENA, Griseb., is a single Cuban species, which the 
author compares with the Australian Microlosna ; but the want of 
any glumes below the articulation places it in Oryzese, not in 
Phalaridese. It is in some other respects allied to Oryza and 
Leersia; but the peculiar inflorescence, the form and proportion 
of the glumes, &c. readily distinguish it. Grrisebach found only 
a single stamen in the flower, a character which I have no means 
of testing, the spikelets in our specimens having already lost 
their stamens. 

The Alopecuroid group of OryzesB consists of four genera : — 

9. Beckeea, Presen., two or three Abyssinian species, in some 



ME. a. BBNTHAM ON GEAMINEiB. 57 

respects intermediate between the two groups. The structure of 
the spikelets, with the two outer glumes very minute or deficient, 
connects them with the preceding genera; whilst the spicate 
inflorescence and three stamens are nearer those of Alopecurus, 
although the spikes are much more slender and several on the 
same stem, on long slender peduncles. The genus is confined to 
those of the § a of Steudel's ' Synopsis ;' the species arranged 
under § h have a very different structure, and form the section 
Becheropsis of Pennisetum. 

10. CsYPSis, Ait. (Antitraffus, Gsertn.), must be limited to the 
original C aculeata, which alone has the characters of the tribe. 
All the other species usually referred to it have the 2-nerved palea 
and other characters of the Agrostese, and were well separated by 
Host under the name of Heleochloa. It is true that some short- 
spiked varieties of Heleochloa scJuxnoides have very much of the 
aspect of 0. aculeata ; but besides the structure of the spikelets 
and the articulation of the rhachilla, they are readily distinguished 
by the rhachis of the spike, which is linear and cylindrical, not flat 
as in Crypsis. 

11. CoEifircopTiE,Linn., is a single Oriental species, very near 
Crypsis, but well characterized by the peculiar inflorescence and 
by the form of the fruiting spike and peduncle, which has supplied 
the generic name. 

12. AioPECiTETJS, Linn., including Cololachne, Beauv., and 
Tozzettia, Savi, is a well-known and perfectly definite European 
and temperate Asiatic genus, with the habit nearly of Phleum and 
the structure of the spikelets that of Oryzeae. Above forty sup- 
posed species have been enumerated ; but at least half of them 
must be regarded as trifiiug varieties of the two or three com- 
monest species, which have now, and perhaps from remote times, 
spread over a great part of the civilized world. 

Tribe IV. Teistegine^. 

This tribe, first proposed by Nees, has been fuUy adopted and 
much extended by Munro, and now consists of thirteen genera, 
which had been variously scattered in Pauicese, Andropogonese, 
and Agrostese, and are really more or less connected with the three 
tribes. They differ from Paniceee and approach Andropogonese in 
the thin, often hyaline texture of the fruiting glume and palea, and 
by the frequent presence of a slender, often bent awn on the 
flowering glume. From Andropogonese they are chiefly separated 



68 ME. G. BENTHA.M ON GEAMUfEJB. 

by their inflorescence; the spikelets are singly scattered or 
clustered along the inarticulate branches of the panicle or, in the 
very few cases where they are in pairs, the two of each pair are 
perfectly similar. Tristeginese are distinguished from Agrosteee 
by the characters which separate the two primary series PanicacesB 
and PoacesB. The tribual name was given by Nees from the genus 
Melinis, which he published as Tristegis, believing it to be new ; 
and although its identity with Beauvois's Melinis has since been 
established, it does not seem worth while now to alter the tribual 
name, which has been pretty generally adopted. Of the thirteen 
following genera, several of them common to the New and the 
Old World, the first four, with three glumes to each spikelet, are 
temperate or subtropical, the following nine all tropical, with four 
glumes to the spikelet. 

1. Thitebeeia is a new name I have been compelled to substitute 
for Qreenia of Nuttall or Scleraohne of Torrey, both of which had 
been preoccupied. The genus is limited to two North-American 
species which Steudel has proposed to unite with Limnas; but 
they differ essentially from it in the awn of the flowering glume 
terminal, not dorsal, in the distinct styles, and other characters 
besides habit. I have named the genus after G. Thurber, who has 
much studied North- American G-ramineae and worked them up 
for S. Watson's Californian Mora. The genus formerly dedi- 
cated to him by Asa Gray has since proved not to be distinct from 
Qossypium, to which it has been reunited by the author himself. 

2. Limnas is a single perfectly distinct species from East- 
Eussian Asia, well described and figured by Trinius. 

3. PoLTPOGON, Desf., a genus readily known by its dense in- 
florescence and the long awns of its empty glumes, is one of those 
which interferes in some measure with general classification. It 
has usually been placed in Agrostese ; but the very decided arti- 
culation of the pedicel removes it from that tribe to the Triste- 
ginese, where in many respects it is allied to Garnotia. It consists 
of about ten species, dispersed over the temperate regions both of 
the northern and the southern hemisphere, one of them almost 
cosmopolitan, but they are rare within the tropics. It was first 
published by Savi under the name of Santia in the Memoirs of the 
Italian Society of Science, a publication which had so little circula- 
tion that the name has not found its way into standard works, and 
that of Desfontaines has now been so long and so generally in use 
in all countries, that it would only create useless confusion now to 



MB. G. BENTHAM ON GBAMINE^. 59 

take up Savi's. Tte Mexican P. elongatus, H. B. K., which is 
Presl's genus Nowodworshya (first described and figured by him 
under the name of JRaspailia), has the pedicels, although claTate 
as in the rest of the genus, yet less decidedly articulate, thus 
forming some real connexion with the Agrostese. 

4f. GrAENOTiA, Brongn. {Miquelia, Nees, Berghausia, Eadl.), 
which sometimes comes near to some forms of Polypogon, has, on 
the other hand, the spikelets in pairs on the inarticulate branches 
of the panicle as in Miscanthus, and thus very closely connects 
Tristeginese with Andropogonese. It has, however, none of the 
long hairs on the rhachilla so common in Andropogonese, and 
cannot well be removed far from Arundinella, whilst Miscanthus 
is too near to Imperata to be rejected from Andropogonese. 
Garnotia comprises about eight species from East India, China, 
and Japan. 

5. ARUNDiNELiiAjEaddi, includes Ooldhachia,Tvm.,Acratherum, 
Link, Thysanachne, Presl, and Brandtia, Kunth. It is the prin- 
cipal genus of the tribe, and comprises about twenty-four species 
spread over the tropical regions both of the New and the Old 
World, but chiefly in Asia. It is generally adopted and fairly 
characterized, though the habit and especially the inflorescence 
vary much, the panicle being sometimes long, narrow, and dense, 
or very large, loose, and spreading, with very numerous small or 
minute spikelets, whilst in a few species it is short and dense, 
forming almost an oval head with larger spikelets: The two 
sections proposed by Nees — MeUosaccharitm, with a small tooth 
on the flowering glumo on each side of the awn, and AcratTierum, 
in which the glume is quite entire, tapering into the awn — do not 
prove to be well defined nor conformable to habit. A. Jiammida, 
Trin., from Brazil and tropical Africa, has neither the habit nor 
the character of the genus, but is in every respect a Trichopteryx, 
with which it was not compared by Nees, Trinius, or Doell, 
because it was at first only known as Brazilian, and TricJiopteryoc 
was supposed to be exclusively African. 

6. PH.aiNOSPEEMA, Munro, is a single Chinese species, nearly 
allied to Arundinella ; but there are three lodicules to the flower 
and no palea (unless one of the lodicules, although apparently in 
the same whorl as the others, be really a small palea), and the 
caryopsis is half exserted from the fruiting glumes as in some 
species of Sporoholus. Phanosperma glolosa, Munro, is a tall 
grass with a very large loose panicle, the slender but rigid 



60 ME. G. BENTHAM ON GEAMINE^. 

branches distantly verticillate along the main rhachis. It was 
first received from the Jardin des Plantes at Paris, where it had 
been raised from seeds brought from China by the Pere David; 
but it has since turned up among Shearer's Kiu-Kiaug plants. 

7. Melinis, Beauv. {Tristegis, Nees, Suardia, Schrank), is a 
single Brazilian species, Nees's original type of the tribe. It is 
very near Anmdinella, but remarkable for the long slender awn 
of the third empty glume, whilst the flowering glume is short, 
without any awn. Doell has reduced the genus to a section of 
Panicim, a view in which I can by no means concur. 

8. TniscENiA, Griseb., is a single Cuban species unknown to 
me, but from the author's description it must be very near to the 
following. 9. AETHEOPoaoif, Nees, a single Brazilian species, 
well described and figured by Kunth. So also is 10. Eetnaudia, 
Eunth, a single "West-Indian species allied to ArtJiropogon ; but 
the awn, longest on the lowest glume, is gradually shortened and 
reduced to a point on the flowering one, and there is no palea: 
there are, however, four lodicules, a condition so unusual in 
Graminese, that we might be tempted to consider the lowest pair 
of lodicules, though close upon the others, as being in fact a 
bipartite palea. 

11. Bhtncheltteum, Hochst., two or three tropical African 
species, which appear to form a fairly distinct genus allied to Arun- 
dinella, but approaching nearer to the A ndropogonese in the long 
hairs of the lower glumes. The generic name was originally Nees's, 
who applied it to a South- African plant of Drege's, which proves to 
be scarcely even a variety of the Panicum {Tricholcena) roseum of 
that country. Hochstetter and Steudel totally misunderstood 
Nees's genus when they added to it their B. grandiflorum, and B. 
ruficomum, which may now, however, retain those names, Nees's 
genus being suppressed. 

12. Thts ANOE^NA, Nees {Myriachata, ZoU. and Mor.), is a single 
tropical Asiatic species, a very tall grass with long broad leaves 
and a very large full panicle, with innumerable minute spikelets 
in dense clusters along its long crowded branches. The flowering 
glumes are more or less covered with rather long hairs ; but these 
hairs are so closely appressed and covered by the empty glumes 
that Steudel could not see them, and published a supposed second 
species as being destitute of them. Trinius figured the plant as a 
Panicum ; by other early Indian botanists it was referred to 
Agroitis. 



ME. a. BENTHAM ON GHAMINE^. 61 

13. CiBiSTAOHNE, 13 a genus I have proposed for two plants, 
one from East India, the other from tropical Atrica, which have 
something of the aspect of Sorghum tropicum ; but the spikelets 
all hermaphrodite, and never in pairs, remove them from the 
Andropogoneae to the TristegineEe. I purpose figuring the genus 
in the forthcoming part of Hooker's loones. 

Tribe V. Zotsiej3. 

I have composed this tribe of two groups or subtribes, which 
might perhaps have been regarded as separate tribes, although the 
difierence between the two is only that which lies between the 
Cenchrus group and Panicese proper. In the first group, Anthe- 
phorecB, the spikelets disarticulate from the rhachia of the inflo- 
rescence or from the pedicels in little clusters of two to six, or 
very rarely more ; in the other group, or Zoysiece proper, the 
spikelets are solitary, or very rarely two together on the pedicels. 
In both groups the structure of the spikelets is generally that of 
Andropogonese, sometimes slightly approaching that of Paniceae, 
but the pedicels are singly scattered or alternate along the inar- 
ticulate rhachis of the spike or general inflorescence. The An- 
thephoresB have hitherto been usually placed in Panicese, as having 
nearly the inflorescence of Genehrus, but of which they have 
not the hardened inner fruiting glume ; the Zoysiese proper have 
mostly been considered as Andropogonese, from which they dififer 
in inflorescence. Of the twelve following genera, the first six 
belong to Anthephoreaj, the remaining six to Euzoysiese or 
Zoysieae proper. 

1. HiLAEiA, H. B. K., in which I should include Pleuraplds of 
Torrey, and, judging from the figure and description, Sexarrhena 
of Pros], comprises five or six species dispersed over the Mexicano- 
Texan region, extending into California. Although the forms 
and proportions of the glumes of each spikelet vary much in the 
difiierent species, or even in difierent spikes of the same plant, the 
genus as a whole is a natural one, and readily recognized by 
each cluster consisting of three spikelets, the central one con- 
taining a single fertile flower, either female or hermaphrodite, the 
two lateral ones each with two male flowers. The spikelets are 
often so closely sessile in the cluster, that it requires some care 
to ascertain which glumes belong to each cluster, and the pairs 
of male triandrous flowers of the lateral spikelets have sometimes 
been described as single hexandrous flowers. The species I have 

LINN. JOrEN. — BOTANT, VOL. IIX. O 



62 ME. Q. BENTHAM ON GEAMINE^. 

seen are H. cenchroides, H. B. K., H. Jamesii (PleurapMs Jamesii, 
Torr.), S. mutica (FleurapMs mutica, Buckl.), -2". serieea (Pleu- 
rapMs serieea, Nutt.), and a West-Texan species {WrigM n. 758 
and 2109, Berlandier n. 168, 1428) very near H. cenchroides, but 
apparently distinct. 

2. iEaopoGOS", Humb. and Bonpl. {HymenotJiecium, Lag., Schel' 
lingia, Steud.), extends in two species from Bolivia to Mexico. 
The genus bas at first sigbt much the aspect of the Asiatic 
MelanoeencTirus, or of some of the very short-spiked species of 
Bouteloua, but the real afiinity appears to be with Silaria. The 
spikelets usually vary from two to six in the cluster, mostly 
with one hermaphrodite flower in each, though there are usually 
one or two empty barren spikelets intermixed ; the clusters are 
in a loose one-sided spike, each one very readily disarticulating 
from its very short pedicel. 

3. Cathestechus of Presl, a single Mexican species, is only 
known to me from his figure and description, which do not agree 
with each other in some important particulars. He says that the 
genus is allied to Mgopogon. I have no means of judging whether 
that be really the case. 

4. Anthephoea, Schreb., is a very well-known and perfectly 
characterized genus of five or six species, of which one is tropical 
American, the others tropical or Southern African. SypudtBurus, 
Hochst., quoted by A. Braun in ' Mora,' 1841, p. 275, and by 
some others, is AnthepJiora abissynioa, Steud. 

5. Teachts, Pers., is a single well-known species from the East- 
Indian peninsula, several times figured by the earlier botanists of 
this century. It is slightly anomalous in the tribe by its spikes 
being two together at the apex of the peduncle, and, as in An- 
ihepTiora, the excessive hardness of the clusters of spikelets after 
flowering renders it difiicult to trace their structure unless exa- 
mined young. The name Trachys was changed by Eeichenbach 
to Trachyogus, and by Dietrich to Trachystachys, as having been 
preoccupied by zoologists, a plea not now regarded as sufficient. 

6. Tbagtjs, Hall. (Lappago, Schreb.), is a single annual very 
well known as a common weed in tropical and temperate regions 
almost all over the civilized world. 

7. I.ATiPEs, Kunth, is a single tropical- African annual, extend- 
ing eastward as far as Scinde, very well described and figured by 
Kunth. It has been united by others with Tragus; but the 
small spikelets, usually solitary or rarely two together on Iho 



ME. G. BBNTHAM ON GEAMINE^. 63 

pedicel, and the very different shape and proportion of the glumes, 
seem sufficient to maintain the genus as distinct. 

8. LoPHOLEPis, Dene., is a little slender East-Indian annual, 
allied in some respects to Latipes, hut with excessively minute 
curiously shaped spikelets, so rapidly ripening and so very deci- 
duous that it is very rare to find any on the specimens in an 
examinable state. The plant was first sent home by "Wallich 
under the name of Solhoellia, and was figured as such by Hooker 
in the Botanical Magazine ; but in the meantime "Wallich pub- 
lished a Lardizabalous genus under that name in his Tentamen 
of a Nepal Flora, and Decaisne therefore changed that of the 
present grass to LopJiolepis. 

9. Netikachne, R. Br., three Australian species, and 10. Peko- 
Tis, Ait. {Xystidium, Trin.), from the tropical regions of the Old 
World, of which the species are variously estimated as from two to 
seven, are both of them well-known genera, accurately described 
and figured. 

11. Leptotheium, Kunth, founded on a specimen brought by 
Humboldt from tropical America, is unknown to me. It is said 
to be very near the Asiatic genus Zoysia, and, from the descrip- 
tion, seems to differ chiefly in its widely distant geographical 
station and in the presence of an additional lower empty glume. 

12. ZoTSiA, Willd. {Matrella, Pers.), is a well-defined genus of 
two or three maritime plants, dispersed over the shores of eastern 
and southern Asia, Australia, and New Zealand, extending also to 
the Mascarene Islands. 

To these Zoysieee I have provisionally added a small Mexican 
plant, the affinities of which are very puzzling, and which I 
have described and figured as a new genus ScHAirifEBA, so named 
after the collector from whom we have received it. At first sight 
it seemed to bear some resemblance to Presl's figure of Gatheste- 
chus ; but the structure of the spikelets is quite different, being 
nearly that of Zoysia, whilst the general inflorescence, though on 
a much smaller scale, approaches that of some species of Andro- 
pogon {Cymbopogon) or of Apluda. 

Tribe VI. AifDBOPOQONEiE. 

This tribe is chiefly characterized by the spikelets in pairs at 
each node of the articulate rhachis of the spike or of the branches 
of the panicle, or in triplets at the end of each branch, and by 
the inner glume under the fertile flower being much smaller and 

g2 



64 MB. G. BENTHAM ON GEAMINEJ!. 

thinner than the lower or outer empty ones, usually hyaline, aad 
often bearing a twisted or bent awn. The two spikelets of each 
pair are either both of them perfect and fertile, or one of them 
is male only or imperfect, or even quite rudimentary, and the 
spikelets are often more or less surrounded by long silky hairs. 
But to each of these characters there are exceptions in single 
genera, which are retained in Andropogonese as agreeing with 
them in most other respects. 

The plants of this tribe are for the most part tropical or sub- 
tropical, although a few are found in more temperate regions, 
chieily in the northern hemisphere. More than eighty genera 
have at different times been proposed, which some botanists would 
reduce to below twenty. Following as nearly as possible the 
principles we have hitherto adopted, I have thought that the 
following twenty-six may be admitted as fairly characterized, 
referring them to four subordinate groups or subtribes — Sac- 
ckarem, Arthraxece, Sotthoelliece, and Andropogonece proper. 

SaccTiareoB comprise seven genera, in which the two spikelets 
of each pair are homogamous, both of them hermaphrodite and 
usually fertile, and the inflorescence paniculate, excepting Pogor 
natherum. 

1. Impeeata, Cyr., three or four species widely spread over the 
tropical and subtropical regions both of the New and the Old 
World, extending northwards to South Europe, China, and Japan. 
In this and the following, Miscanthus, the branches of the panicle 
are exceptionally inarticulate, showing an approach to the Tri- 
steginete ; but the long silky hairs and the very much reduced 
hyaline flowering glume and palea retain them in Andropogonese. 
Munro has shown that the common American I. caudata, Anders., 
is identical with the Old-World 1. ramosa, Anders. ; and I also 
can find no difference between the two, any more than between 
the American and the Old- World specimens of I. artmdinacea. 
Pournier has, however, proposed to separate the American forms 
of the two species generically under the name of Syllepis, on the 
plea of their having the two lodicules connate into a single large 
truncate one, which I have in vain sought for in several difierent 
American specimens. It is possible that Fournier may have 
considered the small truncate palea as a pair of united lodicules, 
but, if so, they are precisely the same in the Old- World species. 

2. MiscAfTTHTJS, Anders,, as now limited, is a genus of eight 
species, of which one is South- African, the others dispersed over 



MB. fl. BENXHAM ON GKAMINEJE. 65 

Eastern Asia from the Malayan archipelago to Japan. It has 
the inarticulate panicle-branches and most other characters of 
Imperata, from which Andersson technically separated it by the 
awn of the flowering glume. Exceptional unawned species occur 
in so many genera where they are usually awned, that this can 
scarcely be regarded as a generic character where there is nothing 
else to separate the two forms. Here, however, if we remove one 
species from Imperata to Miseanthus, inflorescence supplies two 
natural groups. In Imperata the panicle is long, narrow, and 
dense, with short erect branches buried in the copious silky hairs, 
the glumes are never awned, and there is only one, or rarely two, 
stamens ; in Miscanthus the panicle is loose, with long spreading 
branches, the silky hairs are less dense and in one species almost 
wanting, the flowering glume is in most species awned, and there 
are always three stamens. The species known to Trinius were 
by him included in JEulalia ; and Munro, whom I followed in 
the ' Flora Hongkongensis,' restricted the name Eulalia to the 
species now constituting Miscanthus ; but as the true JEulalia of 
Kunth is the type of a section of the very difierent genus Pol- 
linia, I have thought it necessary to adopt Andersson's later name 
Miscanthus. Besides his species, I would include in the genus 
M.fuscus {ISriochrysis fusca, Trin.,^. attenuata, ITees) from East 
India, M. saccharifer {Imperata saccharifera, Anders.), from North 
China, which has the inflorescence and stamens, but not the awns, 
of the other species, and M. cotulifera (Uulalia cotulifera, Munro) 
from Japan, which has scarcely any of the hairs of the other species. 
Steudel proposed the latter as a distinct genus under the name 
of Eocoilopus. 

With 3. SAOCHAErM and 4. EniANTHrs commence the series 
of true AndropogonesB with the branches of the panicle arti- 
culate ; and these two genera are so closely connected that they 
might well be reunited, although they are now almost universally 
recognized as distinct. There might indeed be no great objec- 
tion to consider both, as well as Pollinia and Spodiopogon, as sec- 
tions of one large genus. As now limited, Saccharum is chiefly 
characterized by the compound panicle, usually dense, sometimes 
very large, and the spikelets very small without any points or 
awns to the glumes. The species are supposed to be about ten, 
the typical ones belonging to the tropical or subtropical regions of 
the Old "World, amongst which the well-known sugar-cane is now 
extensively cultivated also in America. The genus would also 



66 ME. G. BENTHAM ON GEAMINE^a). 

include £'. Nareya [Mriochrysis Wareya, Nees) &nA B.longifolia, 
Munro {Uriochrysis longifolia, Munro), from East India, S. pallida 
{Uriochrysis pallida, Munro) from South Africa, and 8. cayennense, 
the typical Uriochrysis of BeauYois, which last differs only iu the 
very dense almost spike-like panicle. 

4. Ebianthus, Mich. (Bipidium, Trin.), would be a more satis- 
factory genus if it could be restricted to the two old species 
U. saccharoides, Mich., from North America, and E. Bavenne, 
Beauv., from the Old World ; but besides the above-mentioned 
connexion with Saccharum, there are several South-American 
species which run very closely into Pollinia. On the whole, it 
seems best to consider Ilrianihus as an intermediate genus 
between SaccTiarwn and Pollinia, having the inflorescence of the 
former, but the flowering-glume more developed into a point or 
awn almost as in Follinia. It would then consist of about twelve 
species, amongst which JE. striata, Nees, from North America, has 
no hairs on the rhachiila, but only a shorb pubescence on the 
glumes. 

5. Spodiopogok, Trin., differs from Pollinia, as Ohrysopogon 
does from Andropogon, chiefly in inflorescence. The short 
branches of the panicle bear three spikelets, one sessile between 
two pedicellate, and occasionally there is a pair of spikelets below 
the three terminal ones ; but the branches never form the regular 
spikes of Pollinia. Besides the original S. sihiricus, Trin., we 
have two additional species, S. pogonantlius, Boiss., from the 
Levant, and & albidiis {Andropogon albidus. Wall. Cat. Herb. Ind. 
n. 8821), from East India. The generic name has also been often 
misapplied. S. angmtifolius, Trin., is a Pollinia ; some others of 
his species with 2-flowered spikelets belong to Ischmmum. Four- 
nier's Mexican Spodiopogons are evidently species of Erianthus ; 
his 8. foliata indeed (Bourgeau, n. 2979) appears to me to be 
identical with the original E. saccharoides, Mich. 

6. Pollinia, Trin., is now a genus of about twenty-five tropical 
or subtropical Old- World species, with the inflorescence of the sec- 
tion Oymnandropogon of Andropogon, and the homogamous spike- 
lets of Saccharum and Erianthus ; the spikelets are in pairs along 
the simple branches of the panicle ; these branches either few, 
almost digitate at the end of the peduncle, or more numerous and 
scattered along the main rhachis. The genus is divisible into 
two very naturarsections : — 1. Eulalia, with the spikes and pedicels 
covered with long silky or rufous hairs as in Erianthus, includes 



ME. a. BENTHAM ON GRAMINEJS. 67 

P. aurea, Bentli. (the original genus JSulalia, Kunth), P. articu- 
lata, Trin. {Pogonatherum contortum, Brongn.), P. eriopoda, Hance 
{Spodiopogon angustif alius, Trin.), P. longisetus {EriantJms lon- 
gisetus, Anders.), P. versicolor {Erianthus versicolor , Nees), P.fili- 
folius (Urianthus filifolius, Nees), and a few others. 2. Lepta- 
therum, with slender spike-like branches, of which the hairs are 
few or short, so as to appear sometimes quite glabrous ; this 
section includes P. glabrata, Trin. (Hulalia glabrata, Brongn.), 
P. nuda, Trin. (P. imberlis, Nees), P. Willdenowianuni (the genus 
Microstegium, Nees, P. lancea, Nees, published also by Nees as 
his genus Leptatherum, and probably also Steudel's Ifemastachys). 
Sprengel's Pollinia would have had the right of priority over 
Tririius's ; but that proved a farrago made up of a few heteroge- 
nous species of Andropogon, Ghrysopogon, and Pollinia. 

