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| VOE! FOREST BOTANY SERIES Part | |

TH

INDIAN FOREST
MEMOIRS

On some Indian Forest Grasses and their
| Oecology

Imperial Forest Service
Botanist at the Forest Research Institute, Dehra Dun, India

Published by Order of the Government of India

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CONTENTS:

INTRODUCTION.
PaGEs.
Importance of Forest Grasslands— (cology defined — Hydrophytes,
Xerophytes, Mesophytes — Formation — Steppe — Savannah — Asso-
ciation — Hygrophytes — Tropophytes — Primary — subdivision into
Grassland and Woodland advisable—Progressive, Regressive and
Parallel Succession—Water supply not necessarily the dominant factor
regulating the distribution of plant-communities—Adaptations to
Temperature—Importance of Light—Plants with different adapta-
tions in same Habitat—Suggested scheme of Cicological Classification
for India—Value of (cology for Forest Officers—Importance of
(Ecology in study of Plant Diseases—Utility of Forest Experiments—
Importance of Field Observations—Lines of work adopted in study
of Forest Grasslands—Species selected—Arrangement of Information 1—35
CHAPTER I.
Description oF Locatrry.
Topography—Climate—Local Types of Vegetation . : : . 39—49
CHAPTER II,
(¢) Devaitep Description or Species.
Saccharum spontaneum ; : ; : : 2 . ° : ; 50—62
Saccharum Munja_ . : : : : : 2 : : 62—79
Saccharum Narenga . : : ; : : : ; 4 : 19— 87
Erianthus Ravenne . : : ; : : : : : ; §7—91
Imperata arundinacea ‘ : : ; : : : . 91—102
Triraphis madagascariensis : : : : f ; ; . 102—105
Aristida cyanantha : : ; : : : : 3 - 105—108
Andropogon monticola : ‘ : : : é ‘ : . 108—114
(zt) Summary or CuiEF CHARACTBRISTICS OF SPECIES DEALT WITH . . 115—117

CHAPTER III.
SUGGESTIONS REGARDING PractTicaL TREATMENT OF LOcAL GRASSLANDS.

Treatment with a view to Fodder-Production—Treatment with a view to
Afforestation j ; 3 ’ ; j 5 : : : ls —=i26

ui
LIST OF PLATES.

Plate 1—Map cf Locality.
I1.—Aristida cyanantha in typical habitats.

I11.—Saccharum Munja and Saccharum spontaneum var. nepalense in their
typical habitats.

IV. —Triraphis madagascariensis in typical habitat.

V.—Acacia Catechu and Dry Miscellaneous Forest in their typical habitats.
Wales)

VIL. |

VILL. }—ELrianthus Ravenne in typical habitats.

oe } —Saccharum Narenga in typical habitats.

XIII.—Loam-form of Saccharum spontaneum in typical habitat.
XIV.—Sand-form of Saccharum spontaneum,
XV.—Loam-torm of Saccharwm spontaneum and var. nepalense.

DEW AL, F
De WALIL ecchanum spontaneum.
XVIII. }
EXGLEXG
XX. >—Saccharum Munja.
Xx1.§
XXII.—Saccharum arundinaceum.
DOINULS 4
XXIV. fase arum Narenga.
xXxV.
a —Hrianthus. Ravenne.
XOXVIDE
XXIX. -—/mperata arundinacea.
XXX.
DXOXOXG]
XXXII. - —Vriraphis madagascariensis.
XX)
UXOXGNGIAV
NOON, ' —Aristida cyanantha.
XXXVI.
JOXOWIL,
XXXVITI. ¢-—Andropogon monticola.
XONGNGIONG:

X.L.--Gola Tappar, a local Grassland.

THE INDIAN FOREST MEMOIRS

VOL. I] 1911. [PART I

Some Indian Forest Grasses and their Cecology.

BW IRs S, IBIOUWE. TEOSSG, TEL. Go, RIES,

Imperial ForestService
Botanist at the Forest Research Institute, Dehra Dun, India.

INTRODUCTION.

1. Indian Forest Grasses are of considerable mmportance
importance on account of the value of the economic products yielded by particular species. Goer
Thus a considerable revenue is realised from the sale of such species as the Bhabar grass
(Ischemum angustifolium, a valuable paper material), Rusa Oil Grass (Cymbopogon
Martini, the source of the valuable Palmarosa or Rusa perfumery oil), Munj (Saccharum
Munja which yields a valuable textile fibre and also a paper material) and others. This
revenue will undoubtedly increase in the future as new industries arise and as the economic
value of the various species becomes better and more widely known.

On the whole, however, there is no doubt that the primary importance of our forest
erasslands depends on two facts :
(a) Their great value as grazing grounds and fodder yielding areas.
(b) The fact that where grass is of comparatively little value, these grasslands occupy
large areas which might produce valuable forest and which it is therefore
desirable to afforest.

2. As regards the value of our grasslands to vatue for
odder and

the people of India for grazing purposes the following figures will indicate their enormous Guaeies
importance. These figures show the average forest area in India and Burma which has
been annually open for grazing and the average annual number of buffaloes, cows, bullocks,

Grasslands
and Afforest-
ation.

2 INDIAN FOREST MEMOIRS.

goats and sheep which have been actually erazed in this area during the four years period
1904-08 :-—

a

NUMBER OF ANIMALS GRAZED ANNUALLY,

Ayea open to Grazing annually in

l
Square Miles. | Cows and | Goats and
| Buffaloes. Ballocke: | Sheep. TOTAL.
1 | |
— }- | ea 2
| | |
197,225 | 1,597,949 | 7,683,682 | 4,489,476 13,771,107

In many parts of India, with the increase in the numbers and in the prosperity of the
local population of recent years, the number of cattle for which grazing is required in the
Government Forests has steadily risen and the question of what measures are to be taken in
order to satisfy as far as possible the urgent needs of the people with regard to fodder and
erazing, without thereby unduly decreasing the area required for the satisfaction of the
wants of the country in respect of wood, fuel and other forest products, for the protection
of the headwaters of streams and for improving the water supply required for cultivated
tracts, is now one of great difficulty and of pressing importance.

In whatever way a solution of this question of securing a satisfactory allotment of
forest areas for these antagonistic interests is ultimately arrived at, it appears certain that
large areas of the greatest value as grazing grounds, or as areas productive of good fodder
grass, to meet present needs, or to serve as a reserve in times of famine, will always be
situated within the boundaries of the Government Forests and such areas will consequently
remain under the management of the Forest Department. It is also clear that with regard
to such areas the forest estate will not adequately fulfil its task of satisfying, to the full
extent of its possibilities, the needs of the people in respect of the produce which it is
capable of yielding, until these grasslands are managed in such a way as to make them
most productive of the best class of article required from them, i.e. until they are made to
yield the maximum quantity of the best fodder grass which they are capable of producing.

3. On the other hand, large areas of grass-
land exist in forests almost throughout India which are not needed primarily for the
production of fodder and which have been for many years carefully protected from fires and
grazing in the hope that they would soon become naturally reafforested, with, in many
cases, so far as can be seen, little or no practical result. Many of our forest grasslands
also are annually burnt under proper control in order to diminish the danger of fire damage
to the surrounding forest which would result from the accidental firing of the masses of
dry inflammable grass contained in these areas in the dry season. Little is known at
present regarding the effect, injurious or otherwise, of these annually repeated fires on the
production of fodder, or on the capacity of the soil to produce good forest growth. The
great importance of initiating an inquiry into the best method of treatment to apply to

Lea

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 3

our forest grasslands (a) with the object of improving the yield of fodder and (b) with the
object of hastening their afforestation having been recognised, the Forest Botanist was
directed in October 1908 to make a beginning with this work by studying the Savannah-
grasses of the Sal (Shorea robusta) forests in Northern India. The present paper includes
the results of the work which it has been found possible to carry out since that date, in the
intervals of heavy educational and other duties in connection with the organisation of the
Botany branch of the Imperial Forest Research Institute at Dehra Dun.

4. In order to obtain useful information
regarding the best method of treatment to be applied to grasslands with the object of
improving the fodder yield on the one hand, and of favouring their natural afforestation
with valuable forest trees on the other, an ecological study of the various grasses and woody

species concerned is necessitated.
5. As the importance of CEcology is often

insufficiently appreciated it appears desirable to explain, somewhat in detail, what is
implied by this term. Cicology has been defined as “ the science treating of the reciprocal
relations of organisms and the external world ( ovxos, a house, doyos, theory)! and this
has been further elaborated by one of the highest cecological authorities as follows :—

“It teaches us how plants or plant-communities adjust their forms and modes of
behaviour to actually operating factors, such as the amounts of available water, heat, light,
nutriment, and so forth.