7. PoGONATHEETTM, Beauv. {HomopUtis, Trin.), is a single tro- 
pical and subtropical Asiatic species, very well marked by its 
slender, much branched habit, the single spikes, and the slender 
awns arising as well from the second empty glume aa from the 
flowering one. 

The ArthraxecB, or second group of Andropogonese, consist of 
three genera, which have the inflorescence of Pollinia ; but the 
second spikelet of each pair is generally reduced to a bare stipes, 
or is even quite deficient, bringing a few species very near to the 
Zoysiese, differing chiefly in their subdigitate spikes, whilst a few 
others, in which the spikes are single, have the rudiment of the 
second spikelet of true Andropogonese. 

8. Apocopis, Nees (Amilyachyrum, Hochst.), has iive or six 
species from East India or the Malayan archipelago, characterized 
by the very broad truncate outer glume enclosing the rest of the 
spikelet. Among the species A. Boyleanus, IN'ees {IsoJimmtim 
paleacewm, Trin., Andropogon paleaceum and A. liimalayensis, 
Steud.), is remarkable for the awn often (but not always) reduced 
to a small fine point, or even entirely wanting ; and A, tridewtata 
{Andropogon tridentatus, Eoyle) has, on the contrary, a very long 
awn, and the young spikes are usually enclosed in a large spathe- 
like bract. 

9. DiMEEiA, E. Br. (ITaplachne, Presl, Didaciylon, Zoll, and 
Mor., PsilostacJiys, Steud., PterygostacTiyv/m, Nees), about ten 
species from the Indo-Australian region, has very slender spikes, 
the lower empty glumes very narrow and rather rigid, and usually, 
if not always, only two stamens. 



68 ME. G. BENTHAM ON GEAMINB^. 

10. Abtheaxon, Beauv. {Pleuroplitis, Trin., Batratherum, 
Nees, Lucwa, Kunth, Lasiolytrum, Steud., Alectoridia, A. Eich., 
Psilopogon, Hochst.), has also about ten species, chiefly from 
the Indo-Australian region, but extending on the one hand to 
China and Japan, and on the other to tropical Africa. The 
spikes are slender as in Dimeria ; but there are three stamens, 
and the lower empty glume is broad but acute, not truncate as in 
Apoeopis. 

Bottloelliea, the third group of Andropogonese, is often re- 
garded as a distinct tribe, characterized by the simple spike, 
the apikelets in pairs at each notch or excavation of the rhachis, 
the one sessile, the other pedicellate, and no awn to the flowering- 
glume. There are, however, as in other subtribes, here -and 
there exceptions to one or more of these characters. "We have 
seven genera. 

11. EtiioinjETJS, Humb. and Bonpl., has about twelve species, 
chiefly South-American or African, with, however, one Austra- 
lian and one from the East-Mediterranean region. They all 
differ little from Rottboellia besides the long silky hairs which 
clothe the spike, thus connecting EottboelliesB with other An- 
dropogonese. -B. Jnrsuta, Munro (SottboelUa Jiirsuta, Vahl), has 
been proposed by Boissier as a distinct genus Lasiurus, as having 
the spikelets in threes instead of in twos at each node of the 
rhachis. But that character is by no means constant ; in several 
specimens I have found the spikelets in threes or even in fours 
at the lower nodes ; but in others they are in the normal pairs 
from the base of the spike. 

12. SoTTBOELLiA, Linn, f., a tropical or subtropical genus widely 
spread, but chiefly in the Old World, has been either extended to 
nearly the whole subtribe or very variously restricted to a small 
number or to a single species. It seems best characterized by in- 
cluding all those which have the simple terete spike, without the 
hairs oiHUonurus or the peculiarities of the four following genera. 
It would contain about eighteen species, amongst which several have 
been proposed as monotypic genera. Coelor}iaoMs,'Biongn.,iB M. mu- 
ricata,'Retz (JR. glandulosa, Trin.); PeZ^qpAorw«,Desv.,is It.myurus: 
in both of these the lowest or outer glume of the perfect spikelet is 
rigid and bordered on each side at the apex by a membranous wing, 
which, however, is also present, but much less prominent, in M. 
rugosa, Nutt. Fhacelur a, (jrvmeb. (Pholiurus,'Svin. in Spreng. JN^eue 
Entd. ii. 67, not of the Eundam. Agrost.), is the Oriental B. digi- 



ME. G. BENTHAM ON &BAMINEJ3. 69 

tata, Sibth. {B. Sandorii, Friwaldsk.), a species striking for tte 
long spikes, occasionally thougb very rarely branched at the base, 
and from the rather large spikelets with acuminate outer glumes 
showing an approach to some Vossice, but scarcely sufficiently 
distinct from Botthoellia to be kept up as an independent genus. 
Gymbaolvne, Eetz, a Bengal grass, has been referred by Willdenow 
to JRotiboellia. Eetz's character does not quite agree ; but the 
plant has not since been identified, and must remain doubtful. 
Apogonia, Pourn., comprises two Mexican species which I am 
unable to distinguish from JRotiboellia : Nuttall's section Apogonia 
of Rottloellia is a species of Ulionurus, very closely allied to, if 
not a variety of, E. ciliaris, H. B. K. 

13. OpHirEra. This genus, as first proposed by Gsertner, in- 
eluded two very different plants separated by Brown as Leptttrus 
and OpMurus. As the latter is now limited, it diifers from Sotf- 
hoellia only in the absence of the second sterile spikelet of each 
node, at least in the upper part of the spike or inflorescence. It 
consists of three, or perhaps four, Asiatic, African, or Australian 
species : — O. corymhosa, GraBrtn. (O. cethiopica, Steud.), O. mono- 
stachya, Presl (0. undulata, Nees), and O. Icevis (Rotiboellia Icevis, 
Retz, R. perforata, Roxb.). The latter species is remarkable for 
having the spikelets in the lower part of the inflorescence in pairs 
at each node as in Hotthoellia, but the two of each pair separated 
by a kind of partition dividing the cavity of the rhachis into two ; 
it has therefore been raised to a genus by Kunth as Mnesithea 
and by Nees as Thyridostachyum. G-enerally, however, in the 
upper part, and sometimes in the whole inflorescence, the sterile 
spikelet is wanting, as in Ophiurus, especially in the young 
spike, for the upper or Ophiurus portion appears to fall away 
very readily, leaving only the Mnesithea part persistent. Lepturus, 
Br., is now classed in the tribe Hordeese. 

14, Eatzebtteqia, Kunth, a very elegant little flat-spiked 
Burmese grass, and 15. Manistjeis, Linn., a common tropical 
weed with little globular spikelets, have both been well described 
and figured. 

16. Hemaetheia, Br., contains two or three tropical weeds or 
maritime grasses, separated from Rotthoellia chiefly on account of 
the flattened and less distinctly articulated rhachis of the spike, 
and the curious way in which the stipes of the sterile spikelet is 
adnate to the rhachis, so as to make it appear sessile and almost 
opposite to a fertile spikelet, which really belongs to the next 



70 ME. Q. BBNTHAM OS GEAMIHE^. 

superior node. These characters, though generally well marked, 
are sometimes more or less obscure. 

17. VossiA, Wall, and Griff., closely connects the EottboeUieae 
with Ischcsmum. As in the former, the flowering glume is always 
unawned, and the rhachis of the spike is rigid and deeply notched, 
but the lower empty glume, at least of the pedicellate spikelet, is 
produced into a long point or awn ; there are generally several 
spikes or simple branches along the common peduncle, and there 
is in each sessile spikelet a male flower below the terminal fertile 
one, as in IscTicemum. The genus was originally established on a 
handsome semiaquatic East-Indian grass, which has since been 
found also in tropical Africa, and two or three additional species 
have reached us from the same country. "We should also refer to 
Vossia the Ischamum speciosum of Nees from East India. Hremo- 
chloa, a Japanese plant described by Biise, is unknown to me ; but 
the character given, if I correctly understand it, agrees well with 
that of Vossia. 

To the fourth group, or subtribe UuandropoffoneiS, may be 
referred nine genera, in which the two spikelets of each pair are 
heterogamous and the flowering glume of the fertile one is more 
or less awned ; and in the first five the spikelets are in many pairs 
along the rhachis of the simple spikes or panicle-branches. These 
nine genera are increased to twenty-one by Andersson and others, 
whilst Steudel unites seven out of the nine under his Andropogon. 

18. Theleposon, Eoth (Jar&'«iffi, Steud.), comprises one East- 
Indian and two or three tropical- African species, all very elegant 
and closely resembling each other. Their infiorescence is that of 
Vossia, whilst the spikelets are nearer those of Ischcemiim, but re- 
markable for the rigid tuberculate outer empty glumes. Nees, in 
working up Wight and Amott's Peninsular grasses, gave Eoth's 
name to a very different grass (Ischcemum semisagittatum, E.oxb.), 
adding the observation that Roth's "description is very bad. The 
fact is, however, that it is Nees who was mistaken in his identi- 
fication, whilst Eoth's description of the true plant is excellent. 

19. IscH^MXTM, Linn., as now understood, has about thirty 
species, widely dispersed over the warmer regions both of the New 
and the Old World, the chief character connecting them being 
that the sessile spikelets have a male flower below the terminal 
fertile one. The spikes are also usually stouter than in Andro- 
pogon, and the genus is a fairly natural one. Beauvois restricted it 
to the I. mwticim, Linn., in which the awn of the flowering glume 



ME. G. BENTHAM ON GEAMINEiE. 71 

is small and hair-like or sometimes entirely wanting, and proposed 
a genus Meoschmm, adopted byNees, for the other species in which 
the awn is more developed. Trinius considered as true Isolimma 
only those in which the pedicellate spikelet has only a male flower 
or empty glume, and added those in which that spikelet has two 
male flowers to his genus Spodiopogon, notwithstanding the dif- 
ference in inflorescence &c. Ischcemopogon, Griseb., is I. latifolmm, 
Kunth, and Hologamium, Nees, is I. laxvm, Br., both species 
with two-flowered pedicellate spikelets, as is also the case in I. 
insculptum, Hochst., and I. maerostaehyum, A. Kich., from tropical 
Africa, and probably also in Porsklhl's genus Sehima, of which 
we have no authentic specimen. I. pectinatim, Trin., I. leersioides, 
Munro, I. ophiuroides, Munro, with a fourth unpublished species, 
all from tropical Asia, form a distinct section {Pectinaria), with 
slender elegant simple spikes, and the larger glume of the sessile 
spikelets pectinate-ciliate. 

20. TEACHTPOGtON, Nees, as limited by Andersson, and 21. 
Heteeopogof, Pers., closely resemble each other in their simple 
spikes with appressed imbricate spikelets and long rigid twisted 
awns ; but in Trachypogon the sessile spikelet of each pair is male 
or sterile and unawned, and the pedicellate one fertile and awned, 
whilst in Seteropogon the sessile one is fertile and awned, and 
the pedicellate one male or sterile and unawned. Andersson enu- 
merates eleven species of Trachypogon, one from South Africa, the 
others from tropical or subtropical America; but several of the latter 
can scarcely be regarded as more than slight varieties. Of Setero- 
pogon there are two well-marked species, H. contortus, Eoem. and 
Schulfc. (S. hirtus, Pers.), now very common in most warm regions 
and extending to the Mediterranean region and to North America, 
and H. melanocarpus, Ell. {H. Moylei, Nees, H. acuminatus, Trin., 
Trachypogon scroMculatus, Nees), which is in North and South 
America as well as in East India. Besides these, three or four 
South- African species have been referred to Seteropogon, but are 
of somewhat doubtful affinity. 

22. Andeopogon, Linn., taking it within the limits assigned to 
it by Munro, including all the species of the subtribe with spike- 
like simple branches to the inflorescence, and without the pecu- 
liarities of the three preceding genera, is still a somewhat poly- 
morphous genus of perhaps a hundred species, very abundant 
within the tropics, but well represented also in Europe, temperate 
Asia, North America, South Africa, and Australia. The fourteen 



72 ME. G. BENTHAM ON GBAMINE^. 

genera into which it has been divided may be fairly reduced to 
the following five, perhaps too artificial, sections : — 1. Schizachy- 
rium, about a dozen species, with the spikes always single upon 
each peduncle. The genus Sohizaohyrium, Nees, was limited to 
a few species in which the spike is slender and not very hairy. 
Dieetomis, H. B. K., is the American A. fastigiatus, Sw., found 
also in tropical Africa, which has a more rigid spike and the 
second empty glume conspicuously awned. HomoeatJierum, Nees, 
is an Asiatic species scarcely to be distinguished from the same 
A. fastigiatus. In 2. Oymhopogon, the spikes, often very silky-hairy 
or woolly, are in pairs on each peduncle, and the peduncle partly 
or wholly enclosed in the sheath of a leafy or spathe-like bract. 
The species are numerous, chiefly in the Old World, and include 
the lemon-grass and its allies. Andersson has divided the section 
into two genera, Qymnanihelia and ISyparrhenia, and perhaps 
more ; but as he has never published their characters, I am unable 
to form any clear idea of them. It would appear, however, from 
the species quoted, that A. scJicenanthus and its allies would be- 
long to Gh/mnanthelia, and A. Mrtus and its allies to Hyparrhenia. 
3. Oymnandropogon, has two or more spikes sessile at the end of the 
peduncle, without any sheathing-bract. The species are nearly as 
numerous as those of Gymbopogon. Amongst them, A. annulatus, 
Porsk., though closely allied to the common A. Ischcemum, forms 
the proposed genus DicliantMum, Willem. ; A. serratus, Eetz, 
with a broad herbaceous outer glume, is Trinius's genus Lepeo- 
cercis ; and it is most probable that Steudel's HuJclastaxon is the 
common American A. virginicus. 4. Amphilophis, Trin., would 
include A. laguroides, DC, and A. argenteus, DC, from tropical 
America, with A. scandens, Eoxb., and A. Vachellii, Nees, from 
tropical Asia, and a few others, difflering from Oymnandropogon in 
the more numerous, usually long and often pedicellate spikes, 
sometimes even divided at the base, forming almost a saccharoid 
panicle. 5. Vetiveria, Thou. {Mandelorna, Steud.), is the well- 
known Vitiver, A. muricata, Eetz, to w hich Munro would redu e 
as varieties A. nigritana, Benth., and Vetiveria arundinacea, 
Q-riseb., a species frequent in East India and tropical Africa and in- 
troduced into America, distinguished by its numerous spikes verti- 
cillate along the axis of a long simple panicle, all glabrous or only 
minutely hairy, and the awn of the flowering glume often very 
much reduced. Beauvois's genus Anatherum, sometimes supposed 
to be specially destined for this plant, included also all the species 



MB. G. BENTHAM OK GEAMINBJD. 73 

of the sections Gymlopogon and Gymnandropogon, in wHcli the 
awn is much reduced or obsolete. Agenimn, Nees, from his cha- 
racter, would also refer to one of these species without prominent 
awns. 

23. Chbtsopogon, Trin. {B}iapMs,Lour., Centrophorum, Trin.), 
and 24. Sobghtjm, Pers. (SlumenhacMa, Koel.), are two genera 
very nearly allied to each other and diii'ering from Andropogon, as 
Spodiopogon does from Pollinia, chiefly in their inflorescence; 
the branches of the panicle bear three spikelets at the end, a 
sessile one between two pedicellate ones, and occasionally only 
one or two pairs below on the same branch. They were both 
included by Linnseus, and afterwards by Brown, in Holcus, a 
name since restricted to that portion of the old genus which 
belongs to Avenacese. Ghrysopogon, as now constituted, has nearly 
twenty species, chiefly tropical or subtropical, but including also 
the European G. Gryllus and some other temperate species. The 
genus may be divided into two natural sections : in the typical 
form the pedicellate spikelets usually contain a male flower ; in 
the section Siipoides, exclusively American, it is reduced to a long 
hairy stipes rarely bearing a minute rudimentary glume. This 
section includes C nutans, G. avenaceus, G. stipoides, G. Minarum, 
and a few others. Sorghum differs from Ghrysopogon in habit, 
in the scarcely articulate branches of the panicle, and iu the 
glumes of the fertile spikelets more hardened after flowering. 
The number of species is very uncertain, for, of the two prin- 
cipal ones, S. halepense is so widely spread as a tropical or sub- 
tropical weed, and S. vulgare so long and so generally cultivated 
in warm regions for a variety of purposes, as to have produced 
a great variety of forms, raised by many to the rank of species. 

25. Anthistieia, Linn. fil. (JPhemeda, Forsk.), if taken as a 
whole, is a very natural genus, of about a dozen species from 
the warmer regions of the Old World, easily recognized by its 
inflorescence. The spikelets are in short dense spikes or clusters, 
usually seven together, of which the four lower ones (two pairs) 
are either empty or with a male flower in each, and are placed 
apparently in a whorl, forming a kind of involucre round the 
three inner ones, which, as in Ghrysopogon, are one sessile between 
two pedicellate ones. In a few species the number of spikelets 
is raised to nine, or even to eleven, by the intervention of one or 
even two pairs of spikelets between the involucral and the ter- 
minal ones. These slight differences in the number or in the 



74 ME. G. BETTTHAM ON GEAMINEiE. 

pedicellation of tte spikelets hare induced the proposal of distinct 
genera for most of the species, and several of them have been 
adopted by Andersson in a monograph most carefully worked up 
in as far as the materials at his command admitted, but in which, 
for want of access to a sufiBciently rich library, he is much mis- 
taken as to several of the synonyms quoted. These proposed 
genera are : — 1. Aristaria, Jungh., for A. frondosa, Br. {A. Jung- 
huhniana, Nees), which forms the section Heterelytron of Anders- 
son, but not Junghuhn's genus of that name. 2. Perohachne, 
Presl, is A. arundinacea, Eoxb., forming Andersson's subsection 
ChrysantTiistiria. 3. Andersson's subsection -BMaM^Ajs^mfl for the 
common A. ciliata, Linn., and its immediate allies, to which some 
botanists would restrict the genus. Andersson distinguishes twelve 
species, adding at the same time that they might well all be re- 
duced to varieties of a single widely-spread species. 4. Andro- 
scepia, Brongn. {Heterelytron, Jungh.), was founded originally on 
the A. gigantea, Cav., but became a very unnatural group when 
made to include A. {Androscepia) anathera, Anders., which very 
closely resembles A. {Euanthistiria) minuta, Anders., and a variety 
arinata, Anders., of A. gigantea, which is much nearer to the A. 
(FeroiacJine) arundinacea. 5. Iseilema, Anders., containing two 
East-Indian and one Australian species, and 6. Exofheca, Anders., 
comprising A. abyssinica, Hochst., from tropical Africa, and A. 
fasciculata, Thw., from Ceylon, have each a peculiar habit and 
characters, sufficient to maintain them as sections. 7. Germainia, 
Balansa, has, perhaps, two closely allied species — A. caudata, Nees, 
from Ehasiya and China, and the typical A. capitata fron Saigou; 
the latter, however, which I only know from Balansa's figure and 
description, is exactly like the Chinese plant, except that there 
appear to be rather more spikelets in the cluster. 

26. Apltjda, Linn., is now universally recognized as a distinct 
and natural genus, limited to the two tropical-Asiatic species ori- 
ginally assigned to it by Linnseiis, though his character was even 
then very imperfect, and rendered still more so by the subsequent 
addition of the very different American Zeugites, which Schreber 
afterwards restored as an independent genus. Beauvois, however, 
threw every thing into confusion; for it is evident from his figures 
that his Diectomis is A. aristata, Linn., and his Galamina is A, 
mutica, Linn,, though in drawing up his character for the latter 
he combined it with some species of Anthistiria. Beauvois's 
Apluda is certainly different, probably a Chrysopogon, 



ME. G. BENTHAM ON GEAMIIfEJE. 75 



Series B. POACE^. 



Having already explained the difference between the two 
primary divisions of Graminese, I need only repeat here that 
the main characters of Poaceee consist, firstly, in the want of any 
articulation of the pedicel below the lower empty glumes, which 
remain persistent after the fruiting one has fallen away, or fall 
away separately, and, secondly, in the male or imperfect or rudi- 
mentary flowers, when present, being above, not below, the fertile 
one. The former character is all but universal; but from the 
latter one exceptions are not very rare, besides that, where there 
is only one flower without any continuation of the rhachilla 
beyond it, the character entirely fails. I should add that in 
some tribes of Poacese there are two or more perfect flowers in the 
spikelet, which is not the case in Panicacese ; and may now pro- 
ceed to examine in detail the eight tribes into which this second 
series may be divided. 

Tribe VII. PHAiiEiDEiE. 

The close aflBnity of this tribe and the Oryzeje has been 
generally admitted, and the two are usually placed in juxtapo- 
sition ; I had even proposed their consolidation into a single one 
in the 'Plora Australiensis.' They have in common the im- 
portant character of the scale immediately under the single 
perfect terminal flower being- keeled or one-nerved, so as to make 
it a matter of discussion whether it be a glume terminal on the 
main axis or rhachilla of the spikelet, or a palea at the base of a 
secondary floral axis. The deciduous part of the spikelet of Phala- 
ridese with its four glumes (or three glumes and a palea) is precisely 
as in OryzesB ; but there are in addition, below the articulation, 
the two persistent empty glumes characteristic of PoaceJB. The 
spikelet, therefore, in this tribe consists of six glumes (or five and 
a palea), the lowest pair empty below the articulation; the second 
pair, above the articulation, corresponding to the lowest two glumes 
of Oryzese, are usually empty and small, sometimes reduced to a 
small bristle, rarely enclosing each a small palea or a male flower; 
the upper pair (or glume and similar palea) enclosing the terminal 
fertile flower and fruit, without any continuation of the rhachilla 
above it. A slight apparent exception will be mentioned under 
Phalaris itself; and in the genus Ginna of AgrostidesB and a very 
few Bambuseee the palea of the fertile flower is, at least apparently, 



76 ME. a. BENTHAM ON GEAMINBiE. 

one-nerved, but otkerwise the character of Phalarideaj is constant. 
They comprise the following six genera : — 

1. Ehehaeta, Thunb. {Trochera, L. 0. Eich.), has twenty-four 
species, of which two are from New Zealand, two from the Mascarene 
Islands, and all the rest from South Africa. In them the glumes 
of the second pair are the largest, empty and usually awned, and 
the fertile flower has six stamens. 2. Miceoij3N"A, Br., including 
Biplax, Hook, f , has five Australian or New-Zealand species, 
differing from EhrJiarta only in the number of stamens reduced 
to four or two. 3. Teteaeehejsta, Br., four Australian species, 
with four stamens to the flower as in MicrolcBna, but the glumes 
are in less regular pairs, all unawned, and the fourth (one of the 
second pair) alone the largest. The panicle is also almost always 
contracted into a spike, not, however, so dense and cylindrical as 
in the following two genera. 

4. Phalaets, Linn., has nine or ten extratropical species, 
chiefly from the Mediterranean region, but also extending to 
North and South America. In this genus it is the lowest two 
persistent empty glumes that are the largest, usually very flat, 
and often winged on the keel, the second pair (like the lowest in 
Oryza) very narrow, sometimes reduced to small bristles, those 
of the upper pair thin and hyaline ; and sometimes in both of 
them, but almost always in the uppermost one, the central nerve 
is very faint or quite obsolete, a character adduced as an argu- 
ment that this upper one is a two-nerved palea on the floral axis, 
and not a glume on the main rhachilla. The two nerves are, 
however, very faint, and the central keel is usually marked by a 
line of hairs on the outside, and the question remains a moot one. 
In the majority of species the panicle is contracted into a dense 
globular or cylindrical head ; but in P. arundinacea, Linn,, a stout 
taU species, forming the geoMsJOigr aphis, Ivm. {Baldingera,QfS&Tin., 
Meg., and Schrad., TypJioides, Mcenoh), the inflorescence, though 
still very dense, is more or less branched or interrupted. This 
genus has also been supposed to be distinguished by the want of 
the broad wings of the outer glumes, so conspicuous in the com- 
mon P. canariensis ; but these wings are very narrow in P. para- 
doxa, Lian., and entirely disappear in P. intermedia, Bosc (P. 
aHimcajjo, Ell.), leaving no available character to separate Z)*^?-^^^'* 
generically. 

5. Ahthoxahthtjm, Linn., has four or five European species, 
of which one is now widely spread over various regions of the 



itE. a. iEKTflAM ON tiRAMllH-Ejtl. 11 

globe, but often only as an introduced weed. One at least of the 
glumes of the lowest pair is the largest of the spikelet, as in 
PAalaris; those of the second pair, though small and without 
flowers, have a dorsal awn. The panicle is usually cylindrical 
and spikelike. 

6. HiEEOOHLOA, Giael.{Savastana,8akraxik,Disarrenum,Tjah\l]., 
Torresia, Euiz and Pav.), about eight species from the colder or 
mountain regions both of the northern and the southern hemi- 
spheres, is usually referred to Avenaces3 next to JSoleus ; but it 
appears to me to be much nearer to Anthomanthum, from which 
it dift'ers in its looser paniculate inflorescence, and in the glumes 
of the second pair being but little smaller than the lower ones, 
and frequently, but not always, enclosing each a male flower. 
Ataxia, Br., one or two Asiatic and two South-African species, 
forms a section of Hierochloa, differing slightly from the typical 
form in the glumes of each pair being more unequal, the lower 
one only of the second pair (rarely both) having a male flower. 
A. mecdcaiia, Rupr., seems to connect the two sections. 

Tribe VIII. AeEOSTBiE. 
The large tribe Agrostese is one of the most diificult to circum- 
scribe satisfactorily, or to divide into definite genera. We have 
taken it nearly in the sense given to it by Trinius, so as to in- 
clude the StipesB, of which other botanists make a distinct tribe ; 
and we have adopted thirty-seven genera, a number which some 
would extend to above eighty, whilst others might reduce it to 
about thirty. Their general character is to have a single flower 
in each spikelet, either apparently terminal as in Panicacese, or 
with a slight bristle-like continuation of the rhachilla beyobd it; 
and from these PanicacesB they are constantly distinguished by 
the pair of empty glumes persistent below the articulation of the 
rhachilla, without any empty glume or male flower intervening 
between the articulation and the flowering glume. The single 
flower in the spikelet, which separates the tribe from the follow- 
ing ones, is not so positive a character, as it occurs also in one 
genus of AvenesB, in a few genera of ChloridesB, and occasionally 
in a few exceptional species of some gene]-a of Festucese, which 
cannot well, from inflorescence or other accessory characters, be 
included in Agrostete. There are also two species of Sporololus 
which approach the Isanthe® in having frequently two flowers ; 
and in Coleanthus the lower empty glumes are entirfily deficient. 

LIIW. JOTJEK.— BOTATfT, TOi. XIX. H 



78 ME. a. BENTHAM ON GKAMrtTEiE. 

Trinius, in his elaborate monograph of the tribe, divided it into 
three primary groups or subtribes — Vilfete with the callus scarcely 
prominent or quite obsolete, Agrostese with the callus globular, 
and StipesB with the callus obconical. In this I feel unable to 
follow him. In the first place he does not appear to have con- 
sidered what the so-called callus really is. It is not, as the name 
would suggest, an appendage to the base of the flowering glume, 
or, as he would have termed it, to the flower, but only the upper 
or principal part of the rhachilla or axis of the spikelet, to which 
the glume and its enclosed flower are attached, and which breaks 
ofi' immediately above the persistent empty glumes. Its shape 
depends on the distance at which the flowering glume is attached 
above the empty ones, a distance very variable throughout the 
Order. And although the long or the short interval may be 
more prevalent or even constant in some genera, yet I have never 
found the variations so precise as to be defined by actual measure- 
ment, and the species are numerous, even in Stipa itself, where 
it is doubtful whether we should call it long or short. It is 
sometimes a useful accessory character, but, I believe, never 
positive enough to be regarded as subtribual. It is true that no 
other simple absolute character has yet been proposed for the 
subdivision of the tribe ; but we are obliged, here as elsewhere, 
to take a combination of characters, to each of which an occa- 
sional exception must be allowed. Acting on this principle, we 
might, whilst following in many respects the arrangements of 
Kunth and others, admit thirty-seven genera of Agrostes, dis- 
tributed in four fairly natural subtribes, all four of wide geogra- 
phical range, but chiefly in temperate regions, the tropical species 
mostly confined to mountain districts, and no genus, except a few 
monotypic ones, exclusively tropical. 

Our first subtribe, Stipeje, is the long-established one of that 
name, slightly extended so as to include Oryzopsis, Muelilen- 
lergia, and their immediate allies, the close connexion of which 
with Stipa has been frequently suggested. The subtribe thus 
formed would be characterized by the paniculate inflorescence 
not condensed into the cylindrical spike of Phleoidese, by the 
rhachilla of the spikelet not produced beyond the flower except 
in the single species of Brachyelytrum, by the awn of the flowering 
glume terminal, not dorsal as in Euagrostea;, and especially by 
the grain being very closely enveloped in the fruiting glume. 
In the majority of species these characters are well marked ; but 



Mb. e. BENIHAM ON SEAMINlSiE. 79 

in the larger genera there occur occasional exceptions more or 
less decided, which prevent our taking any single one of them as 
an absolute teat. The subtribe would include the following eight 
genera, in the first five of which the fruiting glume is more or 
less hardened or rigid as in Paniceae ; in the succeeding three it 
is thinner, though still closely pressed on the grain. 

1. Aeistida, Linn., is now a genus of at least a hundred 
species, abundant in all the warmer regions of the globe, but also 
represented by a few species in Europe and temperate Asia, and 
by several in North America. With few exceptions it is most 
readily recognized by the long, fine, three-branched awns, the 
lateral branches opposite and spreading. Doell adds to the 
generic character three lodicules as in Stipa ; and Nees describes 
three lodicules in some South-African species ; but all other 
Agrostologists describe two only, and I have never found more 
than that number. It is probable that both Nees and Doell 
mistook for the third lodicule the palea, which in many species 
is very thin and scarcely, if at all, larger than the lodicules. The 
genus is divided into three fairly marked sections, which Beau- 
vois, Nees, and some others have raised to the rank of genera. 
In (1) Ghcetaria, Beauv., the flowering glume is continuous with 
the awn without any articulation, and though much longer than 
the empty glumes, and often much attenuated at the end, is 
neither quite awn-like nor decidedly twisted below the branches. 
Amongst its species, Curtopogon was proposed as a genus by 
Beauvois for the North- American A. dichotoma, Mich., in which 
the lateral branches of the awn, instead of diverging from the 
central one, are short and erect at its sides, showing more or less 
distinctly that they are continuations of the lateral nerves of the 
glume. It is probable that this is the case throughout the genus, 
only that the lateral nerves before they diverge are so closely con- 
solidated with the central one as to be undistinguishable from it. 
The genus OrtacJme was proposed by Nees for two or three 
Mexican or Columbian plants, originally published by Kunth as 
species of Streptachne, Br., and afterwards transferred by him to 
Aristida, in which the lateral branches of the awn are very short, 
sometimes minute or even quite obsolete, thus nearly connecting 
the section Ghcetaria of Aristida with the section Aristella of 
Stipa, but in the narrow base of the rhachilla, and some other 
minor points, nearer to the former than to the latter. Ortachne 
retorta, Nees (in Steud. Gram.), is probably a true Stipa. In 

n2 



80 ME. G. BENTHAM ON GBAMINEiE. 

(2) Arthratlwrum, Nees, the awn is dftcidedly articulate ou the 
glume and much twisted above the articulation below the branches, 
the flowering glume itself much shorter than the lower empty- 
glumes, instead of exceeding them as in Ghmtaria. In (3) &ti- 
pagrostis, Nees, the awn is articulate on the glume, as in Arthm- 
therum, but scarcely twisted, and above the branches elegantly 
plumose, the branches also being plumose in some species ; whilst 
in others, forming Kgari and De Notaris's proposed genus Schis- 
tachne, the central awn alone is plumose, the lateral branches 
short and glabrous. All, however, are most conveniently in- 
cluded in the great genus Aristida. 

2. Stipa, Liun., is almost as numerous and as widely spread 
as Aristida. It is also strongly characterized, as to the great 
majority of species, by the narrow, rather hard fruiting glume, 
carrying off a rather long or obconical internode of the rhachilla 
(or so-called callus), by the long undivided awn more or less 
articulate on the glume and usually twisted at the base, and by 
the presence of three lodicules ; but the exceptions to one or more 
of these characters are more numerous than in Aristida ; the in- 
ternode of the rhachilla varies much in length and in shape, the 
articulation aud twist of the awn gradually disappear in some 
species, and the third lodicule, though often as large as the others, 
is sometimes much smaller or even quite obsolete. The genus is 
also not so clearly divisible into sections as Aristida, although 
several genera have been proposed for more or less aberrant 
species. Macrochloa, Kunth, includes 8. tenacissima, Linu., and 
S. arenaria, "Brot., both from the Mediterranean region, remark- 
able for their large membranous glumes, the iiowering one shortly 
bifid at the apex. In Aristella, Bertol., founded on ;8. aristella, 
Linn., a European aud Mediterranean species, in Streptachne, Br., 
a single Australian species, and in OrtTiorapMum, Wees, two or 
perhaps three East-Indian species, the flowering glume is 2-toothed 
or shortly bifid at the apex, the awn scarcely or not at all articu- 
late, and the internode of the rhachilla very short, though still 
perhaps slightly thickened under the flowering glume. The 
S. aristella, however, is very closely connected with typical Stipa 
through iS. sibirica. Lam., S. Redowskii, Trin., and S. altaica, 
Ledeb. Jarava, Euiz and Pav., was founded on S.jarava, Kunth 
{S. eriostacliys, Cav., 8. papposa, Nees), a widely-spread West- 
American species, to which the small spikelets in a long narrow 
dense panicle, with the flowering glumes crowned under the awn 



ME. a. BENTHAM ON GEAMINEiE. 81 

by a pappus-like ring of long hairs, give a very peculiar aspect ; 
but precisely similar flowering glumes are observable in several 
South- American species with very various habits. In the Euro- 
pean S. pennata, Linn,, and a few other American as well as 
Old-World species, the awn itself is (almost entirely, or for a short 
distance above the base) plumose with long spreading hairs. 
Lasiagrostis, Link (Achnaiherum, Beauv.), was proposed as a 
genus for tlie European S. Calamagrostis, Wahlenb., and extended 
by Nees and Trinius to several African and Asiatic species, only 
differing from other small-flowered Stipce in the flowering glume 
itself being plumose with spreading hairs, either below the middle 
or in its whole length ; and in S. mongholica, Trin., forming the 
genus Ptilagrostis of Grrisebacli, these hairs extend to halfway up 
the awn. S. verticillata, Nees, from Australia, and Apera arundi- 
nacea, Hook, f , from New Zealand, two plants closely resembling 
each other, though specifically distinct, connect Stipa with Mueh- 
lenbergia. They have the inflorescence and small spikelets of the 
latter genus ; and in B. verticillata the awn is generally persistent, 
though the articulation is distinctly traceable on the flowering 
glume ; in 8. arundinaeea the awn is very deciduous ; in this 
species there is usually but one stamen, whilst in S. verticillata 
there are the normal three. S. rariflora {MueWenhergiarariflora, 
Hook, f.), from Antarctic America, is another species closely 
allied to the above two ; and all three appear to be better placed 
under Stipa than under Muehlenlergia. 

8. Obtzopsis, Mich. {TTrachne, Trin.), is a genus of about four- 
and-twenty species, from the temperate and subtropical regions 
of the northern hemisphere or from extratropical South America, 
very rare within the tropics, most of them often regarded as 
awned species of Milium, but really more nearly connected with 
Stipa, from which they chiefly differ in the broader fruiting glume, 
often oblique at the top, the awn usually short, slender, and 
twisted, and very deciduous. The genus divides readily into 
three sections, regarded by some as distinct genera, but all united 
into one by Trinius and others. 1. Piptatlierum, Beauv., com- 
prises the Old- World species, often included in Milium as a 
section, with awned glumes, and really connecting in some mea- 
sure the two genera. The obliquity of the fruiting glume is 
much less marked than in the typical species of Oryzopsis ; and 
the rhachilla of the spikelet is glabrous. 2. Euoryzopsis or 
Oryzopsis proper, including the proposed genera CaryocMoa, 



82 ME. G. BESTnAM ON GEAMINEiE. 

Spreng., Piptochatium, Presl, and Nassella, E. Desv., is entirely 
American, with the typical character of the genus, and the rha- 
chilla bearing a ring of hairs under tte flowering glume. 3. Erio- 
coma, Nutt. {Fendleria, Steud.), differs from Euoryzopsis only in 
tte long silky hairs clothing the fruiting glume. 

4. Milium, Linn., was formerly extended to several unawned 
Panieese with only two empty glumes, but is now reduced to five 
or six European or temperate Asiatic species, one of wMch is 
also spread over North America, all removed from Panicaeese as 
having the empty glumes persistent below the articulation. They 
diifer from Oryzopsis chiefly in their obtuse absolutely unawned 
flowering glume. 5. Aoiachnb, Benth., is a single dwarf tufted 
dioecious grass from the higher mountains of Peru and Colombia. 
The female individual, with only one spikelet terminal on the pe- 
duncle, is fully described and figured in the last part of Hooker's 
' Icones.' The male plant, if correctly matched, of which I am 
by no means certain, has a loose almost simple panicle with pre- 
cisely the glumes of the female, but enclosing stamens only. In 
the few specimens seen the leaves are much longer than iij the 
numerous females from various localities, which makes me rather 
doubt the specific identity of the two. 

6. MuEHLENBEEGiA, Schrcb,, has nearly sixty known species, 
chiefly American, extending from the Andes of South America 
over the northern continent generally, with a very few from 
central or eastern Asia. They connect, in many respects, Stipa 
with AgrosUs. In general they come very near in technical 
character to the smaller-flowered Stipas, differing in the still 
smaller spikelets with thinner though still closely appressed and 
narrow fruiting glumes, and usually with a more or less hairy 
rhachilla. Erom Agrostis and its immediate allies they may be 
readily distinguished by this narrow appressed fruiting glume 
with a terminal never dorsal awn ; a very few unawned species 
are scarcely separable from JSpieampes, except by the shape of the 
glume. There is a considerable variety in the inflorescence and 
in the proportions of the glumes, but nothing definite enough to 
establish good sections, although several separate genera have 
been proposed. In the original M. diffusa, Schreb., and its im- 
mediate allies, the panicle is usually long, narrow, and dense, 
and the lower empty glumes are very minute ; whilst in Trinius's 
proposed section Acroxis both the lower glumes or one only of 
them are nearly as large as the flowering one ; but throughout the 



MK. a. BENTPAM OK aKAMIKEiE. 83 

genus the relative size of these glumes appears to vary almost 
from species to species. Vaseya, Thurb., is a Californian species, 
M. comata, Thurb., closely resembling the more common M. syl- 
vatiea, Torr., except in the long hairs surrounding the thin 
flowering glume. Podoseemum, Desv. {Trichochloa, Beauv.), com- 
prises a number of elegant species, in which the spreading panicle 
has a number of small long-awned spikelets on long capillary 
branches and pedicels. Tosagris, originally separated by Beau- 
vois from PodoscBmum on account of the long hairs on the back 
of the flowering glume, was subsequently reunited with it by the 
author himself. Olomena, Beauv., is M. clomena, Trin. (Jf. na7ia, 
Benth.), a dwarf Andiue species, in which the second empty 
glume is the largest of the spikelet, and rather broadly three- 
toothed. The same character is observable in M. gracilis, Trin., 
forming Nuttall's genus Galycodon. 

7. BEACHXEiYTErM, Beauv., is a single North- American species, 
very near to some species of Stipa; but the rhachilla is produced 
beyond the flowering glume into a little bristle, sometimes bearing 
a minute rudimentary glume, which does not occur in any other 
species of the subtribe. 8. Peeieilema, Presl, contains three 
or four tropical or subtropical American species, with much 
of the habit and many of the characters of MueJilenbergia dif- 
fusa, but with the empty glumes awned as well as the flowering 
one. 

Our second subtribe, Phleoideje, is chiefly characterized by the 
inflorescence. The panicle is condensed into a globular or oblong 
head or cylindrical spike ; the rhachilla is, ia a few species only, 
produced beyond the flower into a small bristle ; the flowering 
glume either is awnless or bears one or three terminal awns, and 
when in fruit is thinner than in Stipese, more loosely enclosing 
the grain as in Euagrostese. The following seven genera, or most 
of them, have already been placed in juxtaposition by various 
Agrostologists. 

9. Ltctjeus, H. B. K. (Pfeq^oyow, Nutt.), consists of two closely 
allied American species, perhaps varieties of a single one, readily 
known by the empty glumes as well as the flowering one awned, 
as in Ferieilema, the lowest one having usually two or even three 
awns. The long dense cylindrical spike (or spike-Hke panicle) 
with sterile spikelets intermixed with the perfect ones brings the 
genus in connexion with the subtribe Sesleriese of Pestucese ; but 
there is never more than a single flower in the spikelet, 



84 ME. G. BENTHAM ON GEAMINEiE. 

10. EcHiKOPOGON, Beauv. {Hystericina, Steud.), a single Aus- 
tralian and New-Zealand species, has likewise sterile spikelets 
intermixed with the perfect ones ; but the empty glumes are 
awnless, and the flowering one three-lobed with the middle lobe 
produced into a long awn. 11. Diplopo&oit, Br. {Dipogonia, 
Beauv.), also a single Australian species, has a short awn to 
the empty glumes and three to the flowering one, of which the 
central one is long and twisted. 12. Amphipogoit, Br. (.^yqpo- 
gon, Beauv., not of Willd., Pentacraspedon, Steud.), five Austra- 
lian species, has the flowering glume deeply three-lobed and fre- 
quently awned, and the palea also with two rigid almost awn-like 
lobes. Gamelythrmn, Nees, is the A. turbinatus, Br., separated 
from Ainphipogon only on account of a more distinct elongation 
of the rhachilla between the outer glumes and the flowering 
one. 

13. Heleochloa, Host {Pechea, Pourr.), contains seven or 
eight Mediterranean species, of which one or two are widely 
dispersed over Europe and Central Asia. Kunth referred them 
to a section of Orypsis ; and Host himself subsequently assented 
to the union, probably misled by an apparent resemblance of some 
varieties of H. schaenoides to the true Orypsis aouleata ; but the 
resemblance is apparent only, the two genera are as essentially 
difierent in inflorescence as in the structure of the spikelets. 
The axis of inflorescence, or receptacle, in Crypsis is a flat disk ; 
in Seleoeliloa it is a more or less elongated linear rhachis, cylin- 
drical even in those varieties where the spike-like panicle is con- 
tracted into a sessile head. In Crypsis the empty glumes are 
above the articulation and fall off" with the spikelet, and the 
glumes are quite those of Oryzese without any two-nerved palea ; 
in Heleochloa the empty glumes persist below the articulation, 
and the glumes and palea are entirely those of Phleoideae ; and 
although in the commonest species the spikelike panicle or head 
is short and sessile, yet there are others where it is long, narrow- 
cylindrical, and pedunculate. Pliizocephalws, Boiss., founded on 
Crypsis pygmcea, Jaub. and Spach, makes, with O. amligua and 
C. crucianelloides of Balansa, a very good section of Seleochloa, 
distinguished by the dwarf tufted habit and the spikelets almost 
echinate with the rigid points of the glumes. Beauvois gave the 
same name Heleochloa to a supposed genus, apparently made up 
of a Sporoholus and a Phleum. 

14. Mailiba , Parlat., is the Phalaris crypsoides, Durv., a dwarf 



ME. G. BENTHAM ON GBAMINEJ!. 85 

tufted G-reek plant, with the spilielets flat as in Plialaris, but 
otherwise showing nearly the structure of Phleuin. 

15. Phleum, Linn., about ten species from the temperate and 
northern regions of the northern hemisphere or from Antarctic 
America, is a well-known and already well-defined genus. It has 
been proposed to separate generically Ghilochloa, Beauv. {AcJino- 
don, Link), for the few species in which the rhachilla is produced 
beyond the flower into a minute bristle ; the character, however, 
is in this instance very trifling and uncertain. Aohnodonton, 
Beauv., is P. tenue, Schrad., for which I can find no separate 
generic character. The anomalous Phalaris trigyna, Host, appears 
to have been an individual specimen of Phleum MicJielii, All., 
having abnormally three style-branches instead of two. 

In a third small group or subtribe, SpoEOBOLEiE, I should pro- 
pose to place Sporololus itself, with the three monotypic genera 
Mihora, Coleantftus, and Phippsia. -The subtribe is not very 
clearly defined ; but my previous endeavours to associate Sporo- 
holus with Milium and Isachne, to which I shall recur further on, 
proved still less satisfactory. The plants now grouped together 
have small paniculate or almost racemose spikelets, awn less glumes, 
no continuation of the rhachilla beyond the flower, and the ripe 
grain only half enclosed in and readily falling away from the 
glume — characters sometimes well marked, but in some species 
rather vague. 

16. MiBOEA, Adans. (Ghamagrostis, Borkh., Sturmia, Pers., 
Knappia, Sm.), is a dwarf slender tufted European annual, with 
a simple spike and the lower empty glume at least as long as the 
flowering one. 17. Colbanthtjs, Seid., is a minute annual, first 
found in Bohemia, then in Norway, and more recently gathered 
in the island of Sauvies at the mouth of the Oregon in North- 
west America. It is very near Phippsia and Sporobolus ; but the 
lower empty glumes are entirely deficient. It was first disco- 
vered by Seidel, and distributed by him under the name of Cole- 
anthus sultilis ; but Trattinick in publishing it (as reported by 
Eoemer and Schultes) retained only Seidel's specific name, changing 
the genus to Schmidtia. Ecemer and Schultes in their ' Systema ' 
restored Seidel's name, which Sternberg, rather later, changed 
again both generically and specifically to Schmidtia utriculosa. 
Under these circumstances Seidel is now considered to have pub- 
lished his Coleanthus sultilis sufficiently for general adoption, 
more especially as another very different genus of Grasses has 



86 ME. a. BENTHAM ON GEAMINEiB. 

received the name of Sohnidtia. 18. Phippsia, Br., is a dwarf 
paniculate slender Arctic grass, cMefly distinguishable from Spo- 
rdbolus by the minute lower empty glumes. 

19. SpoKOBOLrs, Br. (Vilfa, Beauv., Agrosticula, Eaddi, Tria- 
chyriwn, Hochst., Gryptostachys, Steud.), is now a genus of about 
eighiy species, spread over the warmer and temperate regions of 
both the New and the Old "World, mostly, however, American, with 
a very few European or Asiatic. Included by the older authors 
in Agrostis, it has since been universally acknowledged as dis- 
tinct, though different characters have been assigned to it. 
Beauvois, who attached primary imp ortance to the presence or 
absence of the awn, referred to it all the unawned species of the 
old genus Agrostis. Brown, who first pointed out the differences 
in the fruit, took as the principal character the loose membra- 
nous pericarp readily detachable from the seed ; but this, though 
very conspicuous in S. indicus, Br., and in some other species, is 
not apparent in the dried state in several others ; and in S. virgi- 
cus, Kunth, and others, it is only when soaked that the pericarp 
can be detached. On this account it has been attempted to 
establish two genera, Vilfa and Spordbolws ; but the character is 
far too indefinite, as well as uncertain, to be available even for 
sectional separation. As a whole, Sporoholus is chiefly distin- 
guished from Agrostis by the total absence of any dorsal awn, 
and by the grain so loosely enclosed in the glume that it usually 
protrudes from it when ripe, and often falls away. The palea 
also generally splits readily into two, and in some species is 
even at the time of flowering divided to the base, a character 
which G-risebach, who only observed it in an Argentine species, 
was induced to take as that of a new genus Diachyrium ; but it 
exists in many other species ; and this divided palea has been 
more than once described, and even figured (as in T. Nees's 
' Genera Moraa Germanicse '), as a two-valved pericarp, a character 
unknown in Graminese. Brown's name Sporoholus was rejected 
by Beauvois, Trinius, and others on the supposition that the genus 
is identical with the older established Vilfa, Adans. That, how- 
ever, is a mistake. Adanson's character of Vilfa is so vague 
that it cannot be identifled by that alone ; but in his index he 
fixes it by quoting two European species, which are certainly both 
of them true species of Agrostis, 

Two North-American species of Sporoholus, S. compressus, 
Trin., and S. serotinus, A. G-r., are exceptional, not only in the 



ME. G. BENTHAM ON GEAMrfTEiE. 87 

genus, but in tte whole tribe of Agrostese, by the spikelets con- 
taining occasionally two flowers (without awns or continuation of 
the rhachilla) as in Isachnese and in some species of Aira and Gol- 
podium ; but the small spikelets and carpological characters are 
quite those of Sporobolus. 