A casual glance shows that species by no means dispose their individuals uniformly
over the whole area in which they occur, but group them into communities cf very varied
physiognomy. Cicology seeks—

(1) To find out which species are commonly associated together upon similar habitats
(stations). This easy task merely involves the determination or description
of a series of facts.

(2) To sketch the physiognomy of the vegetation and the landscape. ‘This is not a
difficult operation.

(3) To answer the questions—

Why each species has its own special habit and habitat,
Why the species congregate to form definite communities,
Why these have a characteristic physiognomy.

This is a far more difficult matter and leads us—

(4) To investigate the problems concerning the economy of plants, the demands that
they make on their environment, and the means that they employ to utilise
the surrounding conditions and to adapt their external and internal structure
and general form for that purpose.”

Having therefore obtained a full knowledge of the cecology of a particular species, such
as the Sal tree, and being able to fully explain why the species thrives under certain condi-

! Geology of Plants. By K. Warming, Eng. Edn., 1909, p, 1.
? Warming, /.e., p. 2.
eh

ecological
Study of
Forest
Grasses
required,

(Ecology
defined.

4 INDIAN FOREST MEMOIRS.

tions and why it cannot exist under others, we shall be able to answer such questions as
whether or not the species can thrive in localities where it does not at present exist, and if
so what are the steps to be taken to make its successful introduction possible; also why the
growth or reproduction of the species is unsatisfactory in a particular locality and what
steps can be taken to counteract its diseases. In other words we shall know what treatment
must be adopted in our Sal forests in order to secure their most satisfactory development
and reproduction and what steps must be taken to extend the area of such forests by intro-
ducing the species where it does not at present exist.

While recognising therefore the vital importance of this study the magnitude and
difficulty of the work must be apparent to the most casual observer; involving as it does a
detailed knowledge of all the living and non-living factors of a plant’s environment, of the
way in which, and of the extent to which, each such factor is capable of influencing favour-
ably or unfavourably the growth and development of that plant and of the extent to which
the favourable or unfavourable effect of these factors may be increased or diminished by
other factors.

These points may be illustrated by a few simple cases : a plant A, for instance, is under
ordinary conditions of its habitat able to kill out a plant B by its competitive action and
this explains the absence of plant B in certain localities. In precisely similar habitats,
however, which are periodically fired, plant B is found to thrive and A to be absent, or
comparatively rare, and this is explained by the fact that plant A is more directly injured
by fire than B, or that fire has caused a change in soil and moisture supply more unfavour-
able to A than B. Firing, in consequence, is a simple treatment to adopt in order to favour
the development of B in such localities where A occurs. Again, species B may be unable
to exist on sandy soils owing to the small capacity of such soils for retaining moisture.

his capacity is greatly increased by an admixture of humus and the introduction of a
species A capable of thriving on dry sandy soil and of producing large quantities of humus
may be the best step to be taken with a view to the ultimate introduction of species B.

In another case a species A may be unable to exist in clay soil owing to the supply of
oxygen being insufficient for the healthy action of its roots. Species B which is able to
thrive on clay soils and is less sensitive as regards the supply of oxygen may be capable of
improving such soils by the drying action of its roots and by the formation of humus,’
and hence the introduction of species B may be the best procedure to adopt with a view to
the ultimate establishment of species A. Again the available nitrogen in a forest soil
depends largely on the rapid decomposition of the humus by bacteria and any factor which
retards the growth and development of these bacteria may result in so impoverishing the
upper layers of soil in respect of nitrogen that the seedlings of a particular species may be
unable to grow in them and the reproduction of that species thus becomes impossible.

The production of good seed and vigorous seedlings also in many cases depends on the
existence of the birds and insects by which the cross-pollination of the flowers is effected

Lise Humus acts as a weak cement and holds together the particles of soil, thus it serves both to bind a coarse-grained sandy soil,
and, by forming aggregates of the finest particles, to render the texture of a clay soil more open.” Zhe Soil by A. D. Hall, 1908, p. 47.

esa

HOLE: SOME INDIAN GRASSES AND THEIR GiCOLOGY. 5

and hence the factors which influence the healthy development and reproduction of such
insects and birds may determine the area within which a particular plant is able to exist.

Finally the absence of a plant in localities suitable for its development may be
accounted for by the simple fact that its seeds have not yet been able to reach those localities.

The importance of the direct injury inflicted by parasitic plants, such as fungi, by
injurious insects, or by browsing animals such as goats, in rendering the existence of a
particular plant in a certain locality impossible, is more obvious and needs no emphasizing
and these remarks will make it clear that the complete cecological study of a particular
species really comprises a complete knowledge of its physiology and pathology.

6. The fact that, to a certain extent, a know- Importance

: ° . of Morpho-
ledge of the physiology of a plant can be obtained by a study of its morphology and anatomy igloo

is of great importance in cecology. Thus the morphological characters of the fruit may Charetars
indicate the agency by which the seeds are distributed, such as wind, water, animals, and‘ “°°°*"
so on; similarly the characters of the flower may indicate the agency by which pollination

is effected, such as wind or insects, while the morphological and anatomical] characters of

the leaf and the habit of the plant may indicate whether it is adapted to exist in a sunny

or shady, dry or wet locality.

By continually tracing the connection existing in different localities between such

characters of the vegetation and the factors of the environment, a mere inspection of the
vegetation existing in localities, not hitherto visited, eventually suffices to give a fairly
accurate idea of the chief factors influencing plant growth which exist there and thus
enables us to decide whether or not a certain species will be likely to thrive there. The
accuracy of such conclusions is greatly increased by a study of so-called indicator-plants.
If a species A, for instance, is invariably found on soil which is known to be suitable for
the vigorous development of species B, it is clear that we may regard the existence of A as
a direct indication of a locality suitable for B, at all events so far as the characters of the
soil are concerned.

Grasses are particularly valuable in this respect and a careful study of the grasses
in forest blanks, in many cases, indicates to the Forester whether or not the conditions of
soil and moisture are such as to render possible the afforestation of such blanks with a
particular species, and similarly the existence of other grasses may indicate that the soil
has become unsuitable for a certain species and may thus explain why reproduction can no
longer be obtained in those forests.

7. As the morphological and anatomical eating

; s of
characters of the vegetative organs, as well as the habit of plants generally, show the most Ecological

ncticeable and striking variations in response to differences in the quantity of moisture ce
available for growth, cecologists up to date have, as a rule, concentrated their attention
primarily on this factor and have made it the basis of their cecological classifications.

Thus Professor Warming notes: “ Water is the condition of life that exercises the
greatest influence in bringing into being external and internal differences among plants;

[28 li

Hydrophytes,
Xerophytes,
Mesophytes.

6 INDIAN FOREST MEMOIRS.

it is likewise water that plays the leading part in determining the creation of plant. com-
munities and their distribution over the soil.”* He accordingly classifies plants into—

1.—Water- or Aquatic- | I1.—lLand or Terrestrial-

Plants= Hydrophytes . “ Plants that pass the Plants : . Those “ that expose at least
whole or the greater their assimilating organs to
part of their lives sxd- the air and hence to trans-
merged in the water, piration.””? Those are sub-
which envelops them divided into—-

completely or, at most, (a) Aerophytes.—“Those
leaves definite floating _ species that are adapted
parts of them un- | to meet the conditions

covered at its sur-
face,”

of strongest transpira-
tion and most precari-
ous water supply.”
(6) Mesophytes.—“ The re-
mainder.’”4
It is, however, pointed out that there is no sharp boundary between these groups “ for
there is a group of plants, marsh-plants (helophytes), which, like water-plants, develop
their lower parts (roots, rhizomes, and, to some extent, leaves) in water or at least in soaking
soil, but have their assimilatory organs mainly adapted to existence in air, as is the case
with land-plants to which they are closely allied,”’ while “ between these two classes
(xerophytes and mesophytes) there is of course no strict. boundary.” It should be noted
that while Warming includes marsh plants in terrestrial plants,’ Clements classes them
as hydrephytes.° For the present these plants may conveniently be considered as Hydro-
phytes.
Some idea of the kind of plants included in the above groups will be obtained from
the following enumeration of a few of the more obvious characteristics of some typical
plants :—

Hydrophytes including Marsh plants.