There remain for the ETTAGEOSTEiE, or fourth and last subtribe 
of Agrostese, about sixteen genera, of which the general character 
is a dorsal usually twisted awn on the flowering glume, the grain 
neither so closely enveloped in the fruiting glume as in Stipese, 
nor so readily exposed as in SporobolesB, and the spikelets usually 
small, loosely paniculate, very rarely condensed into a head as in 
Phleoideae ; but there are exceptions to every one of these charac- 
ters, and the limits of the larger genera are so vague as to render 
this portion of the genera of Gramineae the least satisfactory of the 
whole series. Of the sixteen following genera, the first seven 
show none of that continuation of the rhachilla beyond the flower 
which in the others takes the form of a glabrous or hairy bristle 
rarely reduced to a mere tubercle ; the last four of the series, as 
well as Triplachne, have, besides the dorsal awn, two or four teeth 
to the glume, sometimes produced into straight awns. A few spe- 
cies or monotypic genera have no awn to the flowering glume, 
but otherwise in the structure of the spikelet are nearer to Agrostis 
than to Sporoholus. 

20. Epioampes, Presl, about sixteen species from Mexico and 
the South-American Andes, probably reducible to about two 
thirds of that number, is a genus most embarassing to the syste- 
matist ; for it seems to connect Muehlenbergia and Sporoholus with 
Agrostis. The chief general feature is the long narrow dense 
panicle with very numerous rather small spikelets, the awn 
of the flowering glume, when it exists, much smaller, than in 
Muehlenbergia and often not quite terminal ; the unawned species 
distinguished from Sporobolus by the fruiting glume and grain 
nearly those oi Agrostis, and from the latter genus by the inflo- 
rescence and by the awn when present being very small and 
almost terminal. Several of the published species, however, are 
unknown to me ; and a further study may require considerable 
modification of the generic character and limits. Orypsinna, 
Fourn., which appears inseparable from Epicampes, is founded 
on the E. maoroura (Cinna macroura and G. striata, Kunth), a 
Mexican species remarkable for the very long, narrow, almost 
spikelike panicle. Cinna macroura, Thurb., from California, is a 



88 ME. a. BENTHAM ON GRAMINEiE. 

distinct Bpecies (U. ripens, Beath.) with a still longer and nar- 
rower rigid spikelike panicle often interrupted ; and the spikelets 
are small, with membranous glumes, as in the typical U. siricta, 
Presl, but awnless or nearly so. 

21. BAroHEA, Fourn., a single Mexican species unknown to me, 
might perhaps be reduced to JEpicampes, from which it is said to 
differ chiefly in a great inequality of the two empty glumes. 

22. Ageostis, Linn., even after being shorn of a number of 
heterogeneous plants ascribed to it at various times from general 
aspect, and after the suppression of numerous names given to 
local representatives of cosmopolitan species, is still a genus 
of nearly a hundred species, generally spread over nearly the 
wliole world, but especially common in the temperate regions 
of the northern hemisphere. Among the Buagrostese without 
any continuation of the rhachilla they are generally known 
by the absence of those peculiarities which have induced the 
separation of several of the preceding as well as of the follow- 
ing genera, by the thin short broad flowering glume with the 
dorsal awn below the middle, and by the palea not more than 
half as long as the glume, and often quite minute or even defi- 
cient ; the panicle is also usually loose, very elegant, with nume- 
rous small spikelets or almost capillary branches and pedicels. 
There are, however, here, as elsewhere, exceptional species which 
defy all neat classification ; even the dorsal awn is sometimes re- 
duced to a minute tubercle, or only to be guessed at by the abrupt 
termination of the central nerve of the glume. The American 
species in which the palea is minute or deficient formed Michaux's 
genus Trichodium, which has been extended to the European 
A. canina, Linn., and others witli that character, to which Beau- 
vois restricted the Linnean name Agrostis, whilst he gave the name 
of Agraulus to the species in which the palea is more developed. 

23. CH.a;TrB.rs, Link, is a single Spanish annual, much like 
some species of Agrostis, but anomalous in the group in having 
the lowest (empty) glume larger than the others, and produced 
into a long awn, whilst the flowering one, though shaped as in 
Agrostis, has no awn. The inflorescence is also peculiar, each 
branch of the panicle terminating in the three shortly pedicellate 
spikelets. 

24. Aectageostis, Griseb., is a single arctic species, referred 
by Brown doubtfully to Trinius's genus Oolpodium, but which 
appears to be more nearly allied to Deyeuxia. It is, however, 



ME. G. BENTHAM OIT GEAMHiTE^. 89 

one of those plants which, by irregularity in some characters 
usually very important, is very difficult to place satisfactorily. 
The habit and size of the spikelets are more those of Poa than of 
Agrostis ; but, in the great majority of specimens, the one-flowered 
spikelets without any continuation of the rhachilla are quite 
those of Agrostese, and the palea is fully the length of the glume 
as in Deyeuocia. Very rarely specimens have presented themselves 
with a minute continuation of the rhachilla ; and Brown, in a single 
Melville-Island specimen, found it to bear a,n empty glume or 
second flower, thus showing a connexion, and possibly, in the spe- 
cimen mentioned by Brown, a hybrid between Arctagrostis and 
Foa alpina. 

25. CAiiAMAGEOSTis, Adans., as now limited, comprises four or 
five species from Europe and northern and central Asia, of which 
one has also been found in South Africa, possibly, but not cer- 
tainly, introduced there. Some authors extend the genus so as 
to include the greater part of Deyeuxia, and indeed all tlie 
Euagrostese with a hairy rhachilla ; but it seems more natural if 
confined to the typical species, which, like Agrostis, have no con- 
tinuation of the rhachilla or rarely a very slight one, and bear on 
the flowering glume a fine dorsal awn, rarely reduced to a minute 
point. They differ from Agrostis in the ring of long hairs sur- 
rounding the flowering glume, and generally in their tall almost 
reed-like habit, whence their generic name, and on which account 
they have often been placed in juxtaposition with Arundo. They 
appear, however, to be in every respect true Agrostese ; and there 
are two species, C. tenella, Kunth, and C. olympica, Boiss., which 
are almost intermediate between Galamagrostis and Agrostis, espe- 
cially as a few species of true Agrostis are not entirely without 
hairs on the rhachilla. 

26. CiNNA, Linn. {Ahola, Adans., Blattia, Eries), is limited by 
modern authors to two species from the northern regions of 
Europe and America, with the tall reedlike habit of the larger 
species of Oalamagrostis, but with a glabrous rhachilla, and re- 
markable in the tribe by the palea having only one nerve, although 
there is every reason to believe that it is a true palea, the appa- 
rently single nerve being due to the consolidation of two. Both 
species appear also constantly to have but one stamen in the 
flower. Some botanists unite the two ; but from dried specimens 
they appear quite distinct. Amongst other minor points, the 
original 0. arundinacea, Linn., has generally a minute continua- 



90 ME. G. BENO^HAM ON GEAMINEJ!. 

tion of the rhachilla, which I have never found in G. pendula, 
Trin. (C. expansa, Link, G. latifolia, G-riseb.). Several Americau 
or other grasses published as species of Ginna are now referred to 
Mpicampes or Deyeuxia. 

27. Gasteidhtm, Beauv., has two species from the Mediterra- 
nean region, one of them also found in tropical Africa and in 
extratropical South America, but possibly introduced only in. the 
latter locality. They have the small spikelets of Agrostis, but a 
narrower closer panicle, and are remarkable for the outer glumes 
rather hardened, shining, and ventricose at the base, whence the 
generic name. The older authors included them in Milium on 
account of that hardness in the glumes. 

28. Chjetoteopis, Kunth, is a single Chilian species which 
some would unite with Agrostis, and might well have been joined 
to Hpicampes, but that the rhachilla is produced beyond the 
flower into a rather long hairlike seta. 

29. Teiplachne, Link, is the Gastridimn nitens, Coss. etDur., 
a single Mediterranean species, with the habit, but not the ven- 
tricose glumes, of that genus, and differing both from that and 
from Agrostis in the flowering glume bearing a short awn-like 
point on each side of the awn. 

30. Apeka, Adans. (Anemagrostis, Trin.), has two very closely 
allied European species extending into Western Asia, with the 
technical character very nearly of Deyeuxia, but with the elegant 
panicle and numerous small glabrous spikelets of many sjiecies of 
Agrostis, in whicli they are still included by some under the name 
of Agrostis spica-venti. The New-Zealand plant described by 
Hook. f. as an Apera is now transferred to MueMenbergia. 

31. CiNKAGEOSTis, Q-riseb., from Tucuman in South America, 
is unknown to me, but is said to differ from Deyeuxia in having 
the spikelets unisexual by abortion. It should most probably be 
incorporated in that genus. 

32. Detetjiia, Clarion {Lachnagrostis, Trin.), has now nearly 
a hundred and twenty species, dispersed over the temperate or 
mountain regions of the globe, particularly numerous in the 
Andes of South America, and extending northwards to the Arctic 
circle and southwards to the extreme end of South America. It 
is in some respects polymorphous, running on the one hand 
almost into Agrostis, to which some species have been referred, 
and on the other into Galamagrostis, with which the northern 
species have been often united. It differs from both in the pro- 



ME. G. BBNTHAM ON GEAMINEiE. 91 

longation of the rhachilla into a bristle or stipes usually, but not 
always, hairy, and from Agrostis in the larger spikelets, with the 
palea nearly as long as the glume, and the usually hairy rhachilla. 
There are, however, a few species where one or another of these 
characters fail ; and one or two scarcely diifer from Apera except 
in the habit and in the awn being more decidedly dorsal. Sro- 
midium, Nees (to which belong Bidymochceta, Steud., and Ohamm- 
oalamus, Meyer), contains a few Andine Chilian or Australian 
species, which, with the minute glabrous and perhaps not con- 
stant continuation of the rhachilla, might almost as well be 
transferred to Agrostis, but that they have rather the habit of 
Deyeuxia. Relchela, Steud. {Agrostis sesquivalvis, E. Desv.), 
Cinnastriom, Pourn. (at least as to Deyeuxia poedformis, Kunth), 
Deyeuxia mutica, Wedd., and D. Ireviglwmis, Benth., with a few 
other South- American species, form a little group with a glabrous 
rhachilla and the awn reduced to a small point. In Achmta, 
Fourn., two Mexican species, the awn appears to be deficient ; 
but all the other characters are those of the typical Z>eyeMa;«<e with 
a hairy rhachilla, to which I would also refer the Agrostis aqui- 
valvis, Trin., forming Grisebach's section Podagrostis. 

33. Ammophila, Host (Psamma, Beauv,), comprises four spe- 
cies, two of them widely spread over the northern hemisphere 
chiefly near the sea, and two confined to North America. They 
are distinguished from Deyeuxia by their tall habit, their usually 
dense inflorescence, and especially by their larger paleaceous 
glumes. 

34. DiOHELACHNE, Endl., two Australian or New-Zealand spe- 
cies, with a narrow dense panicle, differs from Agrostis and its 
allies in the flowering glume scarcely smaller than the outer 
empty ones and often toothed, and in the long dorsal awns giving 
the inflorescence a fine bristly aspect. 

35. Teisetaeia, Porsk. (Anomalotis, Steud.), is a maritime 
Syrian and Egyptian plant, very near Uichelachne, but still more 
bristly, the lateral teeth of the flowering glumes being produced 
into fine straight awns, whilst the dorsal one is longer and flexuose. 
Labillardiere and Delile both mention two fertile flowers in the 
spikelet. I have only been able to find one in several specimens 
examined, all from Alexandria; possibly they may have consi- 
dered the rather long continuation of the rhachilla as a second 
flower. 

36. PENTAPO&oif, Br., is a single Australian species, with four 



92 ME. G. BENTHAM ON GEAMINEJE. 

straight awns to the flowering glume, besides the long rigid 
twisted dorsal one, which, as well as the single flower, removes 
the genus from the Pappophorese. 

37. Lagubtjs, Linn., is a well-known widely spread Mediter- 
ranean grass, which, like Trisetaria, has two slender awns to 
the flowering glume besides the more rigid dorsal one, but is 
well marked by the capitate inflorescence, to which the long hairs 
of the linear plumose empty glumes give a peculiar soft silky 
aspect. 

Tribe IX. IsACHNEiE. 

This small tribe is a modification of the subtribe I proposed 
in the ' Plora Australiensis ' under the name of Milieae, and which 
I distinguished from AgrostesB by the absence of the dorsal awn, 
and from Testucese by the single or two equal flowers in each 
spikelet ; and I included in the group both Milium and Sporo- 
bolus. Since I have worked up the , Agrostess of the northern 
hemisphere, however, I find that the presence or absence of the 
dorsal awn is much more uncertain than I had thought, that 
Milium cannot be removed far from Oryeopsis, and that Sporo- 
iolus must be referred back to Agrosteae. But there remain a 
group of genera, nearly related both to Agrostess and to Avenese, 
but never showing the dorsal awn so general in those tribes, 
and enclosing in each spikelet two equal flowering glumes and 
flowers, apparently inserted at the same point without any deve- 
lopment of the rhachilla between them (except in Ccelachie) and 
never any continuation beyond the flowers. The two fl.owers are 
both hermaphrodite and fertile ; or occasionally only one of them, 
usually the upper one, is female or sterile. The tribe thus limited 
would consist of the following seven genera : — 

1. Peionachite, Nees, subsequently republished by the same 
author under the name of Ghondrolcena, is a South-African annual 
with an almost simple terminal spike, distinguished by the outer 
empty glumes as long as the flowering ones, with a rigid pec- 
tinately-toothed cartilaginous keel. Ktenosaohne, Steud., is most 
probably the same plant. 

2. IsAOHNE, Br., comprises about twenty tropical or subtro- 
pical species, chiefly from the Old World, but including a few 
American ones, The small spikelets with the loosely paniculate 
inflorescence and more or less hardened fruiting glumes give them 
the appearance nearly of some species of Panicum, to which 



Ml!. 6. BENTUAil Oif GHAMINEJi. 93 

genus some species of Isachne Lave been referred, but from -whicli 
they constantly differ in the empty glumes persistent below the 
articulation, and in the two flowers both hermaphrodite or female, 
though one may be occasionally sterile. Graya, Nees (judging 
from the reference lo Wight, but not from Steudel's characters) 
is Isacfine pulcliella, lioth {Panieum helium, Steud., P. malaccense, 
Trin.). Fanicum Qardneri, Thw., which, as the author observes, 
is closely allied to Isachne Wallceri, Wight, appears to me to bo 
strictly congener with that species, although one of the flowers of 
the spikelet is frequently, but not always, sterile. It seems to be 
the same as Isachne nilaghirica, Hochst. 

3. Zenkeeia, Trin., very well described and figured in the 
'Linnsea,' vol. xi., now contains two species from the East-Indian 
peninsula and Ceylon, both very near Isachne, but with membra- 
nous fruiting glumes. Amphidonax Seynei and A. fenella, Nees, 
do not differ from the typical Z. elegans, Trin. The second spe- 
cies is Z. oltusiflora {Amphidonax ohtusiflora, Thw.). The ori- 
ginal genus Amphidonax of Nees was founded on a species of 
Arundo. 

4. MiCEAiEA, if. Muell., is a single North-east Australian spe- 
cies recently figured in Hooker's Icones. 5. Coslachne, Br., 
comprises three East-Indian, Chinese, or East- Australian species^ 
G. pulchella, Br., C. perpwsilla, Thw., and O. simpliciuscula, Munro 
{Isachne simpUciuscula,'Wight et Arn.), which, as above observed, 
are anomalous in the tribe by a slight extension of the rhachilla 
between the flowering glumes. 6. Aieopsis, Desv., restricted to 
the single West-Mediterranean A. globosa, is a pretty little 
annual, formerly placed in Milium on account of the hardening 
of the glumes, or in Aira, which it resembles in many respects. 
It shows, however, all the characters of Isachnese, and is indeed 
technically nearly allied to Isachne itself ; but the two semiglobose 
fruiting glumes, closely appressed to each other by their flat 
faces, give the spikelets the peculiar globular shape expressed by 
the specific name. 

7. Eeiachne, Br,, comprises twenty-two species, two of them 
endemic in tropical Asia, the remainder Australian, of which one is 
also in East India. They diff'er from Isachne generally in their 
rather larger spikelets, and especially in the long hairs on the back 
or margins of the flowering glumes, and sometimes in the fine 
straight awns terminating the flowering glumes, or even the teeth 
of the paleffi. Megalachne, Thw. (not of Steud ), is Uriachne triseia, 

LIHK. JOUKN.— BOTANT, TOL. XIX. I 



94 MB. G. BENTHAM ON GEAMINEJE. 

Nees, in which these awns are particularly conspicuous. The 
African species referred by Nees to a section Achneria oi Mriachne, 
form a distinct genus of Avenese, for which Munro has retained 
this n&me AcJmeria, the original genus Achneria of Beauvois having 
been proposed for those true Australian species of Uriachne which 
have uo awn or only a very small one. 

Tribe X. Ateneje. 

This tribe has been more generally recognized and subjected to 
less variations than most of the others. Its general characters — 
the paniculate inflorescence, the spikelets with two or more per- 
fect flowers, the rhachilla"produced beyond the upper flower, and 
a twisted awn to the flowering glume either dorsal or terminal 
between the two lobes or teeth of the glume — sufi'er fewer excep- 
tions than usual. Ai7-a alone has no continuation of the rhachilla ; 
and Anisopogoii alone has only one perfect flower in the spikelets. 
Of the following sixteen genera, the first eleven have the awn 
dorsal and the lowest flower hermaphrodite ; the next three have 
the male or sterile flower below the perfect one ; and the last two 
have the lowest flower hermaphrodite and the awn terminal. 

1. AiEA, Linn,, was once made to include GorynepJiorus, Des- 
champsia, and indeed almost all the Avenese with loosely panicu- 
late inflorescence and small two-flowered spikelets, but has since 
been so thoroughly dismembered by various European botanists 
as not to leave a single species to represent the old Linnean 
name. Taking, however, the widely spread A. oaryophyllea, Linn., 
as a genuine type, and adding to it five or six European species, 
we have a natural genus of elegant, slender, mostly annual grasses 
with fine filiform leaves, the small spikelets always two-flowered 
without any continuation of the rhachilla beyond the upper flower, 
the dorsal awn of the flowering glumes rarely wanting, and the 
ripe grain often adhering to the palea ; the latter character, how- 
ever, is always uncertain. , These six or seven species have all 
been made the types of supposed distinct genera. A. caryopliyllea, 
Linn., and A, prcBCox, Linn., considered as typical Aircz by Par- 
latore and others, form the genus Fussia, Schur, in which the two 
flowers are closely contiguous and the flowering glumes usually 
awned. Morinia, Parlat., is the A. Tenorii, Gruss., distinguished 
by the absence of the awn ; but Q-ussone has shown that it varies 
with or without the awn. Antinoria, Parlat., is the A. agrostidea, 
Lois., with the rhachilla more or less lengthened between the 



ME. Q. BElfTHAU OU CHlAMnfE^. &5 

flowers, and the glume usually uiiawued ; but A. pidchella, "Willk., 
with the glume awned, cannot be otherwise distinguished from 
it. Periballia, Triu., is the A. involucrata, Cav., in which the 
two flowers are as in A. agrostidea rather distant from each other, 
the lowest flowering glume unawned, the upper one awned, but 
both flowers hermaphrodite, as in the rest of the genus. The 
inflorescence of this species is rather peculiar ; the lowest whorl of 
branches of the panicle are usually without any or with only very 
few spikelets, and were regarded by Cavaailles as an involucre : 
but that is not always the case ; I have seen some specimens 
with spikelets on all the branches. A. sabulonum, LabilL, from 
New Caledonia, is a very doubtful plant. Labillardiere's figure 
is a good representation of the Australiasan form of Sporobolus 
virginicus, except that the spikelets are drawn as two-flowered. 
The specimens sent for Labillardiere's plant by Pancher and by 
Vieillard have only one flower in the spikelet. 

2. CoETiTEPHOKTJS, Beauv. (Weingartneria, Bernh.), comprises 
two European grasses, extending into Worth Africa and more 
sparingly to the Levant, with the continuation of the rhachilla 
of Deschampsia, but readily distinguished by the peculiar articu- 
late club-shaped awn of the flowering glumes. 

3. Deschampsia ( Oampeliaj Jjink) is a genus of about twenty 
species, from the temperate or colder regions of both the New 
and the Old World, sparingly represented in mountain regions 
within the tropics. It bears the same relation to Aira that 
Beyeuxia does to Agrostis ; the plants are usually perennial and 
stouter than in Aira, the spikelets larger, and the rhachilla is 
produced beyond the upper flower into a bristle often bearing a 
tuft of hairs, and sometimes an empty glume on even a male 
flower ; the flowering glumes are also frequently more or less 
denticulate. No less than six of the species have been proposed 
as distinct genera: — Vahlodea, Pries, is D. atropurpurea {Aira 
atropurpurea, Wahlenb.), a northern species, with the flowering 
glume not at all or only very minutely denticulate, otherwise 
quite a Deschampsia. Avenella, Parlat. (Lerchenfeldia, Schur), 
is the common D.flexuosa {Aira flexuosa, Linn.), with the flower- 
ing glume surrounded by hairs. The grain is said by Parlatore 
to adhere to the palea, which may be sometimes, but is certainly 
not always, the case. Monandraira, Em. Desv., includes two 
Chilian species, Trisetwn Berteroanum and T. airceforme of Steudel, 
separated from Deschampsia as having but one stamen to the 

i2 



96 UK. G. BEJSTHAM ON 6EAMINEJS. 

flower ; but iu D. Berteroanum I have sometimes fouud two 
stamens, and iii the evidently nearly allied Z). antarctica, Hook, f., 
the stamens, though usually three, are sometimes two only. 
Airidium, Steud., is a species from the Straits of Magellan which 
I am unable to distinguish from the D. antarctica. Bytidosper- 
mum, Steud., is founded on specimens of a Deschampsia closely 
allied to, if not identical with, the common D. C(Sspitosa,in which 
a grub has taken possession of every spikelet remaining in the 
panicle, and has been mistaken by Steudel for the caryopsis, and 
actually described as such. Peyritschia, Fourn., is D. Jccelerioides 
(Aira Jccelerioides, Peyr.), which I have not seen, but which, from 
Peyritsch's elaborate description, must be very near to D. ant- 
arctica, Hook, f., D. nitida, Presl, and D. Jiolciformis, Presl. 

4. AcjiNEEiA, Muuro, contains eight South-African species, 
with one from south-eastern tropical Africa, referred by Nees to 
Eriachne, and by Kuuth to Airopsis, but evidently more nearly 
related to DescTiampsia. 

5. MoNACHTEON, Parlat., is a single species from the Cape- 
Verd Islands, which we have not at Kew, and of which I have 
therefore been unable to verify the character given by Parlatore. 
The specimen he described most probably remained in Webb's 
herbarium, now deposited at Plorence. 

6. HoLCTJS, Linn., formerly included two very different groups 
of grasses ; and Brown specially retained Linnteus's name for that 
one which now forma the genus Sorghum in Andropogonese, 
whilst all modern botanists restrict the genua Holcus to the other 
group, consisting of about eight European or African species, 
chiefly western, of which one or two are common weeds in various 
parts of the world. All are nearly allied to DescTiampsia, but 
have the upper flower of each spikelet male with an awned glume, 
and the lower one unawned and hermaphrodite. Two Spanish 
species have been added by Boissier, H. grandiflorus and H. cass- 
pitosm ; but as they have both the flowers liermaphrodite and 
awned (whence the sectional name Romalachne), they should 
rather be transferred to Bescliaonpsia, although they may have 
the peculiar soft habit of the common species of Holcus. 

7. Teisettjm, Pers., is now known to comprise nearly fifty 
species, ranging over the temperate or mountain regions of both 
the New and the Old "World. All are very near to the section 
Avenastriim of Avena, but differ generally in the flowering glume 
decidedly toothed at the apex, the two teeth often produced into 



MR. G. BENTHAM OS GUA.UIHEX. 97 

straight awns, one on each side of the dorsal twisted one, and in 
the grain glabrous or slightly pubescent at the apex without the 
longitudinal furrow of Avena. The inflorescence is also usually 
more dense than in that genus, with smaller, often shining spike- 
lets. A few African or South-American species, however, such 
as T. hirtum, Nees, and T. antarcticum, Nees (whicb includes 
Bronius antarcticus. Hook, f., and Bromus hicuspis, Nees), closely 
connect the two genera : the flowering glume is more rigid and 
less keeled than in the true Triseta, and the ovary is pubescent 
at the top ;but the grain has not the furrow of Avena. Triehata, 
Beauv., is the Trisetum ovatum, Pers., a species allied to T. sub- 
spicattim ; but the spikelike panicle is more dense and ovoid, or 
almost globular. Acrospelion or Aeropselion, Bess., is Trisetmn 
distichophi/llum, Beauv., not the Ventenata, Link, to which it is 
referred in Lindley's ' Vegetable Kingdom.' Mostraria, Trin., 
was made up of Trisetum neglectum, Eoem. et Schult., and Koeleria 
phJeoides, Pers. 

8. Ventenata, Keel., has two species, V. avenaeea, Koel., and 
Y. macra, Balansa, from the Mediterranean region and Central 
Europe, differing slightly from Trisetum in the longer, more rigid, 
many-nerved glumes, and the absence of any dorsal awn on the 
lower flowering one. 

9. Atena, Linn., as limited by recent authors, comprises about 
forty species, mostly from the temperate regions of the Old 
World, with a few from extratropical North and South America, 
and one or two of the annual ones cornfield weeds in other coun- 
tries. It is generally characterized by the flowering glumes 
rounded on the back and several-nerved, with a dorsal twisted or 
bent awn, and by the ripe grain furrowed in front and more or 
less adhering to the palea, but is divisible into two sections almost 
marked enough in habit as well as character to be raised to the 
rank of genera. In 1. Crithe, Griseb.,the species are all annual, 
usually tall, with a loose panicle of large pendulous spikelets, each 
containing no more than two fertile flowers, and often only a 
single one, and the lower empty glumes 7- or 9-nerved. This 
section includes the common Oat, which has lost its dorsal a^yn 
probably as a consequence of long cultivation; for the plant is 
unknown in a wild state, except here and there as an escape 
from cultivation. In 2. Avenastrum, Koch {Helicotriohum, Bess.), 
the plant is perennial, the panicle usually narrow, with erect or 
rarely spreading spikelets with more than two perfect flowers, 



9 8 MR. e. BEKTHAM ON SKAMINEiE. 

and the lower empty glumes with only one or three, or the second 
rarely with five nerves. 

10. GrAUDiNiA, BeauY., two species, has the spikelets of Avena 
{Avenastrum) ; but they are singly sessile in the notches of the 
articulate rhachis of a single spike, thus showing the inflorescence 
of the tribe Hordese, to which Parlatore would remove the genus ; 
but the dorsal twisted awn places it much nearer to Avena, from 
which some authors would not generically separate it. The com" 
mon O.fragilis, Beauv., is widely dispersed over the Mediter- 
ranean region. The second species, Q. geminiflora, J. Gray, was 
proposed as a genus ArthrostaoJiya, Link, from garden specimens of 
unknown origin ; it has since been detected by Seubert in the 
Azores. 

11. AMPHiBUOMrs, Ifees, is a single Australian species, with 
many-iiowered spikelets. The grain is furrowed as in Avena, but 
glabrous and free from the palea as in Trisetim. 

12. AEEHENATnEErM, Bcauv., contains three European, North- 
African, or Oriental species, often included in ^vewa, but differing 
from that genus as well as from most Poacese in having, as in the 
two following genera, the lower flower male and the upper one 
fertile, though the rhachilla is produced beyond it as in other 
Avenese. 

13. Teistachta, Nees (^Monopogon, Presl), has eight species, 
of which two are tropical American, the remainder African, tro- 
pical or southern, one extending to the Levant. "With the lower 
flower male, as in Arrhenalherum, they are readily distinguished 
by the spikelets always three together, sessile or equally pedi- 
cellate at the ends of the branches of the panicle, and by the 
long twisted awn of the flowering glume being terminal between 
two lobes or straight awns. Amongst Nees's African species, 
T. simplex must be transferred to Trichopteryx. 

14. Teiohopteetx, !Nees {Loudetia, Hochst. ), about ten African 
species, of which one is also in Brazil, has the spikelets of Tri- 
stacJiya; but they are scattered along the branches of the panicle, 
not in terminal triplets. The only Brazilian species, not un- 
common also in tropical Africa, T. fiammea, has, as already men- 
tioned, been rather negligently published and figured as an 
ArvAidinella, of which it has none of the characters and not much 
of the habit. 

15. Anisopogon, Br., is a single "West-Australian species, 
differing from Danthonia in the large spikelets containing only a 



ME, G. BENTHAM Olf GEAMINIiiE. 99 

single perfect flower. Nees added a second species from South 
Africa which I have not seen ; but from his description it can 
scarcely be a congener. Kunth has figured three lodicules in 
the Australian plant ; I have always found only two long lanceo- 
late ones. 

16. Danthonia, DC, is now a polymorphous, almost cosmo- 
politan, genus of nearly a hundred species, of which the greater 
number, however, are South-African, all characterized by the 
spikelets containing three or more perfect flowers, and by the 
awn of the flowering glumes more or less twisted or bent and 
usually flattened at the base, but terminal between two or four 
teeth or straight awns. Notwithstauding considerable diversities 
in habit, inflorescence, and in the size and teeth of the glumes, 
no good natural sections have yet been proposed. Nees's Si- 
mantochate {Streblochcete, Hochst.), with the lateral lobes or teeth 
of the flowering glumes entire and acute or awned, and Penta- 
schiste, with the lateral teeth bifid and one or both teeth awned, 
are purely artificial, and relate to the African species, all the non- 
African ones being Included in SimantocJuste. DeCandolle ori- 
ginally proposed the genus for two European species, D. procum- 
lens and D.provincialis ; Brown showed, however, that they could 
not well be regarded as cougeuers, and removed the former to 
his new genus Triodia. The D. provincialis therefore becomes 
the type of the present large genus Danthonui, though it may be 
somewhat anomalous when compared with the majority of the 
African and Australian ones. DeCandolle's chief character con- 
necting his original species was the great length of the outer 
empty 'glumes compared with the rest of the spikelefc ; and this is 
a general, though not quite a universal, feature of the enlarged 
genus. Since Brown's time the following genera have been pro- 
posed, chiefly upon single species, with characters which appear 
to be of little more than specific value : — Pentameris, Beauv., is 
Z>. Thouarsii, Nees, from South Africa, with nearly the habit and 
inflorescence of Z). pallescens, Nees, but remarkable for the short 
thick grain truncate at the top. Trircvphis, Nees (not of E. Br.), 
is B. radicmis, Steud., from South Africa, nearly allied to Z). crispa, 
Nees. GTicBtolromus, Nees, contains a few South- African species, 
in which one, or sometimes two, of the flowers' in the spikelet are 
imperfect. Monacather, Steud., is B. hipartita, P. MuelL, an 
Australian, species, with the fruiting glumes hardened and oblique 
at the base and bearing a ring of hairs under the lobes. Plin- 



100 ME. Q. BENTUAM ON GEAMINBiE. 

tanthus, Steud., is founded on two Australian spacies, moat pro- 
bably of Banthonia, but which, from the evidently incorrect 
character given, it is impossible to identify without seeing the 
specimens. Grinipes, Hochst., is the Abyssinian species pub- 
lished by A. Eichard as D. dbyssinica, Hochst., in which the outer 
empty glumes are exceptionally shorter than the spikelet. 

Tribe XI. CHLOHIDEiE. 

This tribe is characterized amongst Poaceae almost exclusively 
by the inflorescence. The spikelets are sessile in two rows in 
unilateral spikes, the rhachis of which is neither articulate nor 
notched as in Hordeeae ; and the spikes, sometimes solitary and 
terminal, are more frequently several, either digitate at the end 
of, or scattered along, the peduncle or axis of the single panicle. 
The inflorescence is thus nearly that of Paspalum, whilst the 
spikelets are those of Pestuceje, with the lowest or single perfect 
flower hermaphrodite, and the awns, when present, terminal and 
straight, not dorsal or twisted as in Agrosteas and Avenese. The 
following twenty-seven genera are mostly, but not quite all, tro- 
pical or subtropical ; the first fifteen have one fertile and only 
rarely a second male flower in each spikelet ; the next ten have 
two or more fertile flowers ; all, except a few very small genera or 
exceptional species, have the rhachilla continued beyond the 
flowers, and often bearing one or more empty glumes. The last 
three genera enumerated under the tribe are anomalous dioecious 
grasses, connecting Ohloridese with the subtribe Sesleriese of Fes- 
tucese, but showing the inflorescence of the present tribe at least 
in the male individuals. 

1. MiCEOOHiOA, Br., comprises three species, of which two are 
endemic in Africa and the third widely spread over the warmer 
regions of the New as well aa the Old World. They are slender 
tufted grasses with filiform leaves and single slender terminal 
spikes and small awnless one-flowered spikelets without any 
continuation of the rhachilla. 

2. SoHCENEEELDiA, Kunth, is a single tropical-African species 
with one to four erect spikes at the top of the peduncle ; the 
spikelets are one-flowered without any continuation of the rha- 
chilla as in Microehloa, but not so small; and the flowering 
glumes bearing long capillary awns, give the spikes an elegant 
crinite aspect. 

3. Octodon, Pers. {Bactilon, Vill., Oapriola, Adans., FibicUa, 



- MB. a. BENTHAM ON GEAMIITEJE. 101 

Koel.), a small but mixed genus, of which the typical species is 
a common weed in most warm or temperate parts of the civilized 
world. It has the slender spikes and small spikelets oi MierocMoa ; 
but the spikes are several digitate at the end of the panicle, and 
the vhachilla is produced beyond it into a small point or bristle. 
Three Australian species have, however, been added to it with the 
spikelets of MicrocMoa but with the inflorescence of Gynodon, 
thus closely connecting the two genera. Persoon's generic name 
is far from being the oldest, but has been so long and so uni- 
versally adopted, that the substitution of either of the others 
for it would only breed confusion without the slightest advantage. 

4. Haepeohloa, Kunth, has also two South-African species. 
The spike is single, terminal, dense, and unilateral, often falcate ; 
and there are usually one or two male flowers above the fertile 
one, the glumes all uuawned. 

Of 5. Ctenium, Pauz. (Monocera, EIL), we have seven species, 
of which four from North or South America, three from Africa 
or the Mascarene islands. The spikes are solitary or rarely two 
or even three at the end of the peduncle ; the spikelets, though 
elegantly pectinate as in HarpecMoa, have a very diff'erent struc- 
ture : the second empty glume is larger than the others, and bears 
on the back a fine horizontal point sometimes reduced _^to a small 
tubercle ; the third and fourth glumes are small and empty, or 
only enclose a palea ; the fifth or flowering glume ends in a fine 
awn, and above it are one or two empty ones. 

6. Enteeopogon, ISTees, was founded on an East-Indian grass 
with a single long, often incurved terminal spike ; otherwise very 
near ChJoris except in some minor points. It now includes 
Otenitim Seychellarum, Baker, from the Mauritius, which is scarcely 
specifically distinct from the common East-Indian one, E. macro- 
stacJiya, Munro {Ghloris macrosiachya^ Hochst.), from Abyssinia, 
and an unpublished species from Mayotte in Madagascar, Boivin 
n. 3019, which may be thus characterized : — H. leptophylla, Benth., 
foliis angustissimis siccitate convoluto-subulatis, glumsB florentis 
integrsB arista gluma ipsa longiore. The habit and the long 
unilateral spike are precisely those of the common Indian E. me- 
licoides ; but the leaves are very much narrower and scarcely 
flattened in the lower part, the spikelets rather larger, the flower- 
ing glume nearly 3 lines long, and the awn about g inch, and, 
at least in the spikelets examined, the flowering glume is quite 
entire, scarcely free at the point from the awn. 



102 MB. G, BENTHAM ON GEAMINE^. 

7. Chloeis, Sw., contains about forty species, dispersed over 
the warmer regions both of the New and the Old "World. It is 
for the most part a natural genus, with two or more spikes digitate 
at the end of the peduncle, the one-flowered spike-lets in two 
regular close rows as in the allied genera, the flowering glume 
usually awned, and one, two, or more empty glumes above it ; but 
these characters are not constant, and the structure of the spike- 
lets is somewhat polymorphous. C monostaohya, Pourr., from the 
Mascarene islands, and G. unispieea, E. Muell., from Australia, 
are slender plants with only one or rarely two spikes, and the 
flowering glume as well as the upper empty one are narrow and 
awned. In C aciculare, Br., and G. BoxburgJiiana, Edgew., from 
East India and Australia, and C radiata, Sw., from America 
and Africa, the glumes are likewise narrow and awned, or the 
upper empty one reduced to a mere awn, but the spikes are nor- 
mally several and digitate. O.foliosa, Willd., has also a narrow 
awned flowering glume ; but the upper empty one is a double awn 
(or two awnlike glumes), and the spikes are not so closely clus- 
tered at the end of the peduncle, on which account Doell has 
transferred the species to Oymnopogon, from which it appears to 
me to be much further removed. In O. pumilio, Br., G.pecUnata, 
Benth., and C divaricata, Br., all from Australia, the flowering 
glume is distinctly two-lobed with the awn between the lobes. 
In a considerable number of species the upper empty glumes are 
broad and truncate at the top — these empty glumes being several 
in each spikelet in the Asiatic and Australian species, but one 
only in the American ones. In all the preceding species the 
flowering and upper glumes are awned ; in five or six American 
or African species forming the proposed genus lEtistachys, Desv. 
{Schultesia, Spreng.), botli the flowering and the upper empty 
glume are obtuse and truncate, but without any awn, or only a 
minute point. They are, however, closely connected with the 
typical species of Chloris through C submutica, Eunth. G. villosa, 
Pars., and G. macrantlta, Jaub. et Spach, both of them described 
in detail and figured by Jaubert and Spach, must be transferred to 
Tetrapogon, as having their spikelets with at least two fertile flowers. 

8. Teichlorts, Eourn., comprises two Mexican and two extra- 
tropical South- American species. They resemble Trisetaria in 
their dense oblong crinite panicle and their three-awned flowering 
glumes ; but the panicle is composed of simple crowded or verti- 
cillate spikes, and the spikelets, sessile in two rows on the rhachis 



ME. a. BENTHAM ON GBAMINE^. 103 

with one to three empty awued glumes above the floweriug one, 
are quite those of Ghloris. The two southern species had long 
been indicated and named in herbaria as constituting an inde- 
pendent genus (the one by J. G-ay, the other by Munro) ; but 
never having been published, we must adopt Fournier's generic 
name for the whole. The two southern species (from Chili, 
Tucuman, and Buenos Ayres) are indeed so very near T. fasci- 
culttta, Fouru., that it will require close investigation to estab- 
lish their specific differences. Another different-looking plant 
from Tucuman (Tweedie), much smaller, with a loose inflores- 
cence and short-awned spikelets, shows also the essential cha- 
racters of Trichloris. 

9. Gtmnopogon, Beauv. (Anthopoffon, Nutt.), differs from all 
the preceding one-flowered genera in the spikelets not closely 
crowded, but more or less distant along the slender rhachis of the 
spikes, althoiigh still sessile in two rows and unilateral ; the spikes 
themselves are scattered or verticillate along the common pe- 
duncle. There are four or five American species, northern or 
southern, and one from Ceylon, G. rigidus, Thw., forming Nees's 
genus Dichcetaria, but only differing from the American ones in 
the spikes fewer in the panicle, and the spikelets rather larger with 
longer awns. Doell's G. foliosm and G. pullulans should be 
restored to Ghloris, with which they agree in every respect except 
that the spikes are not quite so closely clustered at the end of 
the peduncle. 

10. MoNoCHjsTE, Doell, a single Brazilian species of which I 
have seen no specimeu, is removed by Doell from Gymnopogon, 
where Martius had placed it, as having no continuation of the 
rhachilla beyond the flower. Nees, however, describes a bristle- 
like continuation, but not bearing any empty glume or awn 
as in Gymnopogon. The genus is as yet, therefore, in some 
measure doubtful. 

11. ScnEnoNNAKDUs, Steud., is the. North- A.merican Leptu- 
rus paniculatus, Nutt., wliich, however, Steudel failed to recog- 
nize. Nuttall indicated its aflS^nity to Gymnopogon, and evidently 
only placed it in Lepturus from not knowing the latter genus 
except from the imperfect characters then published. ScJiedon- 
nardus has now been figured in the last part of Hooker's Icones ; 
the description, accidentally omitted in printing, will appear in 
the next part. 

12. Ceaspedoehachis, Benth., is a single species from east 
tropical Africa, allied to Schedonmrdtts, but differing in the 



104 ME. a. BENTHAM OK GKAMINEJi!. 

flexuose rhachis of the spikes bordered by a narrow merabraue, 
in the flowering glume and palea vey small and thin, ressmbling 
lodicules, and a few other minor points. It is being figured for 
the forthcoming part of Hooker's loones. 

13. BouTELOUA, Lag. (EiUriana, Tela., Actlnochloa, Willd.), 
comprises about twenty-five American species, northern or 
southern, but chiefly western. As ia the four preceding genera, 
the spikes are distant along the main peduncle, and often nume- 
rous, very rarely reduced to one or two ; but they are usually 
short, with the spikelets densely crowded in two rows on one side 
of the rhachis, and the rhachiUa always continued beyond the 
single hermaphrodite flower, bearing one to three empty glumes 
or awns, or sometimes a male flower. The flowering and upper 
empty glumes usually end in three or five lobes, points, or awns ; 
but they are often exceedingly variable in this respect even in 
the same specimen, and it becomes difiicult to make much use 
of them in the arrangement of the species. The following four 
sections, raised by some to the rank of genera, are founded chiefly 
on inflorescence :—(l) Ghondrosia or Ghondrosium, Desv. Spikes 
usually few, often rather long, with numerous spikelets (more 
than twelve) neatly pectinate, and the terminal empty glume 
usually three-awned ; the species rather numerous, especially in 
Mexico, where they run much one into another. (2) Atheropo- 
gon, Muehl., including JSeterostega or Seterosteca, Desv. Spikes 
often numerous, but usually very short with few (rarely above 
twelve) spikelets, crowded but scarcely pectinate, or almost re- 
duced to clusters, the terminal empty glume varying from three- 
awned to entire, or reduced to a single bristle. The species best 
known, B. racemosa, Lag. {Atlieropogon apludoides, Muehl., 
Dinebra curtipendula, DC), was associated by DeCandolle and 
Beauvois with the Dinehra arahica of Jacquin, which, however, 
diflfers essentially in il;s several-flowered spikelets. (3) Tria- 
thera, Desv. Spikes still further reduced than in Atheropogon, 
consisting usually of two to four spikelets so narrow and so close 
together as to appear like a single one, and perhaps sometimes 
really only a single one, the upper empty glume reduced to three 
awns, as in several species of the preceding section. There appear 
to be two species, B. arktidoides {Dinehra aristidoides, H. B. K., 
forming the genus Aristidium, Eudl.), with two to four spikelets 
to each spike, and B. triathera, to which I should, with Munro, 
refer both Triathera, Desv., and Tricena, H. B. K. The spike- 



MB. G. BENfTHAM ON OEAMINEiE. 105 

lets vary in the same panicle, one, two, or three to the spike, 
and are themselves polymorphous. Where there are three, I 
have found the lowest empty glume of the lowest spikelet very 
narrow and awnlike, and very probably that which Kunth has 
described and figured as an awn at the base of the glume ; in 
the uppermost of the three spikelets the lowest empty glume is 
similar to the second, the intermediate spikelet being sometimes 
like the upper, sometimes like the lower one. (4) Polyodon, 
H. B. K. {TriplatJiera, Endl.). Spikes few, short, and crowded at 
the. end of the peduncle with few spikelets, the flowering glume 
three-awned, the two or three upper empty glumes each with 
three or five awns, having together the appearance of a single 
cluster of many awns. We have two species, B. distieha {Poly- 
podon distichum, H. B. K.), including apparently AtTteropogon 
affinis, l^ourn. (not JEutriana qffinis, Hook, f.), with the spikelets 
including the awns under half an inch long, and B. muUiseta (JEu- 
triav.a timltiseta, Nees), with the spikelets and their awns above 
an inch and a half long, giving the plant the aspect oi Boissiera. 
Corethrum, Vahl, is probably the same plant. He received his 
specimen in a collection of Syrian plants sent him by Thouin, 
into which it had probably got misplaced by some carelessness in 
sorting. No special locality is ascribed to it nor any indication 
of the collector; and no plant answering to Yahl's elaborate and 
probably accurate description is known from Syria or any part of 
the Levant. 

14. Melanocencheis, Nees, comprises three species from 
East India or tropical Africa, closely resembling each other, and 
at first sight having the aspect almost of JEgopogon but the 
characters are very nearly those of Bouieloua {AtTieropogon) ; 
the genus is readily distinguished from both by the linear plumose 
empty glumes. 

The preceding genera have all only a single flower in the 
spikelet, the second, when present, being male only ; in the follow- 
ing ten genera there are at least two, and often several more 
fertile flowers. 

15. Teipogon, Eoth {Plagiolytrum, Nees), contains about eight 
East-Indian and tropical-African species, with the single elon- 
gated terminal spike of Enteropogon, but with several-flowered 
spikelets, and the flowering glumes more or less three-awned as 
in Triehloris, TrirapMs, etc., the lateral awns sometimes reduced 
almost to teeth. 



106 MB. G. BESTHAM on GRAMlKE^. 

16. Lepibopyeonia, A. Eich,, is a single Abyssinian species 
which I have not seen, and of which the specimen described is 
said to have been imperfect. .From the figure and description it 
would appear to be allied to the Tripogon cibyssinicus, Nees, but 
with broad, very villous flowering glumes, and the single awn not 
quite terminal. 

17. Tetraposon, Desf., lias four Abyssinian, North-African, 
or "West-Asiatic species, including the above-mentioned Chloris 
villosa, Pers., and O. macrantha of Jaubert and Spach. They have 
one, two, or rarely three terminal erect spikes, resembling Elio- 
nurus in the long silky hairs which cover them, but with the cha- 
racters of Chlorideee, differing from Chloris itself in their several- 
flowered spikelets. 

18. AsTEEBiiA, p. Muell., comprises two or three Australian 
species formerly referred to Dantlionia, from which the habit and 
untwisted awn separate them. In the ' Elora Australiensis ' I 
placed them near PappophoresB on account of their many-nerved 
glumes ; but the inflorescence places them in Chlorideaj, where 
they come in many respects near to Tetrapogon. 

19. Wangekheimia, Moench {Oynosurus Lima, Linn.), from 
Spain and North Africa ; 20. Ctenopsis, DeNotar. {Festuca 
pectinella, Delile), from North Africa ; and 21. Teieaohne, 
Nees, from South Africa, are all single species hitherto referred 
toEestuceffi; but they have all the one-sided spikes with the 
spikelets sessile in two rows of Chloridefe, to which, following out 
Munro's memoranda, I have transferred them. The spikes are 
solitary, erect, and often slightly falcate in Wangenheimia and 
Ctenopsis, several scattered along the common peduncle in 
Tetrachne. 

22. DiJTEBEA, Jacq., remains limited to the original African 
and East-Indian B. arahica figured by Jacquin. DeCandolle, as 
above mentioned, joined it with the section Aiheropogon of 
Bouteloua, of which it has the habit ; but there are always two 
fertile flowers to the spikelet. Kunth referred it to Leptochloa, 
from which it is further removed by its short dense awned spikes ; 
and from both it is separated by the lower empty glumes as long 
as, or longer than, the rest of the spikelet. 

23. Eleusine, Gsertn., taken in the sense given to it by Per- 
soon, is a natural genus of about seven species, from the tropical 
and subtropical regions of the Old World, two of them common 
weeds also in America. The spikes are usually several, digitate 



Mft. 0. BENItHAM OS flEAMINEJE. W? 

at the end of the peduncle, and in some species with the addition 
of others scattered or yerticillate lower down. The flat spikelets 
have been sometimes mistaken for those oi Eragrostis ; but their 
arrangement in two rows is always that of Ohlorideae. The genus 
is often restricted to Gsertner's JS. coracana and E. indica, in 
which the spikes are digitated and rather long, and the membra- 
nous pericarp loose on the ripe seed. This character is particu- 
larly marked in the E. coracana ; but that is probably a plant 
somewhat modified by long cultivation. In the common E. indica 
the pericarp is often as loose, but sometimes remains very thin 
and not so easily detached. In the still more common E. eegyp- 
tiaca, Pers., forming the genus Bactyloctenitm, "Willd., the digi- 
tate spikes are very short and dense, and the very thin pericarp 
appears to wither away or to dry up in ripening, leaving the seed 
apparently exposed and rugose, similar to that of E. indica. In 
E. hrevifolia, Wall, Oat., and E. glaucopTiylla, Munro {Bactylo- 
ctenium glaueopTiyllum, Courb.), the spikes are short as in E. 
(Bgyptiaca, and more or less of the remains of the membranous 
pericarp maybe often seen persistent about the seed. Acraohie, 
Wight and Am., is the E. verticillata, B-oxb., in which the spikes 
are rather long as in E. indica ; but besides the terminal digitate 
ones, there are others scattered or verticillate along the peduncle. 
ArthrochlcBHa, Boiv. in herb. J. Gay, is a remarkable Madagascar 
species which may be thus defined : — E. macrostachya, Benth., 
elata, foliis angustissimis rigidis crassiusculis, spicis 2-3nis ter- 
minalibus, spiculis confertis 18-20-floris, glumis acute cariuatis 
paleaceis bifariam imbricatis. A plant of rushlike habit about 
2 feet high, the spikes about 4 inches long, with very numerous 
spikelets varying from 4 to 6 lines long, resembling those of Era- 
grostis, but much more rigid. 