Roots are usually reduced. Root-hairs often absent. Owing to the buoyancy of water
there is comparatively little need of strengthening mechanical tissue and submerged
plants include no woody species. Owing to the slow diffusion of oxygen in water, all sub-
merged parts of plants are especially liable to the dangers of an insufficient oxygen-supply
for respiration and in consequence large and abundant air spaces in such parts are very
characteristic and serve for the storage of oxygen and for the conduction of this gas from
parts in the air to those in the water or saturated soil. The presence of these air spaces
can often be detected by the naked eye.

Ut 5 Sle,
2 p. 96.
Ton 0 Oe

pp we dlOL,
B59 jd USL,
J A jee Onis
7 p. 97
8

Plant Physiology and Ecology by F. E. Clements, 1207, p. 165.

Ord

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 7

The supply of oxygen can also be augmented by increasing the absorbing surface of
the plant which is in contact with the water and hence submerged leaves and, in some
algee, the entire plant are often cut into a number of long narrow segments.

Oxygen being usually more plentiful in running than in stagnant water, plants in
running water tend to show comparatively less development of aerating devices.

The presence of aerating tissue is also of utility in increasing the buoyancy of the
plant.

In woody marsh plants the so-called pneumatophores, or aerating roots, are often
developed as an adaptation to meet the danger of a deficient oxygen-supply.

To some extent also the characteristics of the plants of this group are due to the
weakening of the light in water and submerged parts of stems and leaves tend to become
long and thin resembling those of terrestrial plants developed in shade.

The epidermis, especially of submerged parts, as a rule exhibits no devices for reduc-
ing transpiration such as a thick cuticle, covering of hair, wax, etc., and hydrophytes, when
entirely removed from water, usually dry and wither with great rapidity.

As examples of this class of plants we may take aquatic Algee or “ sea-weeds.”

rapa bispinosa, the water Chestnut, with its remarkably branched submerged
leaves with filiform segments.

Nelumbium speciosum, the Sacred Lotus, with the enormous air spaces in its sub-
merged creeping stem.

Xerophytes.

The root system is usually strongly developed. Adaptations for reducing transpira-

tion are very common and characteristic and amongst the most usual devices resorted to
are—

(1) A reduction in the surface of those memhers which are, as a rule, most special-
ised with the object of facilitating transpiration, viz. the leaves. Thus
some xerophytes are characterised by the relatively small size of their leaves
and leaflets and others dispense more or less completely with leaves and place
their chlorophyll tissue in their stems or branches.

(2) Development of a thick protective cuticle and often also of thick walled scleren-
chymatous tissue beneath the epidermis which interfere with the free exchange
of gases between the outer air and the moist thin-walled chlorophyll tissue
beneath them. Coatings of wax and hairs serve the same purpose. The
latter can as a rule be easily recognised by eye and the former anatomical
characters are frequently indicated in the field by the firm hard texture and
also by the colour of the foliage, the colour of those leaves in which the chloro-
phyll is protected by thick-walled external cells usually being like that of
leaves which are covered with a thin coating of wax, vzz. a dull, or bluish-
green, as contrasted with a fresh bright green.

et

8 INDIAN FOREST MEMOIRS.

As the stomata are the openings through which gaseous exchange chiefly takes:
place the protective coverings, e.g. of hairs, tend to occur chiefly on the
under-surface, where the stomata in land plants mainly occur.

(3) As cork forms an excellent protection against free gaseous exchange a thick
corky bark is often characteristic of woody plants, the water current ascend-
ing in the wood thus being protected.

(4) The loss of water by transpiration from a leaf depends greatly on the angle at
which the leaf surface is exposed to the sun’s rays, the loss, other things
equal, being greatest when the leaf surface is perpendicular to the direction
of the sun’s rays and least when the surface is parallel to this direction, e. g-
when the leaf exposes its edges only to the direct rays of the sun.

Thus motile leaves and leaflets are exceedingly common and characteristic
and in many species can be recognised at once by the presence of the so-called
pulvinus, pulvinule and glands.

(5) The presence of ethereal oils and latex is believed to reduce transpiration, but
in what way is not at present clearly understood.

(6) Water-storing tissue is often characteristic. Plants obtain their water when
they can get it, even when it is not needed for immediate requirements, and
then store it in these reservoirs where it is available for use when wanted.
The presence of such tissue also diminishes transpiration by checking gaseous.
exchange and the conduction of heat.

The presence of such tissue is often indicated by a succulent, or soft, and
somewhat spongy, texture and by the exudation of water when fresh tissues
are cut.

The above will suffice to indicate a number of familiar plants to Indian Foresters
which, according to prevailing cecological ideas, must from their morphological and
anatomical characters be considered as fairly typical xerophytes, e.g. the desert Prosopis
spicigera with its extraordinarily deep-going root system; the practicaily leafless species of
Tamaria, Casuarina, Euphorbia, Opuntia, Ephedra and Orthanthera; Calotropis with its
glaucous green leaves woolly beneath, Pinus with bluish, or dull green, foliage, the texture
of which is firm and hard and which is characterised by a well-developed cuticle and super-
ficial sclerenchyma: Moringa and Erythrina with thick corky bark; Acacias with motile
leaflets; Euphorbia and Calotropis with milky juice, and many dry locality Rutaceew with
ethereal oils, e.g. Feronia and Agle.

Mesophytes.

Warming notes that these plants “ grow on soil which is of an intermediate character,
and is neither specially acid, nor saline, but is moderately moist, usually well ventilat-
ed, also rich in nutriment and in alkaline humus or in other organic constituents. Meso--
phytic communities occur in very diverse climates, near the Poles or on the equator, yet:

cee

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 9

they can never be exposed to the danger of prolonged drought. Adapted to such conditions
are plants that show a relatively weak development of the above-mentioned arrangements
for regulating transpiration: in this respect these plants stand midway between hydro-
phytes and xerophytes. The leaves are large, and far more varied in form than in xero-
phytes; teeth and other incisions of the margin are common, as are compound or richly
divided leaves; hydathodes seem to be frequent; the vegetative organs are of a fresh green,
and devoid of thick grey coatings of hair or bluish incrustations of wax; the leaves are
usually dorsiventral in structure. Stomata are numerous, often occurring also on the
upper face of the leaf; anatomical peculiarities, such as aqueous tissues, are very rare, or
at least not extremely developed.”*

Among the most typical forest trees of this group is the European Beech and the
Indian tree which is probably most nearly allied cecologically is the Sal.

The anatomical and morphological characters of typical xerophytes may be described
by the adjective xerophytic and the same term may be used for a locality where xerophytes
grow, or for a plant group consisting of xerophytes.

The adjective verophilous(=living in, or suited to, a dry habitat) derived from the
corresponding substantive werophile, is also frequently used. Similarly we have the terms
hydrophilous and hydrophytic, mesophilous and mesophytic.

8. The most important natural plant group, rhe rorma-
or aggregate of individuals, at present recognised in cecological botany is the formation oe
which may be considered as corresponding to the genus of systematic botany. Warming
defines the formation “ as a community of species, all belonging to definite growth-forms,”
which have become associated together by definite external (edaphic or climatic) characters
of the habitat to which they are adapted. Consequently, so long as the external conditions
remain the same, or nearly so, a formation appears with a certain determined uniformity
and physiognomy, even in different parts of the world, and even when the constituent
species are very different and possibly belong to different genera or families.

Therefore—

A formation is an expression of certain defined conditions of life, and is not con-
cerned with floristic differences.”* The principal formations recognised by Warming are
the following which he classifies as shown :—

Hydrophilous—Formations in marsh and water (Hydrophytes and Helophytes).
Xerophilous Formations on sour (acid), cold and saline soil, formations on rock,
sand, gravel and waste land, bush or scrub formations and formations on

desert, steppe and savannah.

1 Lc, p. 135.
2A Pear ieconm is a plant characterised by its habit (eg. tree, shrub, herb, gregarious and sporadic plants, etc.), its period of
vegetative activity, of flowering and fruiting, the duration of its life cycle, by the form and nature of its flowers, fruits, stems, leaves and
roots, in so far as such differences can be ascribed to the influence of the conditions of the plant’s environment,
3 ].c.. p. 140.
[ 9 }

Definition of

Steppe and
Savannah.

Steppe—Is a “ Treeless plain clothed with grasses and other perennial herbs.”*
“The vegetation is xerophilous in character, and does not form a dense, close

which is suitable for good forest growth.

INDIAN FOREST MEMOIRS.

Mesophilous——Good examples of such formations are “ meadows ” and beech, oak

and birch forest of temperate climates. Warming also includes in this group
the deciduous (=monsoon forest) and evergreen forests of India but a more
detailed knowledge of these formations would probably modify this arrange-
ment.

The Indian Forester will at once recognise the very close relationship exist-
ing between desert and the formations of such trees as Sterculia wrens and
Boswellia serrata which nevertheless come under the head of deciduous
forests, while on the other hand some species of the evergreen forests such as
Carallia integerrima, Cyclostemon assamicus and Calamus tenuis occur in
swamps in the deciduous sal forests, and thus form a connecting link be-
tween the mesophilous and hydrophilous formations.

Such terms as marsh, desert and scrub need no definition but the terms steppe and savan-
nah are of value in the description of our Indian forest grasslands and they have been
defined as follows :— :

carpet. Both these features serve to distinguish grass-steppe from the allied
formation, meadow, whose close vegetation consists of fresh-green, soft-leaved,
and broad-leaved grasses and other perennial herbs. On the other hand,
vegetation clothing grass-steppe is closer and taller than that of desert.”

The importance of steppes to the Forester lies largely in the fact that they
are usually considered to be an indication of conditions unsuitable for tree
growth which is naturally of great importance as regards the question of
afforestation.

Savannah.—Schimper defines this as “ xerophilous grassland containing isolated

trees,’* and Warming notes that it differs from steppe “essentially in the
larger dimensions of the constituent plants, and above all, in the presence of
trees. As an excellent example of a savannah we may take the Munj
Grass (Saccharum Munja) formation which occupies enormous areas in
Northern India.

It is very doubtful whether all savannahs can be classed as xerophilous.
Erianthus Ravenna, for instance, forms typical savannahs in Northern India
and this species cannot be classed as xerophilous. With the exception of
Bamboo formation, indeed, savannah probably represents the highest type of
grassland and shows grasses growing under the most favourable conditions
of soil and moisture and thus usually indicates a condition of these factors

1 Warming, L.c., p. 281.
? Plant-Geography, 1903, p. 162.
$'lic., p- 2935

Lope

HOLE: SOME. INDIAN GRASSES AND THEIR GCOLOGY. 11

9. Warming further recognises an ASSOCIA- rhe ata
TION as “ a community of definite floristic composition within a formation; it is, so to speak,
a floristic species of a formation which is an ecological genus.” A forest, for instance,
obviously differs according to its constituent species and this author thus recognises deci-
duous dicotylous forest in the temperate climate as a mesophilous formation of which
Beech Forest, Oak Forest, and so on, may be considered different associations. He also
notes that, as different beech forests are not always identical in character, we may recog-

nise varieties of an association.

10. The classification adopted by Schimper Sehimper’s
differs in some respects from that detailed above. This author also distinguishes Aquatic tion.
Plants (Hydrophytes) from Terrestrial Plants but then proceeds to classify the latter as Teonontiocsa:
follows :—

“Contrivances for expediting the exit of water are characteristic of hygrophytes, or
plants whose conditions of life exclude ali danger of desiccation, and in which a stagnation
of the water, which brings nutritive salts to the parts requiring them, may be feared. On
the other hand, difficulties in obtaining a supply of water lead to the formation of devices
for assisting absorption and limiting transpiration; werophytes are provided with contri-
vances of this kind.’

He also places in a third category—

“all plants whose conditions of life are, according to the season of the year, alter-
nately those of hygrophytes or of zerophytes. All such plants, including, for
instance, the great majority of the plants composing the Central European
flora, should be termed tropophytes. The structure of their perennial parts
is cerophilous, and that of their parts that are present only in the wet season
is hygrophilous.”*

He then distinguishes—
Rain-forest (= Evergreen forest) as a hygrophilous formation of tropical woodland
and
Monsoen-forest (= Deciduous forest) as a tropophilous formation of the same.
Schimper’s hygrophytes and tropophytes as a whole are included in Warming’s mesophytes.
The Indian Forester will appreciate the wide separation of the tropical evergreen from the
deciduous forests of Schimper’s scheme.

11. Warming endeavours to carry classifica- Warming’s
tion a step further by grouping all formations under a series of classes of which the follow- OER
ing are examples :— noes

Psammophytes (formations on sand and gravel),

Eremophytes (formations on desert and steppe),

Halophytes (formations on saline soil).

Camp HLA:
SUC 10) 24.
JG. Tos BY

ri 7

Points which
require con-
sideration in
eological
Classifica-
tion.

Primary Sub-
division into
Grassland
and Woodland
advisable.

12 INDIAN FOREST MEMOIRS.

Sclerophyllous formations (bush and forest),
and so on.

These classes are frequently very difficult to distinguish and we may obviously have
precisely the same formation in a desert (caused by insufficient rainfall) and in a moist
climate (on very dry sand or gravel). The Tamariz thickets on sandy river banks in the
moist Indian region of deciduous forests and in the desert tracts of the Panjab may be
taken as an example. Similarly saline soil formations may be found on non-saline sand,
heavy clay and so on.

Butea frondosa woodlands, for instance, occur both on saline and heavy soils, Tamarix
on saline and sandy soils, and so on. Again much of the woody growth on sand, in almost
desert tracts and on saline soil, might be classed as sclerophyllous bush. A detailed classi-
fication on these lines therefore in the present state of our knowledge tends to become con-

fused and difficult to apply in practice.
12. In endeavouring to draw up an accept-

able scheme of cecological classification the following points demand attention :—
Professor Warming himself clearly states that “the cecology of plants is a subject
still in its infancy; numerous investigations must be made before the found-
ations can be truly and rightly laid, and before a consistent, clear, and natural
classification of plant-communities is achieved.”

For the present, the main desideratum is undoubtedly a lack of accurate descriptions
of the communities which can be readily distinguished in different countries, a lack of
complete and accurate information regarding the factors which compose the environment
of such communities and want of information regarding the factors of dominant importance
which regulate the existence and distribution of the communities. |

As a basis for such work it seems undoubtedly preferable to adopt the two great classes
of vegetation which Schimper terms whodland and grassland; in the former woody plants
give the character to the type of vegetation and in the latter herbaceous plants and grasses.
Schimper also distinguishes desert but here also woody and herbaceous plants exist and the
desert can be regarded as the lowest form of grassland or woodland, according to the
character of the majority of the plants, and does not need recognition as a separate type of
primary importance.

Schimper well says that “woodland and grassland stand opposed to one another like
two equally powerful but hostile nations, which in the course of time have repeatedly fought
against one another for the dominion over the soil,”? but the real significance of the
relations between these great types, at all events from the practical point of view, appears
to lie in this, that, for almost all forms of grassland, there is a corresponding form of wood-
land and the Forester therefore can often find in the nature of his grasslands the answer

to the question of what type of woodland can be introduced with the greatest prospect of
success.

hs 105 We
2 L.c.,\p. 162:

fae

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 13

13. It must be clearly understood that only madvisavie
: e,e ° at present to
after a far more detailed knowledge of plant-communities has been obtained than we at distinguish

, Iinor @eolo-

“present possess will it be possible to adequately appraise their relative importance and thus gical Groups.
to evolve a really satisfactory classification and for this reason it seems impossible and
inadvisable to insist on any distinction at present between formation, association, and
minor groups. All that seems immediately necessary is that those cecological groups which
exist and which can be readily recognised in different localities as distinct, either on
account of a different physiognomy, or habit of growth (tree, shrub, erect, climbing, etc.), a
different period of vegetative activity, of flowering or fruiting, a different admixture of
species, or on account of marked differences in the dimensions attained, in the density of
stocking, state of reprceduction, and so on, should be separately described and, for the
present at all events, these may conveniently be termed communities. As our knowledge
increases these can be subsequently aggregated into larger groups or disintegrated into
smaller groups, as may seem advisable.