24. Leptochioa, Beauv. (Oxydenia, Nutt.), about twelve spe- 
cies, tropical or subtropical, in the New as well as the Old World, 
and extending on the one hand into North America, and on the 
other into extratropical Australia, is one of those genera which 
interfere provokingly with our classifications. Nearest allied to 
Eletcsine, ithas also considerable affinity with Gynodon, Biplachie, 
and Poa, to which some of the species have been occasionally 
referred ; and one has beeu figured as a Gynosurus. The chief 
character consists in the slender spikes scattered along the 
common peduncle, with numerous small flat spikelets, giving the 



108 Me. G. BENtHAli ON OlUMlNEji. 

plants a different habit from that of the several preceding genera. 
The species may be distributed into two rather distilict sections. 
(1) Pseuclocynoclon, with only one or two flowers in the spikelet, dif- 
fering hut little from Gynodon except in inflorescence. To this sec- 
tion heloiigL.uniJlora, Tloch.sb.(Ci/nodonffraoilis, Nees), from Abys- 
sinia, Ii. Neesii {Gynodon Neesii, Tlnv.), from Ceylon, and L. poly- 
stachya, Beuth. {Qynodon polystachyus, Br.), from Australia. (2) 
Euleptochloa, with two or more flowers to the spikelet, comprises 
the remainder of the species. Tliose which have a point or short 
awn to the flowering glume were formerly generically separated 
by Beauvois under the name of RabdooMoa. Amongst the pub- 
lished species, L. arabica, Kunth, is the genus Dlnelra ; L. Lind- 
leyana and L. mollis, Kuuth, are referable to Triodia. L. duhia, 
Nees {Chloris duhia, H. B. K.), and the North-American L. (Di- 
plachne) fascicularis , A. Gray, appear both to be true species of 
Diplachne. Lorentz's South-American plant disti-ibuted by G-ri- 
sebach as L. fascicularis appears to be a mere variety of Panicmn 
sanguinale. L. Wightiana, Noes, is an JSragrostis. L. plu- 
mosa, Anders., is a Triodia. 

There remain three anomalous monotypic genera from the 
Mexican-Texan region — 25. BucnLOE (Sesleria, afterwards Oalan- 
thera, Nutt.), 26. Jouvea, Fourn., and 27. Opizia, Presl, which 
connect in some measure Chloridese with the subtribe Sesleriea', 
and are all dioecious, and very remarkable for the great dissimi- 
larity in the spikes and spikelets of the two sexes. Of the first, 
numerous specimens fully confirm Engelmann's excellent jigures 
and description ; the other two are unknown to me, and remain 
somewhat doubtful. Of Jouvea, Pournier only knew the female, 
which he says is allied to BucMoc, with a very different habit. 
Opizia is said by him to have the male plant exactly like Buchloe ; 
the female figured by Presl must be very different, though his 
and Pournier's descriptions do not agree in all points. ' 

Tribe XII, EESTtrcEiE. 

The large tribe Festticea presents considerable difficulties to 
the systematist. Of the seventy genera on our list (about a 
hundred and ten of some botanists), the greater number are perhaps 
better defined than those of Agrostese for instance, and afibrd a 
much greater variety of characters ; but none of the various 
arrangements' proposed for distributing them into groups or sub- 
tribes have proved satisfactory, and the two largest genera Poa 



ME. a. BENTHAM OV GBAMINE^. 109 

and Festuca are connected by a number of smaller ones, wbich. 
are more variously associated together or separated by European 
botanists than almost any others of the Order. As a whole, 
Testucese should include all the Poacess with two or more perfect 
flowers to the spikelet, which have neither the peculiar inflores- 
cence of Chloridese or of Hordeese, nor the dorsal or twisted awn of 
AvenesB, nor the peculiar habit of Bambusese. But we have seen 
that there are a few species where the awn is wanting, but which 
must yet be left in Avenege ; we shall find that in Diplachne, Oreo- 
cJiloa, and even in Festuca itself, there is occasionally an inflores- 
cence very nearly that of Chlorideae ; and with regard to Bambusese, 
distinct as is the habit and foliage of the great mass of genera, yet 
it is exceptional in Planotia, and in the subtribe Centotheceae of 
Festucese there is some approach to that of the Bambusese. The 
subdivisions proposed of the Order into subordinate groups 
have been so various, and often on such plausible (though some- 
times contradictory) grounds, that it is not without hesitation 
that I have selected for adoption the following eight subtribes : — 
Subtribe 1. Pa^popTiorecB, has often been raised to the rank of 
a substantive tribe, but with various limits ; and it really is only 
distinguished from Triodiese by the more nnmerous teeth, lobes, 
or awns of the flowering glumes. There are- five well-established 
genera, requiring little comment. 

1. PoMMEHErLLA, Linn, f., is a single East-Indian annual, 
with short spikes almost enclosed in the upper leaf-sheaths, and 
remarkable for the presence of two empty glumes between the 
ordinary lower pair and the flowering ones, as in Ctenium, Fre- 
mochloa, BrylMnia, and Uniola. 

2. Pappophoeitm, Schreb., has nearly twenty species from the 
warmer regions of both the New and the Old "World, distributed 
in two sections, regarded by some as distinct genera : — Poly- 
rhaphis, Trin., a few American species in which the flowering 
glume has thirteen to twenty-three very unequal awns ; and 
Frmewpogon, Desv., chiefly, but not exclusively, from the Old 
"World, in which the flowering glume has nine awns, all nearly 
equal, or five of them rather external and slightly different from 
the four inner ones. 

3. CoTTEA, Kunth, is a single tropical American species, 
differing from PappopJiorum in the looser panicle, and in the 
flowers usually more than two, instead of only one or two, ia 
each spikelet. 

LINH. JOUEN. — BOTANT, VOL. XIX. K 



110 ME. G. BENTHAM ON GBAMINE^. 

4. BoissiEHA, Hoclist., is a single Oriental species, which, 
besides the characters derived from the glumes, has a very short 
membranous dilated two-lobed style, different from that of every 
other known grass. 

5. ScHMiDTiA, Steud., contains two species closely resembling 
each other, one from thB Cape-Verd Islands, the other from 
tropical and South Africa, with a narrow but loose panicle, and 
the flowering glumes ending in four inner lanceolate lobes, and 
live outer subulate lobes or awns. 

Our second subtribe, Triodiem, is not so definite as could be 
wished. There are usually more than two flowers to the spike- 
let ; and the flowering glumes have rarely more than three nerves, 
and end in three teeth, lobes, or awns. These characters are gene- 
rally very prominent ; but in a few species of Triodia and Dipl- 
achne the teeth are scarcely more than what occur occasionally 
in some other subtribes, and in one or two species of Triraphis 
the nerves of the glumes are more numerous, bringing them 
technically near to Pappophorese. The panicle in all the genera 
is usually narrow, dense or loose, but very rarely spike-like, and 
in a few species loose and spreading. "We include six genera. 

6. Tbiodia, Br. (Uralepis, Nutt.), as at present limited, com- 
prises about twenty, extratropical species, northern or southern, 
with a very few extending into the tropics in America or Africa. 
It has the typical characters of the tribe without the peculiarities 
of the other genera, the lobes of the flowering glumes reduced to 
short teeth or points, or the central one rarely lengthened into a 
short awn. It must be admitted, however, that it is still both a 
vague and a polymorphous genus, comprising three rather dis- 
tinct sections and a few anomalous isolated species : — 1. Isotria, 
three Australian species (J. MitchelU, Benth., I. pungens, Br., 
and I. Ounninghamii, Benth.) with the three lobes of the flower- 
ing glumes narrow lanceolate and equal ; 2. Uralepis {Sieglingia, 
Bernh., Merisacline, Trin.), about sixteen American or Australian 
species with one European one, in which the lateral teeth of the 
glumes are broad and not pointed and sometimes very minute, 
the middle one a point or very short awn ; 3. Tricuspis, Beauv. 
(Windsoria, Nutt.), three North- American species differing from 
Uralepis in the nerves of the lateral teeth produced into short 
points. Besides these, Leptocarydion, Hochst., is an Abyssinian 
species, T. plumosa {LeptocTiloa plumosa, Anders., Diplachne alo- 
pecuroides, Hochst.) with the dense soft panicle almost of Tri- 



ME. a. BENTHAM ON GEAMINEiE. Ill 

cMoris, but with the spikelets of Triodia. Triehoneura, Anders., 
is Leptochloa Lindleyana, Kunth, from the Galapagos Islands, 
with the habit nearly of Diplaahne fasoicularis, but with the 
characters of Triodia. Leptochloa mollis, Kunth, from Senegal, 
which I have not seen, would appear from his figure to be a 
Triodia, near to the T. plumosa, but with the loose panicle of 
T. Lindleyana. Rhombolytrum, Link, is T. filiformis, Nees, a 
Chilian species, very near to the North-American T. albescens, 
and T. trinerviglumis, Munro, and also to T. KerguelensiSj 
Hook, f., and T. antarctica (Gatahrosa antarctica, Hook, f.), from 
extreme southern America. 

7. DiPLACHNE, Beauv., now comprises about fourteen species, 
dispersed over the tropical and temperate regions both of the 
New and the Old World, and variously referred to Triodia, 
Leptochloa, or Molinia by different agrostologists ; and the genus 
is really closely connected with the two first-named, but more 
especially with the Triodice of the. typical section Uralepis. 
Prom these it chiefly differs in inflorescence : the branches of 
the panicle are long and slender ; the spikelets, almost linear, 
scattered along the rhachis and sometimes sessile or nearly so 
in two rows, but not regular and unilateral enough to place the 
genus in Chloridese, to which it is sometimes referred. The cha- 
racteristic teeth of the flowering glumes are also sometimes very 
minute. Whatever position, therefore, we give to the genus, it 
must be more or less an arbitrary one ; but that next to Triodia 
seems the least objectionable. 

8. Teiplasis, Beauv. {Biplocea, Eafin.), has two North-Ameri- 
can species (Uralepis cornuta, Ell., and TI. purpurea, Nutt.), with 
a narrow, slender, slightly-branched panicle, and the flowering 
glumes deeply divided into three narrow lobes, the central one a 
slender awn. The South-American Triplasis setacea, Gtviseb., is 
a Biplachne (D. spicata, Doell). 

9. ScLEEOPOaoN, Philippi {Lesourdia, Fourn.), has four species, 
one from Chile, the others from the Mexican-Texan region, all 
remarkable for the unisexual spikelets, those of the two sexes so 
different in- aspect that without positive evidence it would have 
been difficult to suppose th§m to belong to the same plant. The 
Mexican ones have been very well described and one of them 
figured by Fournier, who, from his specimens, supposed them to 
be strictly dioecious ; but we have specimens with the two in- 
florescences upon different branches of the same individual. 

k2 



112 ME. O. BENTHAM OIT GBAMINEJB. 

Fournier also was not aware of the Chilian species previously 
published by Philippi, whose generic name necessarily takes the 
precedence over Fournier's. 

10. Eeemochloa, S. Wats., contains two New-Mexican or 
Californian species, well described and illustrated by S. Watson. 
The genus is exceptional in the subtribe in its one-flowered 
spikelets, and in the two empty three-awned glumes between 
the lower pair and the flowering one. 

11. Teieaphis, Br., comprises five or six species, one as yet 
unpublished from South Africa, the others from Australia. In 
all, the three lobes of the flowering glumes are awned, whilst in 
one species, T. mollis, Br., otherwise inseparable from the genus, 
there are additional small membranous lobes between the awns, 
and at least five nerves to the glume, showing a close connection 
with the Pappophoreae. T. microdon, Benth., from Australia, is 
a very anomalous plant with unawned glumes, which should be 
removed from the genus ; but it is diiflcult to say where it should be 
placed T. capensis, Nees, is a Danihonia {D. radicans, Stead.). 

The third subtribe, Arundinece, though often regarded as a 
substantive tribe, has no definite character beyond the tall habit 
with a rich panicle, as designated by the common name Meed, 
and the long hairs surrounding the flowering glumes, either 
arising from the rhachilla or from the glumes themselves. This 
character, however, is no more exclusive here than in Saccharese ; 
for there are other genera, such as Oalamagrostis, Qraphephorum, 
&c., which have the hairs nearly as long, but which on other 
accounts cannot be included in Arundinese. The genera certainly 
belonging to the subtribe are four : — 12. Gyneeittm, Humb. 
and Bonpl., three tropical or subtropical American species with 
strictly dioecious spikelets. 13. Ampelodbsmos, Beauv., two 
Mediterranean species with rigid five-nerved flowering glumes. 
14. AEirNDO, Linn. {Donax, Beauv., Scolochloa, Mert. and Koch, 
Amphidonax, Nees), three or four species from the warmer 
regions of the New and the Old "World, very abundant all round 
the Mediterranean, with three-nerved flowering glumes ; and 15. 
Pheagiiites, Trin. {Arundo, Beauv., Czernya, Presl, Trichodon, 
B-oth), two species, one of them almost cosmopolitan, only differ- 
ing from Arimdo in the lowest flower of the spikelet being male 
only. These last three genera are frequently, and perhaps with 
reason, regarded as sections only of Arundo. Two Mexican 
monotypic genera proposed by Fournier, 16. Gotjinta, and 17. 



ME. G. BENTHAM ON GEAMIMEiE. 113 

Calamochloa, are unknown to me. Tliey are placed by the 
author in ArundinejB ; but both would appear, from the short 
character given, to have a rather different iiiflorescence. 

We have formed our fourth subtribe Sesleriece of ten genera, 
all except the monotypic Elytropliorus extratropical, and con- 
nected together for the most part by inflorescence as well as by 
structural characters. These are, however, not without excep- 
tions. The spikelets are usually collected together in little heads 
or close clusters, which are themselves closely clustered in a dense 
globular or spike-like panicle ; and at the base of the head or 
panicle are usually a few barren spikelets or empty glumes, or, 
sometimes single glumes, which I have elsewhere, though as I 
now believe erroneously, designated as bracts subtending the 
branches of the inflorescence. In two genera the whole inflo- 
rescence is reduced to two or three spikelets sessile in a cluster 
of floral leaves. In structure the flowering glumes are variable 
in their nerves ; but the styles, in almost all the genera, are long, 
with barbellate or very shortly plumose stigmatic branches, 
forming an exception to the tribe, which has induced several 
botanists to refer Sesleria itself to the Panicese, from which it 
differs so widely in other characters. "We include the following 
genera in the subtribe : — 

18. MoNANTHOCHLOE, Engelm., is a single Texan species, ano- 
malous in habit and character, well described and figured by 
Engelmann. It has been compared to Bitchloe on account of its 
unisexual spikelets and creeping habit ; but the two sexes in 
Monanthochloe are very similar to each other, and there is no 
indication in the inflorescence of any affinity with Chlorideae. 

19. MrrsEOA, Torr., has now three or four species, two or three 
from extratropical South America having been recently added to 
the original Mexican-Texan one, figured in the last part of Hooker's 
Icones. The genus is a perfectly isolated one, showing only 
some slight affinity with Monanthochloe, especially in the very 
few spikelets sessile within a cluster of floral leaves ; they are not, 
however, unisexual as in that genus. 

20. EoHiNAEiA, Desf. (Panicastrella, Moench), a single weU- 
known Mediterranean species ; 21. Ammochloa, Boiss. {Cephalo- 
chloa, Coss. & Dur.), two Oriental or North-African species ; 
and 22. Ueochljsna, Nees, a single South- African species figured 
in the last part of Hooker's Icones, require no further comment 
on the present occasion. 



114 ME. a. BENTHAM OK GEAMINEJi). 

23. Sesleeia, Scop., about eiglit European or "West-Asiatic 
species, is an old-establisted distinct genus whicli has been but 
little interfered witli, except that S. tenella, Host, has been pro- 
posed as a genus by Link under the name of Fsilathera, but 
upon characters which do not appear to be of more than specific 
value. 

24. Elxteophghtts, Beauv. {Ilchinolysium, Trin.), is a single 
exceptionally tropical species, widely spread, but limited to the 
Old World. The little heads of minute spikelets forming a spike 
longer than usual in the subtribe, and often interrupted, and the 
wings on one or both the keels of the palea readily distinguish 
the genus. 

25. Fingeehuthia, Nees, a single species figured in the last 
part of Hooker's Icones, is exceptional in the whole primary 
division Poaceae in having the very short pedicels articulate 
below the spikelet as in Panicaces3, whilst the male flower or 
empty glume is above the fertile flower as in Poacese. Its geo- 
graphical range is also peculiar : rather common in South Africa, 
it has been recently gathered by Dr. Aitchison in Afghanistan, 
without its ever having been observed in any intermediate station. 

All the preceding genera have the long styles of the subtribe ; 
but there remain two with the styles scarcely longer than in the 
other Eestucete, whilst the barren spikelets at the base of the 
dense inflorescence or of its branches are very conspicuous, and 
show a close affinity with the Seslerie83. These are 26. La- 
MAECKIA, Moench {Chrysurus, Pers., Pterium, Desv., Tincea, 
Garzia), a single Mediterranean species, and 27. OxNOSTJErs, 
Linn., three or four species with a much wider geographical 
range over the temperate regions of the Old World, and one of 
them at least now naturalized in several other countries within or 
without the tropics. Both genera are remarkable for the lower 
barren spikelets of the clusters or spikes elegantly pinnate with 
numerous bifarious empty glumes. The two were united by the 
older botanists, and have been again brought together by some 
modern ones under the Linnean name Cynosurus ; but they appear 
to be sufficiently different in habit and character to be maintained 
as separate genera ; and Gynosurus itself has two very distinct sec- 
tions, raised by some to the rank of genera : — Cynosurus proper 
for the C. cristatus, Linn., and its annual Algerine variety O. 
polyhracteatus, Poir., altered to C muliibracteatus, Eoem. & Schult. 
(C crista-galli, Munby), in which the spike-like unilateral panicle 



ME. a. TBENTHAM ON GBAMIlTEJi!. 115 

much resembles the spikes of Chloridese, and the glumes, though 
very pointed, are unawned ; and Falona, Adans., altered by Du- 
mortier to Phalona, for the C echinatus. Linn., and O. elegans, 
Desf , with the panicle or head more like that of a Bactylis, but 
with awned glumes. 

The characters of our fifth subtribe, Eragrostece, like those of 
Eufestuceee, are chiefly negative. The two together comprise all 
the FeatuceaB which have not the peculiarities of either of the 
other six subtribes, and differ from each other in the Eragrostese 
having three prominent nerves to the flowering glumes, the Eufes- 
tucess five or more nerves, sometimes rather obscure. Trifling as 
this character may be, it is a fairly constant one, the exceptional 
species being exceedingly rare ; and I have found no other one 
so useful in distributing these numerous genera into groups. 
We have included in EragrostesB twelve of them, though the last 
of them {JEctrosia) might be equally well placed under the fol- 
lowing subtribe Melicese. 

28. KcELEEiA, Pers., has about twelve species, of which ten 
are European, temperate Asiatic, or North African, one of them 
extending over extratropical America north and south and South 
Africa ; the two remaining species are endemic, one in South 
Africa, the other in the Sandwich Islands. The genus has been 
generally admitted with little variation ; but it is difficult to assign 
to it any positive character. The panicle is usually dense and 
narrow, often spike-like; and the glumes are more scarious, espe- 
cially on the margins, and have fainter nerves than in the others 
of the subtribe. It has been divided into two sections, maintained 
by some as genera : — 1. Airochloa, Link, to which Eeichenbach 
restricts the name of Koeleria, with the glumes obtuse or acute 
but without distinct points; and 2. Lophochloa, Eeichenb., to 
which Link restricts the name of Koeleria {JSgialitis, Trin., 
altered by Schultes to ^gialina), in which the flowering glumes 
have a distinct point or short awn at or just below the tip. This 
section includes, besides the species enumerated by Cosson and 
Durieu in their ' Flore d'Algerie,' K. Gerardi, Munro,from South 
Africa, and K. glomerata, Kunth {K. vestita, Nees), from the 
Sandwich Islands. The last species differs slightly from the 
genus in the long loose panicle, which, however, is more dense in 
our specimens than it is figured by Kunth. C. Koch's genus 
Wilhelmsia, from the Caucasus, is, according to Grisebach, only 
a depauperated specimen of K. phleoides, Pers. 



116 ME. (J. BEHTHAM ON GEAMIKE^. 

29. Atellinia, Parlat., is a single "West -Mediterranean annual 
with the hahit of SMsmus, and placed by Savi in Bromus, by 
Grussone in Avena, and by DeCaudolle in Koeleria. It is well 
marked amongst Eragrosteee by the outer glumes, of which the 
lowest is almost reduced to a bristle, and the second broad, mem- 
branous, and the largest of the spikelet ; the flowering glumes 
awned. 

30. Eatonia, Eafin. {Rehoulea, Kunth, Colohanthus, Trin.), 
two or three closely allied North -American species, with the 
second empty glume the largest of the spikelet, as in Avellinia ; 
but the habit is very different and the glumes all unawned. 

31. DissANTHELiTjir, Trin. (Phalaridium, Nees, Stenochloa, 
Nutt.), comprises two, or perhaps three, species from the Andes 
of South America and the coasts of Mexico and California, 
figured and described in the last part of Hooker's leones. They 
have most of the characters oi Schismus, but, besides the widely 
distant geographical stations, they differ in the nerves of the 
flowering glumes always three, not five. 

32. MoLiNiA, Moench {Unodium, G-audin), a single well-known 
European and temperate Asiatic species, and 83. Sphenopus, 
Trin., a very pretty little Mediterranean annual, require no fur- 
ther comment on the present occasion. 

34. Catabeosa, Beauv., can only be distinctly characterized if 
reduced to the single C. aquatica, Beauv., placed by some authors 
in Aira, by others in Glyceria. In it the three nerves of the 
flowering glume characteristic of Eragrostese are very promi- 
nent. The two or three Oriental species added to it by Trinius 
belong to the genus Coi^odium. 0. antarctiea, Hook, f., is a Triodiq,. 
C. glaucescens, Phiiippi, and C. magellcmica, Hook, f., are true 
Glyceria. 