14. Every type of vegetation is to a greater Or Formations
Jess extent able to increase the quantity of soil available in a locality and to increase the neesseeely
capacity of that soil for retaining moisture through the addition to it of humus. Rocks ?"™"""™
-are gradually weathered and broken up by the mechanical action of heat and cold and the
chemical action of the atmosphere and water. Plants also to some extent aid in
this work of decomposition by keeping the rocky surface moist, by the eroding action of
their absorbing roots and by the mechanical action of those roots which penetrate into cracks
and crevices and which by their continued growth are able to split the rock. Plants how-
ever are of still greater importance in this respect owing to their power of accumulating in
their neighbourhood wind and water-borne dust and débris and of resisting erosion. It is
thus clear that the continued existence of plants in a locality may alone be sufficient to
convert a xerophytic into a mesophytic habitat and the gradual covering of naked rock
with luxuriant vegetation through a succession of (1) Algee, lichens and mosses, (2) Xero-
philous grassland or woodland, and (3) Mesophilous grassland or woodland has been fre-
-quently emphasized by cecologists.

Again plants which are able to resist the injurious effect of a considerable degree of
shade and which are in consequence able to establish themselves on the ground already occu-
pied by other plants frequently have more moisture available for their growth and develop-
ment than those plants which establish themselves directly on bare unoccupied areas,
owing to the absence of shade and humus in the latter.
Shade-bearing species therefore are frequently hygrophilous, or mesophilous, and are
able to enter into the composition of what must be regarded as a higher type of vegetation
than that which is usually constituted of light-demanders.
A succession which thus proceeds from a xerophilous to a mesophilous and finally a Progressive
hygrophilous type of vegetation, 7.c. from a simple to what must be regarded as a more rene
highly developed type, may be termed a progressive succession. Such a succession may

[ 138 7

Regressive
Succession.

14 INDIAN FOREST MEMOIRS.

probably be recognised in (a) the xerophilous light-demanding woodlands of Acacia Catechu
in the stony, sandy, riverbeds of Northern India, (>) the dry miscellaneous forests of the-
same region, in which the proportion of A. Catechu has diminished and a number of addi-
tional species have made their appearance of which several have a marked power of increas-
ing the humus content of the soil, ¢.g. Ehretia levis, Holarrhena and Adhatoda Vasica.
Such woodlands are clearly less xerophilous than the pure woods of Acacia. (c) the markedly
mesophilous shade-bearing Sal forest in which the sal predominates. This type of vege-
tation also extends, in parts of Bengal and Assam, into the region of the hygrophilous
tropical evergreen forest which must be regarded as the highest type of woodland.’

On the other hand the reverse succession from a highly developed to a more simple type
may be termed regressive. An example of such a succession is seen when mesophilous
forest is cleared, or more gradually destroyed by fire and grazing, on steep slopes, the:
resulting erosion converting the area into a rocky hillside only capable of supporting the
poorest and most xerophilous types of vegetation. Fire is a very potent factor in causing
regressive successions, for not only is it capable of temporarily depriving the soil more or
less completely of its covering of vegetation, but it also directly dries the soil and destroys
the humus. Fire may in this way be responsible for the existence of xerophilous grassland,
or woodland, in localities which once supported mesophilous, or possibly hygrophilous,
vegetation. .

Grazing again, by destroying the undergrowth and keeping a forest open, may so reduce.
the humus content of the soil as to render impossible the reproduction of the mesophilous
species constituting the forest and may thus cause a regressive succession.

The effect of artificially clearing an area of its covering of vegetation in this connec-
tion is often insufficiently appreciated although this is of vital importance to the Forest
Officer. The entire removal of the tree-growth on an area, such as takes place under the
treatment of clear-cuttings, or coppice-fellings, results in the drying of the soil, the rapid
disintegration of the humus already contained in it and a temporary interruption in the
supply of additional humus. In this way the capacity of a sandy soi! to retain moisture
may be greatly reduced, while a clayey soil becomes denser, less aerated, liable to water-
logzing in the rainy season and to become very dry at other seasons owing to the moisture
which is evaporated not being replaced rapidly, on account of the sluggish movements of
water by capillarity in the dense soil. Conditions may therefore be thus created which are
unsuitable for the development of young plants of the species which composed the original
woodland. It is therefore easy to understand from this point of view why, in Satin

1 Nothing said above regarding the details of a succession which under certain circumst S i
succession must necessarily always follow the same sequence, or that a Sal forest, for RUM Eo are ee a ae enttee
occupied by Acacia or certain other miscellaneous species. On sandy soil in the locality dealt with in the present paper ri Se geet
of the succession described above can be clearly recognised, but, on the other hand, it is certain that all the Sal fords OF ‘the G ©) oe
not originated in the same way, There is, for instance, little doubt that the Sal can naturally establish itself in this lo lity locality, nave
sandy or well drained clayey soil, provided sufficient moisture is available by percolation or otherwise, (2) on nde Lars ia i) ditectlagen
plants like Adhatoda has considerably increased its humus content, (3) on clay which has supported a growth of plants lik mi aie ae bee
Trewia nudiflora, and others. The texture of the clay is improved by the draining action of the roots of these plants - db epee EES
of humus so that the aeration of the soil is improved and the movement of water from the subsoil to the surface Te canes S ee
y capularity ae ed.

le te. 4

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 15

localities and for certain species, methods of treatment which result in the more or less
complete removal of the vegetative covering are unsuitable, while a method of treatment
like the Selection System under which the exposure of the soil is reduced to a minimum is
the most successful.

A point, also, which is frequently not appreciated is that, by making clearings in the
middle of woodland, conditions are often created which are quite unsuitable for the healthy
development of the species composing the surrounding woodland and which are caused by
the absence of free air-circulation.

The grassy plains, so common in the Sal forests of India, frequently illustrate this
point. These areas are usually clearings which were originally made in the forest for
cultivation and then abandoned. Owing to the absence of air-circulation caused by the
surrounding forest belt, such areas are intensely heated by the sun in the day and hot season
and cooled by radiation at night and in the cold season. In dry localities this action is
intensified and in such places the stcol shoots, springing from any stumps of the felled
trees which happen to have been left in the ground and to have survived, are annually cut
back by frost, while the absence of humus and dryness of the soil prevent the establish-
ment of new seedlings. The natural re-establishment of the mesophilous Sal forest is
therefore rendered impossible, except very slowly in the narrow belt immediately adjoining
the forest. The Forest Officer who wishes to quickly afforest such areas, therefore, will be
most successful if he recognises the fact that a regressive change has taken place, that a
habitat which recently supported mesophilous vegetation has now become strongly xerophy-
tic and that xerophilous and frost-hardy species like Acacia Catechu should accordingly
be introduced.

Coppice fellings in the middle of forest growth may similarly cause a regressive
succession.

Finally there is a type of succession which we may distinguish as paraliel succession.

Types of both grassland and woodland are found in all kinds of habitats, ranging from
the most xerophytic to the most hygrophytic, and it is of great importance for the Forest
Officer to realise that, for each type of grassland, there is, as a rule, a corresponding type
of woodland capable of thriving under similar conditions of the environment, seeing that
this has a direct bearing on the question of afforesting grasslands. When a type of grass-
land, such as Munj savannah, is replaced by a parallel type of woodland, e.g. dry miscel-
laneous forest of Acacia Catechu, Dalbergia Sissoo and others, we may therefore regard it

as a case of parallel succession to distinguish it from the progressive and regressive changes
considered above.

Parallel changes can be frequently efiected more easily and rapidly than progressive
changes and with reference to such questions as the afforestation of grasslands and the
extension of woodlands parallel changes are probably as a rule of more practical import-
-ance.

[ 15 }

Parallel
Succession.