35. Eeageostis, Beauv., an almost cosmopolitan genus of 
above eighty species (multiplied by Steudel and others to about 
two hundred and fifty), is a very natural one so far as the great 
majority of species are concerned, and distinctly limited if we 
include the three-nerved glumes amongst the essential characters. 
Yet in other respects there are exceptional species which have 
been variously referred, even by modern botanists, to Poa, Fes- 
tuca, Priza, Dactylis, Eleusine, or Leptochloa, which they in some 
measure approach respectively ; and some have been proposed as 
substantive genera ; but it has appeared to me that the genus 
may be best defined if retained entire, dividing it into the six 



MR. a. EBNTHAM OIT GBAMIITEJE. 117 

following fairly distinct sections : — 1. Cataclastos, Doell, includes 
J?, ciliaris, Link, ^. peruviana, Trin., and a few other tropical 
species, which, on account of their short spikelets with few 
flowers and fragile rhachilla, have been restored to Poa by Four- 
nier and some others ; but the shape and nervation of the glumes 
are quite those of Mragrostis, and the inflorescence, though pecu- 
liar, is not more that of Foa than of JSragrostis. Macrohlepharus 
of Philippi, judging from his description, does not seem to differ 
from the true H. ciliaris. 2. PlagiostaoJtya comprises some 
African and East-Indian species, which, with the flat several- 
flowered spikelets and continuous rhachilla of Uragrostis proper, 
have an inflorescence approaching that of Chlorideae. The species 
are rather dissimilar in habit. ^. iifaria, Steud., has the long 
simple terminal spike nearly of Tripogon, with obtuse glumes. 
E. SoMmperi {Sarpachne Schimperi, Hochst.) has a shorter simple 
spike and acuminate flowering glumes. I], hrevifolia and E. 
CoelacJiyrum, Benth., the latter forming the genus Goelachryum, 
Nees, and figured in the last part of Hooker's Icones, have 
nearly the habit of some species of Eleusine (Dactyloctenium) or 
of .^luropus. jE. congesta, Oliv., and H. cynosuroides, Eoem. et 
Schultes, have very numerous short sessile spikelets crowded or 
clustered along the long terminal peduncle. 3. Myriostachya is 
an East-Indian species, E. Wightiana {LeptocMoa Wightiana, 
Nees), allied to H. cynosuroides, but with a more complicated in- 
florescence. 4. Fteroessa, Doell, or Eragrostis proper, is charac- 
terized by the usually many -flowered spikelets with the rhachilla 
continuous or rarely articulate when old, the flowering glumes 
usually deciduous, leaving the palea persistent on the minute 
floral axis with its back to the rhachilla. The species are nume- 
rous, and may be distributed into three rather distinct series : — 
CylindrostachyoB, three or four Australasian species, with narrow- 
linear almost terete spikelets ; Leptostachyce, including the cos- 
mopolitan E. pilosa, TBeauv., and its allies, with narrow-linear flat 
spikelets ;, and Megastachyce, including the widely-spread E. mega- 
stachya. Link, and many other, chiefly American, species, with 
broader linear or oblong flat spikelets. The generic name Mega- 
stacJiya has undergone many vicissitudes. It was first founded by 
Beauvois on the Poa mucronata of his Elora of Oware and 
Benin ; but in drawing up the generic character for his ' Agro- 
stographie ' he had in view chiefly the common E. megastachya. 
Eoumier, more recently, founded a genua on those American 



118 ME. 0. BENTHAM ON GEAMINE^. 

species (^. reptcms &c.) whici have the spikelets more or less 
unisexual, the males usually flatter, longer, and with more flowers 
than the females ; but this separation of the sexes is very 
variable, and not always accompanied by any difference in habit. 
Some species may be occasionally quite dioecious ; in others the 
males and females are in different panicles on the same plant ; 
others, again, are variously polygamous ; and here, as in the 
Chilian Pow, the character is too inconstant to justify generic 
or even sectional separation. Megastachya, Pourn., does not in- 
clude JE. megastachya, Beau v. 5. Flatystachya, includes a few 
Africaner Arabian species with broad, flat, many-flowered spikelets 
with rather paleaceous glumes, and the rhachilla articulate as in 
Cataclastos. Amongst these Munro would include as E. genicu- 
lata the Uriza geniculata, Thunb., which in its thickened spikelets 
appears intermediate between Briza and JSragrostis, but has the 
prominently three-nerved glumes of the latter. 6. Sclerostachya, 
has three African or Asiatic species. The paleaceous glumes and 
articulate rhachilla are those of Flatystachya ; but the spikelets 
are not so broad, and the rigid habit with long and rush-like or 
short and pungent leaves are those oi ^luropus. 

36. Ipntjm, Philippi, is a single Chilian species unknown to 
me. From the author's description, it would appear to differ 
from the section Pteroessa {Cylindrostachyis) of Eragrostis in the 
articulate rhachilla and in inflorescence. 

37. CuTAjTDA, Willk., is a genus proposed for the European or 
North-African Festuca maritima, DC, F. philistcsa, Boiss., F. 
memphitica, Boiss., F. divaricata, Desf, F. inorassata, Salzm., and 
F. lanceolata, Eorsk., which, notwithstanding a general resem- 
blance to Festuca in the shape of the spikelets, could not remain 
in that genus without an essential alteration in its generic cha- 
racter, the glumes being strongly keeled and three-nerved from 
the base. The inflorescence is also peculiar. , 

38. Oreochloa, Link, is a single European mountain species, 
formerly included in Sesleria, which it approaches in its short 
compact inflorescence and the slightly elongated stigmas; but 
there are no barren spikelets, the spike is simple, with almost 
sessile bifarious spikelets like those of some species of Festuca 
(Scleropoa), and the glumes are those of Eragrosteae. 

39. EcTEosiA, Br., comprises three or four Australian species 
allied both to Eragrosteae and to Melicese, but technically rather 
better placed in the former subtribe. 



ME. G. BENTHAM ON SEAMIKEJil. 119 

As a Bixth subtribe, Melicece, I have collected five genera 
allied botli to Eragrostese and Festuceae, but technically connected 
with each other by their spikelets containing two or more empty 
glumes above the flowering ones. This character is also observed 
in JSctrosia and in Lojphatherum, whicli I had formerly included 
in the group ; but on other accounts the former appears better 
placed in Bragrostese, and the latter in Centothecese. 

40. Cetptochloeis, Benth., a single species probably from 
Patagonia, 41. Heteeachnj;, Benth., two Australian species, and 
42. Anthochloa, Nees, one or two species from the Andes of 
South America, are very distinct dwarf grasses, described and 
figured in Hooker's Icones. 

43. Melica, Linn., contains above thirty species dispersed over 
the temperate regions of the northern hemisphere, and extending 
down the Andes into extratropical South America, represented 
also in South Africa. This genus, the typical representative of 
the subtribe, has, been universally recognized since the days of 
Linnseus, and less tampered with than any other genus of equal 
extent, although it may be in some degree polymorphous in habit 
as well as in character. In the typical Melicas, however varied 
the panicle, long and narrow, or very loose and spreading, the 
spikelets are generally nodding or pendulous, with rarely more 
than two flowers ; the flowering glumes more or less scarious on 
the margins and never awned, and the terminal empty glumes 
one within the other, form an obovoid obtuse mass. In a section 
proposed by Thurber for four North-west American species under 
the name of Bromelica, the spikelets are erect, with more rigid 
glumes occasionally awned and three to eight flowers, the upper 
empty glumes narrower and not so closely packed, giving the 
plants altogether so difierent an aspect, that I have much hesi- 
tated whether I should not, as suggested by Thurber, raise the 
section to the rant of a genus. In both sections the flowering 
glumes have five or more nerves. M. stricta, Boland., is in some 
measure intermediate between the two. Chondr achy rum, Nees, a 
Brazilian species which I have not seen, would seem from the de- 
scription given to be a true Melica. 

44. DiAEEHENA, Eafin. {Korycarfus, Gorycarpus or IRosmeria, 
2i&&, Onoea, Franch. and Sabat.), two species, one from IN'orth 
America, the other from Japan, is very near Melica ; but the 
flowering glumes have only three nerves and are hardened round 
the grain, which usually exceeds them, and the stamens are reduced 



120 ME. a. BENTHAM ON GEAMINEJ3. 

to two or one. The habit is nearly that of the section JBromelica 
of Melica. 

Our seventh subtribe Centotheceoe is formed of a small number 
of tropical grasses, several of which have been occasionally re- 
ferred to Bambusese, but expelled from that tribe by all vrho have 
specially worked at it. The structure of the spikelet is that of 
some EufestucesB or Meliceae ; but the foliage is unusual, the 
lamina of the leaf is broad and flat, and between the numerous 
longitudinal veins are small transverse veinlets not observed in 
any others of the Order except in a few Bambusese. There is, 
however, none of that articulation of the lamina on the leaf-sheath 
which is almost universal in the latter tribe. The Centothecese 
comprise five genera. 

45. Centotheca, Desv., has two or three species from the tro- 
pical regions of the Old World. They are tall grasses with a 
loose panicle, the spikelets awnless with usually more than two 
flowers, without any, or with only one empty glume above them. 
In the common C. lappacea, Desv., the flowering glumes have on 
their back a few reflexed rigid hairs or bristles ; and that has 
been generally relied upon as the essential character of the genus ; 
but the bristles are sometimes reduced to one or two minute 
tubercles, or even wanting, and in an African species (probably 
the Foa mucronata, Beauv.) there is no trace of them, and yet 
the plant is in all other respects an undoubted Centotheca. 

46. Oethoceada, Beauv., is a single tropical- American species, 
with the habit, foliage, and inflorescence of Centotheca ; but the 
spikelets contain only one, or rarely two, fertile flowers, the second 
flower being usually male only ; the glumes have never the re- 
versed bristles of Centotheca ; and the spikelets appear to be fre- 
quently unisexual. 

47. LoPHATHEEUM, Brongn. {Acroelytrum and Allelotheca, 
Steud.), has one, or perhaps two, species from tropical and 
Eastern Asia. The habit, foliage, and inflorescence are those of 
Centotheca and Orthoclada ; but above the single fertile flower 
are several small empty, very shortly-awned glumes, densely 
crowded in a little unilateral tuft or crest, bringing the genus 
into connection with Meliceae. 

48. Steepto&tne, Beauv., is a single species sparingly scat- 
tered over East India, tropical Africa and America, and the 
southern states of North America, allied perhaps in some re- 
spects to Lophatherum, but quite isolated in habit and a variety 



ME. G. BENTHAM ON GEAMINEiE. 121 

of characters. The long narrow spikelets are few or numerous 
in a long, rigid, terminal unilateral spike ; and the exceedingly- 
long capillary styles become spirally twisted together far beyond 
the glumes. There is a considerable variety in the number 
of spikelets, in the number of flowers in each from one to 
four ; and even the stamens and styles are sometimes two, some- 
times three ; but I have been unable to trace any connection of 
these diversities either with each other or with geographical 
stations so as to mark distinct species. 

49. Zeugitbs, Schreb. (Senitis, Adans., Bespretzia, Kunth, 
KrombhoUzia, Pourn.), has five or six tropical -American species. 
They have not all the tall habit of the preceding genera ; for the 
best-known West-Indian and Central-American species, origi- 
nally described by Linnseus under Apluda, is a much weaker 
plant with smaller leaves ; they are, however, broad and flat, 
with the characteristic venation of the subtribe. The spikelets 
in the genus generally have one fertile flower with two to five 
male ones above it. 

I have already adverted to our eighth and last subtribe Mufes- 
tucece, or Festuceae proper, as difiering from Eragrostese in having 
five or more nerves to the flowering glumes instead of three or 
one only. Generally speaking, they have not the several-awned 
glumes of PappophoresB and Triodieae, nor the barren spikelets nor 
long styles of Sesleriese, nor the long hairs surrounding the 
flowering glumes of Arundinese, nor the cluster of upper empty 
glumes of Meliceee, nor the peculiar foliage of CentothecesB ; yet 
there are here and there exceptional species showing an approach 
to one or another of these characters, and interfering much with 
any definite line of demarcation. We include in the subtribe 
twenty genera, the last five of which are further characterized by 
the adherence of the grain to the palea ; but, as already observed, 
this character is not quite constant even in Festuca, and is occa- 
sional in such genera as Foa and Briza, where the grain is 
usually free. I have been unable to discover any other cha- 
racter which would distribute the genera of the subtribe into 
more satisfactory groups. 

The first two genera have a simple racemose inflorescence. 
50. PiETiEOPOGOTr, Br. {LophocJilcsna, Nees), has three species, 
one arctic, the two others Californian, distinguished by the keels 
of the palesB bearing a linear tooth or flat crest ; and 51. Betl- 
EiNiA, r. Schmidt, a single Japanese species, with two empty 



122 ME. G. BBNTHAM ON GBAMINE^. 

glumes close under the flowering one besides the two lower per- 
sistent ones. 

52. "Uniola, Linn. (Trisiola, Eafln., Ohasmanthium, Link), 
has four genuine North-American species, tall plants somewhat 
variable in inflorescence, but all with flat broad spikelets in which 
the three to sis lower glumes are empty, but in size and shape 
pass gradually into the flowering ones, which vary from three to 
about twenty. If U. racemiflora, Trin. (Z7. virgata, Grriseb.), from 
the "West Indies, be retained in the genus as having flat spikelets 
with more than two empty glumes below the flowering one, it 
must be considered as a very exceptional species with the inflo- 
rescence nearly oi LeptoMoa among Chlorideae. The small spike- 
lets are closely sessile in two rows in unilateral spikes ; and these 
spikes, shorter than in Leptochloa, are very numerous and crowded 
along the long peduncle. It would be better perhaps to regard 
the plant as a section of LeptocMoa rather than as a distinct 
genus. Pournier has added three Mexican species of Uniola 
which are unknown to me ; but, from his short characters, they 
would scarcely seem to be true congeners. U. prostrata, Trin., 
and its allies are now included in Distichlis. 

53. Distichlis, Eafln., comprises four or five closely allied 
species, or perhaps varieties of a single one, extending from North 
America down the Andes to extratropical South America, one of 
them found also in Australia. They are generally, but not 
always, maritime plants, with few spikelets nearly sessile in a 
dense panicle, and generally if not always strictly dioecious, though 
the two sexes diflFer but little in habit. The glumes are rather 
rigid and paleaceous, which induced Link to join the only American 
species with which he was acquainted, with the Mediterranean 
Poa sicula, Jacq., as a genus Srieopyrvm, a name retained by 
Presl and by ITournier for the American species. The European 
S. siculum, however, and some African congeners have the spike- 
lets hermaphrodite, a more regular bifarious inflorescence, and 
otherwise differ sufiiciently from the American forms to be main- 
tained as a separate genua which has a primary title to Link's 
name. 

64. .ZELTJEOPirs, Trin. (CflZo^^ecfl!, Spreng., ChamcBdactylis,'SBeB), 
has three species from the Mediterranean region. Central Asia, 
and East India, formerly included in Dactylis, but diflering in 
their creeping or prostrate branching habit, short, rigid, often 



ME. Q. BENTHAM Olf GEAMIBBJ3. 123 

pungent leaves, more numerous flowers in the spikelets, and some 
other minor points. 

55. Daottlis, Linn., is now limited to two species : — the com- 
mon and well-known Z>. glomerata, Linn., which from Europe and 
temperate Asia has spread over many parts of the civilized world; 
and Z). ccBspitosa, Porst., the celebrated Tussock grass of the 
Falkland Islands, which, though a much larger plant, appears to 
be strictly a congener. 

56. Lasiochloa, Kunth, has three or four South-African 
species with a close almost spikelike panicle and hairy glumes, 
allied in many respects to Koeleria ; but the inflorescence as well 
as the many-nerved glumes bring them nearer to Dacfylis. 

57. Beizopteum, Link, as now understood, is specially founded 
on the Mediterranean Poasicula, Jacq., to which are added three 
South- African species. The flat broad spikelets with coriaceous 
glumes are nearly those of Eragrostis sect. Flatystachya ; but the 
flowering glumes have seven nerves, and the spikelets are nearly 
sessile in a bifarious spike, or especially the lower ones closely 
clustered. I have already referred under DisticMis to the Ame- 
rican dioecious plants for which the name Brizopyrum. has been 
retained by Presl and by rournier ; the true Srizopyra are all 
hermaphrodite. 

58. SoLEEOCHLOA, Beauv., is limited to the S. dura, a small 
Mediterranean annual well characterized by the inflorescence and 
sliape of the glumes ; the other species, sometimes referred to 
Sclerochloa, belong chiefly to Outanda. 

59. Beiza, Linn., about ten species, of which the typical ones 
are chiefly European, though one has now spread over the greater 
part of the civilized world ; two sections are entirely American, 
tropical or northern. All are characterized by the very concave, 
sometimes almost vesicular, glumes enclosing a much smaller broad 
flat palea, the grain much flattened from back to front, and some- 
times, but not generally, adhering to the palea. The three best 
known European species have a very loose panicle with the spikelets 
hanging from capillary branches ; the Oriental B spieata, Sibth., 
differs in its narrow closer panicle. The American species have been 
separated as two distinct genera, which may be. retained as sec- 
tions, though with little difierence in essentia] characters. Ghas- 
colytntm, Desv., has the awnless spikelets of the European species ; 
but the "panicle, though branched, is much more compact, the 



124 ME. G. BENTHAM ON GEAMIJTBi. 

spikeleta almost sessile. Galoiheca, Desv., with a loose spreading 
panicle, has broadly scarious awned glumes. 

60. ScHisMUS, Beauv. {Electra, Panz., Hemisacris, Steud.),ha8 
three or four species, one of them widely spread from the Medi- 
terranean region, eastwards to Afghanistan and Arabia and 
westwards to the Canary Islands, the others South African. All 
are annuals with a narrow panicle, and distinguished by the long 
empty glumes quite enclosing the flowering ones. 

61. Nephelochloa, Boiss., limited to the original Oriental 
species, is a very elegant little grass with the habit of Aira invo- 
lucrata, and is figured in the last part of Hooker's Icones. The 
species added to the genus by G-risebach, for which he was obliged 
to alter Boissier's character, are now restored to Poa. 

62. Poa, Linn., is a cosmopolitan genus, chiefly extratropical, 
which, after frequent extensions and reductions, has now become 
fairly limited to a series of about eighty species. They form a 
group natural enough as to the great majority of species, dif- 
fering from JSragrostis in their five-nerved flowering glumes, from 
Glyceria and Festuca in their glumes keeled from the base ; but 
here, as elsewhere, there are species apparently intermediate 
between these large genera, and several smaller ones are only 
separated by characters of little importance. Poa has also been 
distinguished from Pestuca by the obtuse, always unawned glumes, 
and by the non-adherence of the grain to the palea. The former 
character is general, but not absolute ; several species of Poa 
have acute glumes, and in P. lanuginosa, Poir., they bear a fine 
point which might almost be termed a very short awn. And as 
to the grain, though it is usually free, there are several Chilian or 
Australian species and some Asiatic ones where it is adherent to 
the palea, as in Festuca, and even in the common Poa pratensis 
it is often more or less adherent, whilst there are several true 
Pestucas where it is quite free. 

Most of the widely spread species of this genus are so variable 
that it would require much more research into specific detail 
than I can at present bestow upon them to distribute them into 
natural groups or sections ; and I can only refer to the following 
as having been proposed as sections or separate genera : — Pseudo- 
poa, proposed by C. Koch as a section oi Festuca, includes P.per- 
sica, Poir., and two other temperate-Asiatic species, with very 
small spikelets and with nearly the habit oi Nephelochloa, to which 
Grisebach has referred them, but which appear inseparable from 



ME. G. BENTHAM ON GEAMINEiE. 125 

Foa notwithstanding the adherence of the grain to the palea. 
Leucopoa, Grriseb., is the temperate-Asiatic P. alhida, Turcz., 
with the spikelets rather larger than usual, and somewhat sca- 
riose and shining glumes like those of several Chilian species. 
Bioicopoa, E. Desv. {Dispar, Doell), is a section proposed for 
P. chilensis, Trin., P. lanuginosa, Poir., and their allies, in which 
the spikelets are usually, but perhaps not always, dioecious. In 
the dried state there is very little external difference between 
the male and the female panicle ; and there are certainly Chilian 
specimens otherwise very near P. chilensis which have herma- 
phrodite flowers. Doell also places P. lanuginosa in his herma- 
phrodite section, whilst Emile Desvaux describes it as dioecious, 
as I have generally found it, The stamens, however, are very 
deciduous, and the ovary at the time the stamens are still enclosed 
very minute ; and it requires careful observation to ascertain the 
real absence of the one or the other. It is probable that many 
species hitherto supposed to be perfectly hermaphrodite are more 
or less polygamous. P. lanuginosa shows, moreover, an appi'oach 
to Festuca in the fine though short points to the flowering glumes, 
and in the adherence of the grain to the palea. Poidimn, Nees, 
a Brazilian species, was separated by I^ees from Poa on account 
of a reduction in the number of flowers to one or two, which Doell 
finds to be by no means constant. 

63. Oolpodium:, Trin., about ten species, from the Levant and 
Eussian Asia, might perhaps be regarded as a section of Poa. 
It diflfers in very little besides the small spikelets containing only 
one or two flowers, thus connecting Poa with the Agrostese. 
The arctic plant, published by E. Brown as a doubtful Oolpodium, 
now forms Grisebach's genus Arctagrostis, included above under 
Agrostese. 

64. G-EAPHEPHOETJM, Desv., including Scolochloa, Link (Plu- 
minia, Fries), and Dvpontia, Br., contains seven North-American, 
North-European, or North- Asiatic species, very well worked up 
and distributed into four sections by Asa Gray. They are all 
very near Qlyceria, differing chiefly in the hairs surrounding the 
flowering glumes, which induced several botanists to refer the 
genus to Arundinese, though very different in habit and in the 
shape and venation of the glumes. The hairs of the spikelets 
are, moreover, very variable, shorter than in true Arundinese, very 
short in the section Arctophila, and not entirely abseot in one or 
two species of Qlyceria. 

LINN. JOTJKN. — BOTANY, VOL. XIX. L 



126 ME. G. BENTHAM ON GBAMINEiB. 

65. Gltcebia, Br., if we include Atropis, Eupr., is a genus of 
nearly thirty species spread over the extratropical region?, northern 
or southern, both of the New and the Old World. It is very 
nearly allied both to Poa and Festuca, differing from the former 
in the flowering glumes rounded at the base without any promi- 
nent keel, from Festuca in the broader more obtuse glumes, and 
the grain usually free from the palea, and from both in the short- 
ness of the nerves of the glumes. The habit is somewhat variable, 
but as much so in each section as in the whole genus. The two 
sections into which it has been divided, often raised to the rank 
of separate genera, are : — 1. Hy&rochloa, Hartm. (Forroteranihe, 
Steud., Exydra, Endl., Olyceria proper of many botanists), with 
the lodicules connate and truncate or deficient, and the thick grain 
only marked on the inner face with a very narrow lined furrow or 
quite smooth ; and 2. Atropis, Eupr. {Fuocinellia, Parlat.), with 
two distinct lodicules and the grain more or less compressed from 
front, to back, with a broad furrow or almost fiat on the inner 
face. But these characters are not constant. The lodicules in 
the typical SydrocMoa, Q. fluitans, Br., though thicker than in 
Atropis, and usually connate, are readily separable and occasion- 
ally spontaneously free ; in Q. aquatica, Sm., they are so short as 
to render it diiBcult to say whether they do or do not cohere, and 
in Q. nervata, Trin., and in Q. pallida, Trin., I can find no trace 
of them ; in Atropis they are usually, but not always, more deve- 
loped and thinner. The shape of the seed and of its furrow seems 
to vary from species to species, in so far as I have been able to 
procure it well ripened. 

66. Eestuca, Lian., is one of the genera as to whose limits 
botanists are the least agreed. "With the exception of the exclu- 
sion of Cutanda and Frizopyrum, we have followed generally the 
arrangement proposed by Cosson and Durieu, which would include 
between seventy and eighty species (estimated by some at above 
two hundred and thirty), almost cosmopolitan in their geogra- 
phical distribution, but most abundant in the northern temperate 
regions of the Old World, with not many American and very few 
tropical species. They are generally distinguished in the sub- 
tribe by the flowering glumes rounded without any prominent 
keel at least at the base, and acute or awned at the end, and by 
the glabrous grain adhering to the palea. But there are excep- 
tions to each of these characters ; and some species run very much 
into Foa, whilst others are scarcely distinct from Fromus. The 



MR. O. BENTHAM ON GRAMINEjE. 127 

following are the most prominent groups established as sections 
or proposed by some as independent genera: — 1. Vulpia, Gmel. 
{Mygalurus, Link), panicle narrow, dense, and usually unilateral, 
the outer glumes very unequal, one often minute or almost obso- 
lete, the flowering glumes awned, and frequently, but not always, 
only one stamen. If we had only the common European species, 
this might well have been kept up as a genus ; but in the South- 
American F. ulochteia, Doell, and Jl leptothrix, Trin., the panicle 
is loose, as in Mufestuca, and in F. delicatula. Lag., F. setacea, 
Parlat., the awn is sometimes very short and the inflorescence 
rather that of Eufestuca. The proportions of the outer glumes 
vary from species to species. 2. Eufestuca, comprises the greater 
number of the species, with a loose, spreading or narrow panicle, 
the outer glumes nearly equal, the ilowering ones acute or mu- 
cronate, rarely short-awned, and three stamens. Amongst them 
Grrisebaeh has distinguished a section PhceocMoa, with the ovary 
slightly hairy at the top as in Bromus : but the character is very 
variable in F. sylvatica, F. varia, and their allies. Doell has pro- 
posed a section Mallopetalum for the Brazilian F. ampliflora, 
Doell, apparently the same as the Mexican F. amplissima, Rupr., 
characterized by the lodicules villous at the top. I find these 
lodicules fringed with long hairs at the top exactly as in F.fim- 
Iriata, Nees, which Doell places amongst his Festucce legitimcB 
with glabrous lodicules. Selleria, Fourn., is proposed as a genus 
for the Mexican Bromus lividus, H. B. K., which Sprengel after- 
wards and Kunth himself removed to Festuca, of which it has all 
the characters of the awned species. The inflorescence is at first 
very like that of some varieties of Bromus tentorum ; but as it 
advances the spikelets become very much divaricate or refiexed, 
giving the plant a peculiar habit. 3. Schedonorus, Beauv. {Am- 
phigenes, Janka), comprises F. pratensis, Huds., F. sylvatica. 
Host, F. nutans, Host, F. littoralis, LabilL, F. Hookeriana and 
F. scirpoidea, P. Muell., and a few others, tall plants, with loose, 
narrow or spreading panicles, awnless glumes, and the grain quite 
free from the palea, thus connecting the genus with Poa. Fries 
and other Swedish botanists, whilst they rightly referred Beau- 
vois's species of Schedonorus back to Festuca, transferred his generic 
name to a very difi'erent group, which now forms the sections 
Festueoides and Stenobromus of Bromus. 4. Gatapodium, Link, 
including Micropyrum, Link, differs from Fufestuca in the inflo- 
rescence, which is nearly the simple spike of Hordeese ; but the 



128 ME. G. BENTHAM ON GBAMINE^. 

rhachis is not notched and tlie spikelets are not quite sessile, the 
lower ones often two or three together on a very short branchlet, 
not collateral. Nardurus, Eeichb., is the F. unilateralis, Schrad., 
differing from the rest of the section in the flowering glumes 
mucronate or shortly awned. Oastellia, Tineo, is the -F. tubercu- 
lata, Coss. and Dur., in which the flowering glumes are minutely 
tuberculate and the spike often shortly branched. Nardurus 
montanus, Boiss., scarcely differs from F. {Vulpia) delicatula. 
Lag., and F. cynosuroides, Desf.,is also referable rather to Vulpia 
than to Gatapodivm. F. loliiim, Balansa, may really be said to 
be intermediate between Festuca (Oatajaodium) and Lolium. F 
unioloides, Kunth, is Brizopyrum siculum. Gatapodium fusiforme, 
Nees, is Tripogon hromoides, Nees. 5. Scleropoa, Griseb. {Sclero- 
chloa, Eeichb., not of Beauv.), annuals, often small, with one- 
sided panicles, the short rigid branches bearing few almost ses- 
sile spikelets, at flrst erect, then spreading or reflexed, giving 
nearly the habit of Cutanda, but the glumes entirely those of 
Festuca, 

67. Pantatheea, Philippi, and 68. Podophoeus, Philippi, are 
monotypic genera from the island of Juan Pernandez, both very 
near Bromus, but scarcely reducible to it. 