16 INDIAN FOREST MEMOIRS.

15. It may be asked that if such parallelism
really exists between grassland and woodland as has been indicated above and if the:
existence of a certain type of grassland in a particular habitat really indicates that that
habitat is suitable for a certain type of woodland, how is it that we do not find this wood-
iand already in existence there and how is it that the grasses are able to remain in posses-
sion? The answer to this question is to be found, it is believed, in the exceedingly efficient
mechanism for seed-distribution by wind, and probably also to a considerable extent by
water, possessed by grasses by means of which they are frequently enabled to reach un-
occupied ground before woody plants. Anyone who has watched the wind blowing across.
a mun) savannah, several miles in extent, and wafting the myriads of tiny cottony fruits
in every direction far and wide, finds no difficulty in realising the undoubted fact that to
a great extent the banks of alluvial deposits thrown up by the rivers, although suitable for
the growth of such trees as Khair (Acacia Catechu) and Sissoo (Dalbergia Sissoo),
are first seized on and occupied by the munj. Once grassland has thus gained
possession it is obvious that the establishment of woody plants becomes more
difficult seeing that their seedlings have to face a competition with the grasses
both in the soil for moisture and in the air for light. A very clear case where
an area of recent alluvium has been first colonised by munj in this way but from which it
has later been driven out again by the khair has been seen by the writer in an area at the
foot of the southern slopes of the Siwaliks near Mohan. In part of this area munj 1s still
dominant and vigorous, but young plants of khair are just appearing scattered here and
there, in other portions the khair are more numerous, larger and older and many of the
munj clumps between them can be seen dead and dying, while elsewhere a dense pure pole-
wood of khair has become established under the shade of which can still be seen the decayed
remains of the munj clumps which had first colonised the spot.

The same thing has been observed in other types of woodland and grassland and it is
believed that, as a general rule, woodland soon succeeds in nature in ousting the correspond-
ing grassland from its habitat unless this is prevented by local factors such as fire, grazing,
frost, or by factors which prevent the formation and efficient distribution of good seed.

In consider- 16. It must never be forgotten that the-

ing the adap-

tations and principal problem which cecology has to solve is to supply an answer to the question “why
distributi ; ;: ; ; ; A ; : :
of Plants it is Cac species has its own special habit and habitat.” CE&cologists in regarding water as the

told aivine actor which “ plays the leading part in determining the creation of plant-communities and

avoid givin

uniue pso- their distribution over the soil” are apt to lose sight of the fact that a plant’s environment

g single is composed not of a single factor but of many factors and that each plant must be perfectly
adapted not only to the available water supply of its habitat but also, and no less perfectly,
to the conditions of light, temperature and other factors to which it is exposed. Hence
although we see that a plant is clearly adapted as regards its periodicity, habit of growth
and structural peculiarities to survive in a habitat where certain moisture-conditions

prevail, we must guard against the idea that the available water supply has necessarily”

Ee)

HOLE : SOME INDIAN GRASSES AND THEIR C2COLOGY. ai

been the dominant factor responsible for the fact that that plant is only able to exist in
certain habitats and not in others, 7.e. for its distribution as we now find it.

Differences in the amount of available water for instance offer no obvious explanation of Necessity of
. : : s : _ avoiding
such remarkable facts as that, in the deciduous Indian forests, on the same soil and in the Supra-maxi-

5 - : : is a “ ' . mal Tempera-
same climate, some species like Schleichera trijuga, Bassia latifolia, and others, come into ture may

full vegetative activity in the hottest dryest season of the year, while the majority, like Teak, Beton ad
wait until the near approach of the monsoon insures an easy and plentiful water supply. Pericaicity.
If the water supply is here the factor of dominant importance what advantage can have
been gained by the plants which first showed a tendency to become active in the hot
season, which can have accounted for the evolution and survival of such species? No such
advantage can I think be shown from the point of view of the water supply, but there is a
real advantage from the point of view of temperature. Transpiration undoubtedly reduces
the temperature and it is quite possible that some species avoid the risk of a rise of tempera-
ture above the supra-maximal by transpiring actively during the most dangerous season of
the year, although other species are able to exist with different periods of activity, owing
to their being less sensitive to injury from high temperatures, to their possessing more
adequate protective arrangements during the resting season, and so on.*

A most remarkable fact which possibly indicates the importance of active transpira-
tion during the hottest season is that in the swamps of the Dehra Dun valley, at an
elevation of less than 2,000’ above Mean Sea Level, the following temperate species occur,
which are usually found in the Himalayas at an elevation of 4,000’ to 8,000’, and this may
be due to the fact that vigorous transpiration is possible in the hot season owing to the
abundance of water available :—

Hedera Helix which often ascends to 9,500’ in the Himalayas and Quercus incana.

17. Again if we consider the danger from Xerophilous

. os . Characteris-
infra-minimal temperatures, no Forester who has had experience of the forests of Central ties, general.

and Northern India would be inclined to doubt the immense importance of frost as a factor tale
influencing the distribution of trees and it is a remarkable fact that almost all the adapta- he pas

tions ncted in paragraph 7 above as tending to reduce transpiration and to be characteristic (re¢"
of xerophytes also tend to reduce liability to injury by frost and low temperatures. The
reduction of the radiating leaf surface and the movements of leaves and leaflets reduce
radiation, while protective coverings of sclerenchymatous tissue, cuticle, cork, wax and
hairs all protect the living tissues from contact with chilled external air.

It should be noted also that two of our most typical Indian xerophytes are also two of
our most frost-hardy species, viz. Acacia Catechu and Zizyphus Jujuba. Just as these
adaptations moreover may be of service in affording protection from extreme cold, to

scarcely less an extent may they be of service in affording protection from extreme heat.

1 Plant-competition is also a factor which may possibly be responsible for the evolution of the remarkable periodicity noted above,
Species capable of growth at seasons which are unfavourable for the majority of plants may, on this account, be able to survive and hold
their own in the struggle for existence with more vigorous plants, the period of vegetative activity of which, however, is either shorter
or occurs altogether at a different season.

[ Ww ]

Importance
of light.

18 INDIAN FOREST MEMOIRS.

The presence of ethereal oils also often appears to have a remarkable effect, at all
eyents in regard to low temperatures, and during the phenomenal winter of 1904-05, when
all indigenous woody plants suffered more or less severely in Dehra Dun, the plants which
showed the most conspicuous immunity were the exotic Cinnamomum Camphora and
Eucalyptus citriodora.

The fact here emphasized that the so-called xerophilous adaptations may be of great
utility in enabling a plant to withstand extremes of temperature is obviously of practical
importance, inasmuch as xerophilous plants are in consequence the most suitable species
to introduce in localities not only where there is a scarcity of available water and a pro-
bability of damage by drought but also where there is a danger of injury by extreme heat
or cold.

In this connection it must be remembered that the mere presence of water in the habi-
tat of itself directly tends to reduce the danger from extreme temperatures. The presence
of water vapour in the air tends to reduce both solar and terrestrial radiation and the
presence of water in the soil similarly tends to maintain an equable soil temperature.

It is instructive to note that non-indigenous mesophilous and hygrophilous species of
bamboo which have been introduced and grown successfully in comparatively moist and
protected localities in Dehra Dun have been destroyed or greatly injured by frost in exposed
grasslands such as Ramgarh Tappar.

Such a character as the possession of a thick corky bark, also, may often be of special
utility in the way of decreasing danger from fire damage.

18. Even if such adaptations as the above can
be proved by experiment to be the ordinary response of the plant to the stimulus of a
variation in the water supply, this does not alter the fact that, in many cases at least, the
vital importance of such adaptations lies in this, viz. that they enable the plant which has
learnt to respond to this particular stimulus in this particular way to resist extreme
temperatures and thus to survive in the struggle for existence, while other plants which
responded in a different way, no less efficient as regards securing a sufficient water supply,
but failing to afford the necessary protection from temperature, perished. Seeing there-
fore that the main object of cecology is to explain “why each species has its own special
habit and habitat,” it is obvious that we should avoid as far as possible laying undue stress
on a single factor, however important in itself, but which in a multitude of cases is probably
not the dominant factor responsible for the existence of forms of vegetation, as we now
find them in their natural habitats.

19. As indicating how an insistence on the
primary importance of the water supply may quite possibly obscure the vital factor of
dominant importance, we may take the case of those evergreen trees which exist in regions
of moderate rainfall, such as Laurus nobilis, Ilex aquifolium and Prunus Laurocerasus in
the Mediterranean region and species of Persea, Laurus, Phebe, Ilex and Myrica in the
Canary Isles, which are characterised by a more or less xerophilous type of foliage and

f 18 j

HOLE: SOME INDIAN GRASSES AND THEIR GiCOLOGY. 19

which are believed to owe their cecological peculiarities to a deficiency in the summer rain-
fall and to the fact that they must remain active through the winter to utilise the rain
falling at that season. Such plants are consequently supposed to be peculiar to regions
where rain falls chiefly in winter. Species of Ilex, Persea, Phabe and Myrica however
all occur in India in regions where the rain chiefly falls in summer, the season when the
majority of the associated species show their maximum activity. The Sal tree also
curiously enough is practically evergreen and its foliage is unusually xerophilous for a
mesophilous species. It will also be noted that all the trees mentioned have a more or less
pronounced capacity for standing shade. No Forester doubts the extreme importance of
light as a factor capable of influencing the distribution of plants and, in the case of the Sal,
its capacity for withstanding a considerable degree of shade is almost certainly the factor
of most importance in enabling it to gain a footing in forests of miscellaneous species and
to form gregarious forests of its own. Diminution in intensity of light, however, must to
some extent mean less assimilation and therefore less vigour for growth and vital functions,
and from this point of view it can be understood why a plant, which has become adapted to
the utilisation of diffuse light and which has to compete in the struggle for existence with
plants which can utilise light of greater intensity, should find it advantageous to maintain
its foliage throughout the year. At the same time the fact that the foliage has to persist
through unfavourable seasons and withstand the cold of winter and heat of summer explains
why a somewhat xerophilous structure should characterise the foliage. The dominant
factor therefore which here seems responsible for the existence of a remarkable type of
vegetation appears to be not water but light.

In this connection also it should be noted that there are several plant characters which
may be considered adaptations to deal with light or water. The highly polished surface of
many xerophilous leaves for instance may be of service in reflecting some of the incident
light, while, on the other hand, it may be of use in dealing with excessive moisture, in
causing the water to run quickly off the surface. A covering of pilose hairs is similarly
doubtful :—

“The epidermal cells project as countless papillae which give to the leaf a velvety
appearance. Between the papille the water rapidly spreads by capillarity
as a thin film extending over the whole leaf blade, so that the water can eva-
porate more quickly than if it were not thus spread out. The opinion has
also been expressed most recently that these papille serve to increase the
amount of light supplied to the leaf.”*

20. Again, whatever may be the degree of
adaptation exhibited by a formation to the conditions of moisture now prevailing in its habi-

tat, there is no doubt that in a great number of cases the dominant factor responsible for
the reduction of forest to bush and of bush to virtual desert has been man, by reckless

1 Warming, p. 340.

[. 1a J

Doubtful
Adaptations.

Action of
man often
primarily
responsible
for the
creation and
distribution
of communi-
ties.

Importance
of specific
differences.

20 INDIAN FOREST MEMOIRS.

fellings on hillsides and by subjecting woodlands in the plains to excessive grazing and
injury by fires. Similarly savannahs and steppes have in many cases undoubtedly resulted
directly from forest by fellings followed by fires and in some cases frost and excessive
insolation.

It is of importance to realise this for it emphasizes the possibility of reconverting
desert, or grassland, to forest by the selfsame agency, viz. man, and shows that deserts and
grasslands need not necessarily be regarded as permanent on account of wide differences
in climate or soil. While we may therefore utilise such words as xerophilous and hygro-
philous as useful terms conveying some idea of the characteristics of a form of vegetation,
of some of the conditions of its environment and of the way in which the economy of the
plant is adapted to these conditions, we must continually guard against the idea that the
available water supply has necessarily been the dominant factor responsible for the exist-
ence and distribution of that type. In studying the cecology of any plant or plant-com-
munity, we must continually remember that its existence and distribution depend on a num-
ber of factors to each one of which the plant must be just as perfectly adapted as it is to that
of water supply and that a slight change in any one of such factors may suffice to render
the continued existence of such a type impossible. If we wish to understand the cecology
of any plant, or plant-community, we must study, not only the adaptations of its vegetative
members, root, stem and leaf to the water supply, temperature, light, air currents, other
living plants and the animals of its environment, but also, no less, the adaptations shown
by the flowers and fruit and the arrangements necessary for the production of vigorous
seed and its suitable distribution, such as the season which various plants find most advan-
tageous for the production of flowers and fruit and the reason for this.

21. A point that appears to be frequently
overlooked in cecological work is that, although the nutrition of ordinary terrestrial plants
is carried out broadly on the same main lines, differences in detail undoubtedly exist.
There is reason to believe, for instance, that owing to a difference in the osmotic strength
of the substances produced in the cell-sap of the root-hairs some species may be able to
obtain sufficient moisture from concentrated solutions of salts, or from very retentive soil,
whereas others cannot do so easily, and the latter must therefore resort to devices which
aim at decreasing transpiration, the storage of water, and so on. If this is the case we
should naturally expect to find plants with, and without, obvious xerophilous adaptations
in the same habitat. The fact that Butea frondosa which is believed to possess no very re-
markable xerophytic adaptations can exist on saline soil and that it is in full vegetative
activity during the hottest dryest period of the year on the retentive Black Cotton Soil in
the plains of India possibly indicates some such peculiarity. The needs of different species
in respect of the quantity and quality of various mineral salts, of the quantity of oxygen
for respiration, of the intensity of light for assimilation and of temperature undoubtedly
vary considerably and we should therefore naturally expect to find considerable differences
in the nature and magnitude of the adaptations exhibited by different species in response

[ 20 ]

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 21

‘to variations in the same factor. Very little is known about this and hence of the extent
to which structural differences can be relied on to give a correct conception of the intensity
of any particular factor of the environment, but we must obviously be prepared to find
plants with widely different adaptations existing together in the same habitat and forming
a part of the same community.

22. Connected with the above question also
is the possibility of precisely the same object, viz. successful existence, being attained in
different ways by different adaptations. Some cecologists, for instance, do not admit that
two plants, the one with an obvious xerophilous structure and the other with a hydrophilous
structure, can arise through evolution in nature in the same habitat. Thus Clements ex-
pressly states that “it is impossible for the same habitat to produce both hydrophytes and
xerophytes,”* and he explains the well-known fact that many swamp species exhibit obvious
xerophilous adaptations by considering that the latter are stable structures which “ were
developed when these species were growing in xerophytic situations, and not by the hydro-
phytic habitat in which the plants are found at present,”* in other words he apparently
considers that such plants are not in complete harmony with their existing environment.
From what is known of the conditions of plant life and the universal perfection of the
adaptations developed by plants in response to their environment we should naturally ex-
pect that any species out of harmony with its surroundings would, if plastic, rapidly adapt
itself to those conditions, or that, if not plastic, it would fail to survive and would there-
fore not be found existing in nature under such conditions. If, on the other hand, a species
proves stable under different conditions of environment we should naturally be inclined to
think that no new adaptations are required, in other words that the existing structure is in

‘complete harmony with the conditions of the habitat. In the case of swamp plants for
instance the great difficulty in the majority of cases appears to consist in securing an ade-
quate supply of oxygen for the submerged roots. If the oxygen supply is short the normal
functions of the roots will be depressed and a diminished water supply will result. This
can be met either by increasing the oxygen supply and thus the rate of absorption at the
roots, by the provision of more adequate aerating arrangements (e.g. wide air spaces,
pneumatophores, etc.) or by diminishing the loss of water by transpiration so that it may
keep pace with the slow absorption at the roots and thus there is no reason to believe that
plants, exhibiting xerophilous structures designed to diminish transpiration, are at all less
perfectly adapted to a swampy habitat, than are plants possessing hydrophilous arrange-
ments designed to increase the oxygen supply. Indeed of the two types of structure the
former is probably the most perfect and on the whole the most successful, inasmuch as it
may enable a species like Saccharum spontaneum to exist in a very dry and sandy habitat,
where there is a great scarcity of available moisture, in a heavy clay soil where there is at
-some seasons a large quantity of water and little oxygen for the roots and at other seasons

———

1 1c.) DP. 169.
2 L.c., p. 170.

[ #1 ]

The problem
of Successful
Existence
under a given
set of Con-
ditions may
be solved in
nore than
one way.

Convenient
to classify
Terrestrial
plants as

Xerophytes,

Mesophytes

and Hygro-

phytes.

22, INDIAN FOREST MEMOIRS.

a scarcity of water owing to the slow movement of water by capillarity, and finally in a
swamp where there is a maximum of water but very little oxygen, and all this with very
slight changes in structure, the leaves of the swamp form still being clearly xerophilous.