69. Beomps, Linn., is a fairly natural genus of about forty 
species, generally distributed over the temperate regions of the 
northern hemisphere, with a very few tropical or southern spe- 
cies. Very near Festuca, with which it is closely connected 
through Festuca gigantea, Vill. (Bromus giganteus, Linn.), it differs 
generally in the flowering glumes distinctly notched or shortly 
two-lobed at the end, with an awn between. the notches often not 
quite terminal and sometimes slightly twisted, showing an ap- 
proach to Avena, and in the grain (always adnate to the palea) 
crowned by a little appendage or tuft of short hairs. These cha- 
racters are, however, not quite constant ; and the four following 
sections into which the genus has been divided run also much into 
each other, though some of them are often regarded as separate 
genera : — 1. Festucoides, Coss. and Dur. (Schcenodorus, Griseb.), 
consists ol B. asper and 5. inermis, Linn., B. ereetus, Huds., and 
their allies, tall perennials, coming nearest to Festuca, with the 
awns usually very short or reduced to small points. 2. Steno- 
Iromus, Griseb. (Anisanthe, C. Koch), mostly annuals, with narrow 
spikelets and iong-awned glumes. Schedonorus of Fries and other 
Swedish botanists, but not of Beauvois, includes both Festucoides 



MR. G. BENTHAM ON GEAMINEiB. 129 

and Stenohromus. 3. Zeohromus, Griseb. (Serrqfalcus, Parlafc.), 
spikelets usually broad and thick, the flowering glumes awned, 
and the nerves of all the glumes more numerous than in the pre- 
ceding sections. Libertia, Lejeune {Michelaria, Dumort.), is the 
B. ardennensis, Kunth, differing from S. (Zeohromus) secalinus in 
the lateral lobes or teeth of the flowering glumes produced into 
slender points or very short awns. Triniusa, Steud., is the B. 
DanfhonicBjTnn., very near B. (Zeohromus) macrostachyus, Desf. ; 
but most of the flowering glumes, especially the upper ones of 
the large spikelets, bear three long recurved awns. 4. Cerato- 
chloa, DC. (or Beauv.), three or four American species, extra- 
tropical or Andine, with flat spikes not unlike those of Uniola, but 
at length often thickened as in Zeohromus, and the flowering 
glumes scarcely notched at the end, and the awn very short. 
Fouruier rightly retaios the 5. (Ceratochloa) purgans, Linn., in 
5roMMS (under the name of B. Hooheri), but keeps up the genus 
Ceratochloa for the original C unioloides, DC, as having the 
lodicules connate. I have examined a number of specimens, both 
wild and cultivated, and have always found the lodicules attached 
by a broad base and contiguous, but quite free or only exceedingly 
shortly cohering at the very base. 

70. BBACHTPODirM, Beauv. {Hemibromus, Steud.), has five or 
six European or temperate- Asiatic species, one or two of which 
are also in Mexico, Colombia, and tropical and southern Africa. 
They closely connect Festuca with Agropyrum ; the spikelets are 
those of the former though usually longer, and the simple spicate 
inflorescence is that of Agropyrum, except that the rhachis is not 
articulate and not at all or scarcely notched, and the spikelets are 
not so closely sessile, usually few and distant. TracTiynia, Link, 
is B. distacJiyum, Uoem. and Schult., which differs from the rest 
of the genus as an annual, with only one or two spikelets at the 
end of the peduncle. 

Tribe XIII. Hoedee2B. 
This tribe, one of the most definite of the Poacese, is charac- 
terized chiefly by the inflorescence. The spike is always simple, 
except in abnormally luxuriant cultivated varieties or monstrosi- 
ties, the rhachis notched and often, but not always, articulated, 
the spikelets (one- or several-flowered) singly or two or more 
collaterally sessile at each notch. The genera, mostly very dis- 
tinct, belong to the temperate regions of the New as well as the 



130 ME. G. BENTHAM ON GEAMINEJE. 

Old "World, chiefly in the northern hemisphere ; and scarcely any 
species, except as introduced weeda or escapes from cultivation, 
penetrate within the tropics. The twelve genera are readily 
ranged in three distinct subtribes, and require but little comment 
on the present occasion. 

The first subtribe, Triticew, comprises four genera, in which 
the spikelets have three or more, or very rarely only one or two, 
flowers, and are singly sessile at each notch of the rhachis. 

1. LoLiTJM, Linn., is at once distinguished from all others of 
the tribe by the position of the flat spikelets with their edge 
to the rhachis. Steudel enumerates twenty-two species ; most 
authors reduce them to three or four, which run much into each 
other. De Rouville published at Montpellier a detailed mono- 
graph, in which he rejects all the old species and redivides the 
genus into three primary and several subordinate races, to which 
he gives new characters and new names, doing little but add to 
the prevailing confusion. Two genera have been founded on in- 
dividual species or forms — CroBpalium, Schrank, is the L. temulen- 
tum, Linn., and .Arthrochortus, Lowe, is a Madeiran species or 
variety very near to L. rigidum, Gaudin {L. strictum, Parlat.), 
and to some varieties of L. temulentum. 

2. A GEOPTEUM, J. Gsertn. {Elytrigium, Desv.), contains about 
twenty species, formerly regarded as congeners of the cultivated 
Wheats, from which they differ much in habit and technically in 
the lateral nerves of the flowering glumes connivent at the top 
or confluent into the terminal awn. They are well distributed 
into two sections : — 1. Agropyrum proper, mostly perennials, 
with the spikelets more or less distant along the common pe- 
duncle or rhachis, the outer empty glumes usually very unequal- 
sided and not keeled. To this section belong the common A. re- 
pens, A. junceutn, A. caninmn, and a few others. Sosgneria, 
C. Koch, is, according to G-risebaoh, a species closely allied to 
A. caninum. AnthosacJine, Steud., is the Australasian A. scabrum, 
Beauv. {Festitca scalra, Labill.), which, with the closely allied 
East-Indian A. semioostatum, Nees, and the Oriental A. longearis- 
tatum, Boiss., differs from the commoner species in the denser 
spikes and narrower glumes tapering into long awns at length 
diverging. A. pectinatvm, Beauv., is an Australian species still 
further connecting .4yropy>'jwre proper with JLremopyrum. 2. Ere- 
mopyrum, Ledeb. {Cremopyrum, Schur, perhaps by a clerical 
error, Costia, Willk.), mostly annuals, with the spikelets distichous 



MB. &. BENTHAM ON GKAMINEiE. 131 

and close together in a short dense spike, the narrow empty 
glumes nearly equal-sided and keeled. Two species, A. villosum 
(Secale villosum, Linn., Saynaldia, Schur) and A. hordeaceum, 
Boiss., form the proposed section Dasypyrum, Coss. and Dur. 
{Pseudosecale, Gren. and Godr.), differing slightly from the other 
species in the empty glumes rather unequal-sided, and one lateral 
nerve on one side of the keel very frequently as prominent as the 
keel itself, giving the glume the appearance of being two-keeled. 
Heteranthelium, Hochst., from the Levant, is a species very near 
A. {Eremopyrum) orientate, with a dense villous spike, and several 
of the spikelets, especially near the base and apex of the spike, 
often sterile with empty glumes. 

3. Sj3CAiiE, Linn., is now reduced to two species or perhaps 
varieties, the cultivated Rye, of which S. montanum, Guss., is 
supposed to be the original spontaneous form, and S. fragile, 
Bieb. The genus differs slightly from the section Eremopyrum 
of Affropyrum in the dense cylindrical spike, and in the spikelets 
usually containing only two flowers. 

4. Teiticttm, Linn., excluding Affropyrum and including ^yi- 
lops, can scarcely reckon more than ten botanical species ; the 
most prominent character separating them from Agropyrum con- 
sists in the shape of the spikelets not so flat, and especially in the 
lateral nerves of the flowering glumes not conaivent, but remain- 
ing parallel or nearly so, and either stopping short of the apex or 
produced beyond it into distinct teeth or awns. There are three 
rather distinct groups : — 1. The cultivated Wheats, of unknown 
origin, in which the flowering glumes are keeled at the end and 
sometimes from the base, and terminate in a single awn, the 
lateral nerves usually barely reaching to the end of the glume. 

2. CrithoMum, Link, founded on T. monocoecmi, Linn. {T. boeoti- 
cum, Boiss.), in which the spikelets have only one fertile flower, 
and the flowering glume is keeled from the base and ends in 
a single awn, T. bicorne, Torsk., with two or even three fertile 
flowers and the lateral nerves of the flowering glumes sometimes 
produced into short teeth, may be referred to the same section. 

3. ^gilops, Linn.< above forty published species, which Munro 
reduces to seven or eight, differing from the cereal wheats in the 
flowering glumes more rounded at the back and not at all keeled, 
and in the lateral nerves of the flowering glumes often produced 
into long awns, especially in the upper end of the spike. The 



132 ME. G. BENTHAM ON GRAMINE^. 

extreme readiness with which some species hybridize with the 
cultivated wheats has given rise to the suggestion, strongly advo- 
cated by some, positively rejected by others, that it is in some of 
the common species oi j^gilops that we must look for the original 
of our cereal wheats. 

The second subtribe, Lepturem, is characterized by the slender 
spikes and the spikelets solitary at the notches, each with only 
one or rarely two flowers. We refer to it five genera, placed by 
Kunth and some others in Eottboelliese, from which they differ in 
the outer empty glumes, when present, persisting below the arti- 
culation of the rhachilla. 

5. LEPTTTErs, Br., including Fholiurus, Trin., has six species, 
five of them with the ordinary geographical range of the tribe, 
the sixth, L. repens, Br., exclusively Australasian or South Pacific 
and maritime. They are distinguished in the subtribe by the 
rigid outer empty glumes, one or two in number, much longer 
than the hyaline flowering glume, thus showing the nearest 
approach to Eottboelliese. They differ from each other sufficiently 
to have been referred by different botanists to different genera. 
L. cylindricus, Trin. (-£. subulatus, Kunth), included by Link in 
Ophiurus, by Keichenbach in Monerma, has one outer empty glume 
and one flower with no empty glume above it. The Australasian 
L. repens, a much larger plant than any of the others, has one 
outer empty glume, one flower, and above it a glume either 
empty or enclosing a palea, but no flower. L. persica, L. incur- 
vata, and L. filiformis, Trin., have two lower empty glumes, one 
flower, and no empty glume above it. li. pannonicus, Kunth, 
forming Trinius's genus PhoUurus, and referred by T. Nees to 
Ophiurus, has two outer empty glumes and two perfect flowers. 

6. PsiLUEiTS, Trin. {Monerma, Beauv., partly, Asprella, Host 
but not of Willd.), is a single annual, near Lepturus, but with 
only one minute empty glume, a single narrow and awned flower- 
ing glume, and only one stamen in the flower. 

7. Naedus, Linn., is a single well-known small perennial, the 
position of which in the system is rather puzzling. The spikelet 
has only one flower without any empty glumes below it or pro- 
longation of the rhachilla above it, which might have decided its 
relatiouship either to Panicacese or to PoacesB, and its long simple 
style might indicate an afSnity to some Panicese or to Sesleriese ; 
but on the whole it seems nearest allied to the Lepturese, a sup- 
position which might be confirmed, if we regard the rather pro- 



ME. G. BENTHAM ON GEAMINE^. 133 

minent lower margin of the notches of the rhachis as a rudimen- 
tary glume. 

8. Kealikia, Coss. and Dur., is a single Algerine species 
unknown to me, but well described, and evidently rightly placed 
in the present group. 9. Oeopetium, Trin., is a dwarf East- 
Indian species remarkable for the cylindrical spike, with per- 
fectly immersed spikelets as in some MottloelUas and Ophiurus, 
but with the outer persistent glumes of Hordeese. 

The third subtribe Eh/mecB comprises three genera, in which the 
spikelets are two or more, collaterally sessile at each notch of the 
spike, or the lateral ones very shortly stipitate. 

10. HoBDJsrM, Linn., was restricted by Beauvois to the common 
cultivated barley, S. vulgare, Linn., which in a great variety of 
forms is of very ancient cultivation, and whose indigenous origin 
is no more known than that of our wheats. Amongst these 
forms the East-Indian H. agiceras, Eoyle, has been proposed by 
E. Meyer as a genus under the name of CritJio ; but it cannot be 
otherwise considered than as a luxuriant monstrosity. The really 
spontaneous species of Hordeiim amount to about twelve, distin- 
guished from Mymus by the single flower in each spikelet, and 
distributed into three sections: — 1. Zeocriton, Beauv. (Critesion, 
Eafin.), for the S. mttrinum, H. hulhosum, H.jubatum, Linn., and 
some others, in which the central spikelet alone of each three is 
fertile, the lateral ones sterile or reduced to empty glumes ; 
2. OritJiopsis, Jaub. and Spach (sect. Medusather of Elymus, 
Grriseb.), for the H. crinitum, Desf., and its allies, with two per- 
fect spikelets at each notch, the intermediate one deficient or 
rarely represented by one or two empty glumes ; 3. Ouviera, 
Koel., for the H^ sylvaticum, Huds. (Ulymus eurojpceus, Linn.) , 
with three collateral spikelets. 

11. ELTMrs, Linn., as now generally limited, comprises about 
twenty species, distributed into three sections, all distinguished 
from Sordeum in having two or more flowers to each spikelet : — 
1. Sitanion, Eafin. {Polyantheriie, Nees), for the North- American 
M. Sitanion, Schult., with the flowering glumes usually three- 
awned; 2. Glinelyna, Griseb., with the spikelets usually two only 
to each notched and the flowering glumes with one long awn ; 
and 3. Fsammelyna, Griseb., taU rigid species with often more 
than two spikelets to each notch, and the flowering glumes un- 
awned or with only very short awnUke points. 

LIHN. JOTJEir. — BOTANT, TOl. XIX. M 



134 MB. G. BENTHAM ON GEAMINEJE. 

12. AsPEELLA, Willd. {Systrioo, MoencL, Qymnostichum, 
Schreb.), has three species, of wliicli two are Nortli-American, 
tte third from New Zealand. There are two or three collateral 
spikelets to each notch as in the preceding genera ; but the outer 
empty glumes are entirely deficient, except sometimes one or two 
slender ones to the lower spikelets of the spike. Willdenow's 
name has the priority over Schreber's ; for although the Beschrei- 
bungen of the latter author bears the date 1769 on the titlepage, 
the third part, in which the present genus was proposed, was 
only issued in 1810. 

Tribe XIV. Bambuse^. 

The Bamboos have been so admirably monographed by Munro 
in the twenty-sixth volume of the Linnean ' Transactions,' that 
I have very few notes to make on the present occasion. Since 
the appearance of that memoir, Balansa has published a New- 
Caledonian Bamboo forming the distinct genus Greslania ; a fur- 
ther acquaintance with Thamnocalamus has induced its reunion 
with Arvmdinaria ; and, on the other hand, Merostaohys capitata. 
Hook., is so very different in inflorescence from the rest of the 
genus, that I have proposed to separate it under the name of 
Achroostaehys. I have also proposed as a new genus Melocala- 
mus, the Pseudostachyitm oompaetiflorum, Kurz, published since 
Munro's monograph. There is also much confusion in the generic 
term BeesTia, which, though used by Eheede for a Peninsular 
species of Bamboo, was first characterized by Kunth chiefly from 
the more eastern Bamhusa laccifera, Eoxb., now Melocanna lam- 
lusoides. He did indeed also include the Peninsular and Ceylon 
species ; but that was first properly characterized as a separate 
genus by Thwaites, under the name of Ochlandra, which it seems 
advisable to adopt, though the genus may include Eheede's Beesha, 
a name which it seems best to consider only in the specific sense 
first given to it. 



DE. W. E. m'NAB on 4ECTIC DKIET WOODS. 135 

Eeport on the Arctic Drift Woods collected by Captain Eeilden 
and Mr. Hart in 1875 and 1876. By W. E. M'Nab, M.D., 
P.L.S., Professor of Botany, Eoy. Coll. of Science, Dublin. 

[Bead November 3, 1881.] 

In the following Eeport I have endeavoured to detail the results 
of my examination of the drift woods brought from the Arctic 
regions by the naturalists attached to the recent Arctic expedition 
under Captain Nares. There are thirteen specimens of drift 
wood and one specimen of bark, collected in different localities by 
Captain Feilden and Mr. Hart ; and these were placed in my 
hands for examination by Professor Oliver, P.E.S. The speci- 
mens of wood are all completely devoid of bark ; hence it was im- 
possible to distinguish the genus to which some of the Coniferous 
woods belong, as, for example, Pioea and Larix, the genera to 
which most of the woods may be referred. In general the 
woods were well preserved and in good condition, except on the 
very surface ; hence there was little difficulty in obtaining proper 
sections for microscopic examination. The woods were all cut 
in three directions, as is usual in examining dicotyledonous and 
coniferous woods ; and the sections were viewed both dry and 
when mounted in Canada balsam. Careful comparison of the 
drift woods with sections of named woods has not enabled me to 
identify the species in any case ; hence the whole of the results 
obtained must be considered unsatisfactory. In the following 
list the specimens are lettered A to 0, the letters &c. correspond- 
ing to the labels on the slides of the preparations accompanying 
the Eeport*. 

1. PiNus sp. (One species.) 

Two portions of wood are referable to Pinus. These I have 
indicated by the letters A and B. Both the pieces in the collection 
are marked as from the same locality, viz. " Head of Discovery Bay, 
April 1876," but one of them (B) as having been " 100 yards from 
the water, embedded in sand." The woods are quite similar in 
outward appearance, and are portions of comparatively large 

* I have also examined a small oolleotion of drift woods made by Staff-Sur- 
geon Edward L. Moss, M.D., B.N., Surgeon of the 'Alert.' His specimens are 
similar to those obtained by the naturalists of the Expedition , with one excep- 
tion, Tiz. a portion of a stem of a species oi Juniperus. The stem must have 
been of some size, and belongs apparently to a North-American species. 



136 



BE. W. E. m'NAB on the AECTIO DEIPT WOODS 



trunks or branches. The central portions of tlie stem are well 
preserved ; but the outer part is soft and partially destroyed. 
A has well-developed annual rings, some of the rings near the 
periphery of the stem being, however, small and feebly developed, 
thus indicating a deficiency in growth. The wood of B pre- 
sents the same general characters ; and I am inclined to consider 
them portions of the same species of Finns, if not parts of the 
same tree. 




Sketch Map showing the route of the late Arctic Expedition between the 
parallels 78°-83° N. lat., with the localities generally where the drift 
woods were obtained. 



COLLECTED BY CAPT. FEILDEN AND ME. HAET. 137 

2. Abies sp. (One species.) 

One piece of wood of small size ; but probably a portion of a 
larger stem cut and rounded by man, belongs to the genus Abies 
(C). By comparison with other sections, it seems to come very 
close to Abies pectinata. The specimen is marked " Discovery 
Bay, 20 feet above sea-level, Aug. 1875, Captain Peilden." The 
annual rings of wood are large and well developed. 

3. PiCEA or Laeix. (Three species.) 

Seven pieces of drift wood are to be referred to one or other of 
these genera ; but, owing to the absence of the bark, it is impos- 
sible to decide definitely. One of the specimens (D) comes very 
near Larix, and differs from aU the other woods in the collection. 

? Laeix sp. This is the specimen D, marked " Dumb-bell 
Harbour." The stem has been large, and is well preserved, and, 
by comparison with named sections of Larix, seems to come very 
near L. eiiropwa. 

? PiCEA sp. Two specimens seem to belong to one species, viz. 
E, "Label incomplete. Upon Moe. Sept. 12, 1875," and P, 
" On Ploe, lat. 82° 30' N. Capt. Peilden." These are portions 
of well-preserved woods, white and firm, and having the same 
microscopical characters. They are probably not portions of the 
same stem, as I believe E is almost certain to be from Mr. 
Hart's collection, while E is Captain Eeilden's. The annual 
rings are well developed in both specimens. 

PPiCEA sp. Four specimens. Gr, H, J, K. All similar in 
microscopic character, and belonging either to Picea or Larix. 

(G) " 1 mile inland and 150 feet elevation at 'Alert ' winter- 
quarters, Eeb. 1876. Captain Eeilden." This piece of stem has 
well-developed annual rings. 

(H) " Drift wood. Bottom of Musk-ox Ejord. Sept. 16, 
1875." 35-5 inches long, 16 in circumference. Portion of a 
large stem with well-developed annual rings. 

(J) " No. 1. No locality nor date." A small piece of very 
much waterwom drift wood with well-developed annual rings, 
and probably a portion of a large stem. 

(K) " No. 1009^ No locality." Small portions of a large stem 
in a good state of preservation, and having woU-developed annual 
rings. 

PiCEA sp. Bark only. The specimen (L) is from pieces of 
bark evidently of a Ficea, and marked " On floe in Dumb-bell Bay, 



138 DH. w. E. m'nab on aectic drift woods. 

Sept. 1875. Found by Commander Markham." It is not impro- 
bable that this may be the bark of the commonest drift wood, and 
may therefore help to identify the genus of the six specimens of 
wood just described. 

4. Taxus sp. (One species.) 

A single sample of very much waterworn pieces of drift wood, 
which have probably been embedded in mud, is referable to the 
genus Taxus, having the spiral markings clearly shown in the 
wood prosenchymatous cells. It is marked M, "Out of cloth 
bag ; no locality." The annual rings are extremely imperfect and 
very numerous. 

6. PoPDLTJs sp. (One species.) 

Two pieces of drift wood are to be referred to Fopulus (near 
tremuld), and are interesting as being the only species of dicotyle- 
donous wood in the collection. One of the specimens (N) is 
marked " Drift wood. Musk-ox Bay. Sept. 1875;" and the other 
(0), " East Gary Island. Capt. Feilden." The former is a por- 
tion of a large stem, and is in an excellent state of preservation, 
while the latter is equally well preserved, but is only a part of a 
small branch. In both of the woods the annual rings are well 
developed. 

Of the 14 specimens submitted to me for examination, 13 are 
samples of wood, and 1 is of bark alone without any trace of wood, 
the bark being evidently coniferous and to be referred to the genus 
Picea. Of the 13 woods, 11 are coniferous, and only 2 dicotyledo- 
nous, both belonging to the same genus, Populus, and to the same 
species. 

The 11 coniferous woods belong to four, or perhaps five, genera, 
there being 1 species oiFinus, loiAbies,loiZaria;?,2o{Pmus 
or Larix, and 1 of Taxus. Of Pinus there are 2 specimens, of 
Abies 1, Larix or Picea 7, and Taxus 1 — the commonest form being 
some kind of Picea, probably an American Spruce. 

I have not been able to identify the species, but, from careful 
comparison of specimens, am inclined to think that most of them 
are North- American ; and as the annual rings are usually very well 
developed, the trees must have grown in the more northern tempe- 
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EVENmO- MEETINGS OF THE LINNEAN SOCIETY. 

To BE HELD AT BUBLINGTON HoUSE, SESSION 1881-82. 

On Thursday Ereninga, as undermentioned, the Chair being taken at 8 p.m. 
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March 2 








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