If a deficiency of oxygen in the soil tends to depress root activity and thus diminish the
quantity of water which roots can obtain from the soil, we should expect a heavy non-
aerated soil like clay, or the water-logged soil of a swamp, to be physiologically dry, just as
a sandy soil, through the mere absence of water, may be physically dry, and consequently
we should expect to find the same species not infrequently on dry sandy soi! and on heavy
clay, or ina swamp. This is the case and in India Bombax malabaricum, Odina W odier,
Garuga pinnata and others are often found on physically dry, stony or sandy soils and
also in swampy localities, while Phyllanthus Emblica, Avgle Marmelos and others are often
found on dry, sandy, stony, soils, as well as on heavy clay.

It must also be remembered that a clay soil may be physically dry, as well as physiolo-
gically dry, owing to the fact that the dense nature of the soil prevents the free movements
of water by capillarity and the soil may thus lose water by evaporation at the surface far
more rapidly than it can be replaced from the subsoil water below.

It has also been pointed out above that xerophilous structures frequently serve equally
well as adaptations with reference to temperature conditions, as to moisture conditions,
and it is therefore clear that we must be prepared to find one and the same species pro-
vided with a certain set of adaptations existing in a number of different habitats and
entering into the composition of a number of different communities. It may grow equally
well in different communities; it may be dominant in one and form a small proportion of
others; it may attain its highest development in one and show inferior dimensions in
another; it may reproduce itself readily in one and with difficulty in another, and so on.

23. With particular reference to the vegeta-
tion of India it will probably be found convenient, at all events for the present, to confine
the word Hydrophyte to a true aquatic plant, as defined by Warming (paragraph 7 above)

and then to subdivide terrestrial plants (including marsh plants) into Xerophytes, Meso-
phytes and Hygrophytes.

It is necessary however to consider what is really the essential meaning of these terms.
(Ecologists frequently overlook a fact of great importance, viz. that a xerophilous leaf-
structure, as a rule, merely indicates that transpiration is slow, that when assimilation is
proceeding the actual quantity of water vapour passed off from the leaf in a unit of time
is relatively small, but it by no means follows that the total quantity of water which a
plant with xerophilous leaves may be able to utilise during a period of say one complete-
year is also insignificant. Owing to the fact that a plant with xerophilous leaves is.
frequently able to remain vegetatively active during periods when another plant with
mesophilous leaves is obliged to cease activity, the actual quantity of water utilised in a.
year, and the vigour of growth, as indicated by the dimensions attained, may be actually
greater in the case of the former than in the case of the latter. It is known that, when the:

[ 2 ]

HOLE : SOME INDIAN GRASSES AND THEIR G:COLOGY. 23

quantity of moisture in the soil falls below a certain limit, the movement of water by capil-
larity in the soil becomes very slow. It is therefore conceivable that under these conditions
the roots of a mesophilous plant, in order to keep pace with the rapid transpiration from
the leaves, will remove water more rapidly from the soil than it can be replaced by capil-
larity. Vegetative activity would therefore obviously become impossible. A xerophilous
species on the other hand with its slower transpiration and absorption might be able to
continue activity.

It is believed that the savannah grasses Saccharum Munja and S. Narenga, dealt with
in this paper, supply an illustration of the point here considered. The glaucous, wax-coated
leaves of the former are obviously xerophilous, while the dark-green leaves of the
latter are comparatively mesophilous. Notwithstanding this fact the former has frequently
been seen in the locality dealt with in this paper exhibiting more vigorous growth and
attaining larger dimensions than the latter. Plants of both species experimentally culti-
vated on loam, however, showed that, in February when the moisture-content of the surface
soil was comparatively small, although both species were growing under similar conditions
of soil and moisture, while S. Munja was growing vigorously (as judged by actual measure-
ments of the internodes of the culm), S. Narenga was vegetatively inactive.

Warming rightly notes that “ Tropical rain-forest constitutes the climax in the develop-
ment of vegetation for the whole world.”* We may therefore accept the evergreen tropical
rain-forest, which Schimper classifies as a hygrophilous formation, as the most highly
‘developed cecological type of vegetation, the xerophilous as the least highly developed
and the mesophilous as an intermediate type.

In view of the remarks made above, however, it is obvious that we must not depend on so-eaned
the so-called xerophilous and hygrophilous characteristics of plants as an unfailing criterion SEOs
of the type of vegetation to which they belong, but must consider them also in connection P10"

characteris-
with periodicity and therefore regard them rather as an indication of the manner in which ‘i; Neven

unfailing

particular plants deal with the supply of water they are able to obtain, hygrophilous charac- Cees

teristics usually enabling a plant to deal with a large quantity of water in a short time and ree ere)
xerophilous adaptations enabling a plant to utilise its water supply slowly in small io
quantities. As the true criterion of a xerophilous or hygrophilous type of vegetation, emetanon:
however, it seems certain that we must consider the total quantity of moisture which plants

-are able to utilise throughout their existence and this would be approximately indicated

by the vigour of growth and dimensions attained. On the whole it seems probable that the

best factor to accept would be the actual dry weight of organic material produced per unit

of area at the period when the majority of the individual plants composing the vegetation

have attained maturity. Thus for grassland the weight of dry grass, and for woodland

the weight of wood, per acre, would probably be an approximate criterion of most practical

utility, although strictly speaking the weight of subterranean as well as of aerial parts

ought to be considered and also the weight of the ieaves and fruit produced by woody

17.c., p. 340.
Leo

Classification
of Woodland,
Bamboos.
Palms,

Conifers.

24. INDIAN FOREST MEMOIRS.

plants. In the case of woodland we must obviously consider the wood produced by all the:
species which enter into the constitution of each type and by all the plants existing in
both overwood and underwood at the age of maturity. From this point of view it will be
seen that, other things being equal in each case, the most hygrophilous type of vegetation will
be composed of individuals attaining the greatest dimensions, while vice versé the most
xerophilous type will contain the individuals which show the smallest dimensions; that the
most hygrophilous type will include the most densely stocked and the xerophilous type the
most open and poorly stocked communities; that the most hygrophilous type will include
the plants with the longest period of vegetative activity and the most xerophilous type the
plants with the shortest period of vegetative activity; and that the most hygrophilous type:
will include the individuals with the most marked hygrophilous adaptations and the most
xerophilous type the individuals with the most marked xerophilous characteristics.*

At present, owing to the absence of statistics regarding the weight of organic material
produced per unit of area by the different types of vegetation, it is obviously impossible to-
do more than indicate an approximate classification of communities as hygrophilous,
mesophilous and xerophilous, paying particular attention to such characters as the
dimensions attained by the constituent plants, duration of the period of vegetative
activity, and the density of stocking. Such a classification therefore can only be accepted
as tentative and provisional and will be liable to correction as statistics are gradually
accumulated.

24. Bamboos constitute a peculiar type of
vegetation more or less intermediate between grassland and woodland but which is most
conveniently dealt with as woodland and of which it is believed that xerophilous, mesophi-
lous and hygrophilous types exist.

Woodland composed of Palms will also perhaps have to be dealt with as a separate
type of woodland.

Warming places all coniferous woodland into a separate class which he considers
xerophilous stating that “the soil upon which coniferous forest occurs varies widely, yet,
so far as reliable information is available, it is always physically or physiologically dry.”
Schimper however rightly points out that “it is by no means admissible to include them
(i.e. coniferous trees) among xerophytes, as Warming has done. The term xerophyte may
to a certain extent apply to several species of Pinus and of Juniperus of dry, sandy, and
stony soils; * * but it does not apply to most species of Abies and Picea, our silver-fir
and spruce for instance, which require as much moisture as broad-leaved trees.”®

Coniferous woodland should be dealt with as a separate type, of which xerophilous,.
mesophilous and hygrophilous forms probably exist.

1 In connection with what has been said above it is interesting to note that, so far as dimensions of plants are concerned, the highest
development is attained by evergreen and not by deciduous trees, e.g. the giant trees of the world belonging to the genera Sequoia and.
Eucalyptus.

2iiicepyolO:

8 Lc. p. 564

HOLE: SOME INDIAN GRASSES AND THEIR (ECOLOGY. 25

Similarly in the case of Dicotylous Woodland deciduous and evergreen woodland yerenen
c . . and Veciaduous
must each be dealt with separately. Of the latter xerophilous, mesophilous and hygro- Woodland.

philous forms certainly exist.

It is probable that as our knowledge extends we shall have to deal separately in this
way with other types of vegetation consisting of plants with similar needs, as indicated
by close systematic relationship, or of plants which, although not closely related, have
adopted a similar mode of economy, as indicated by similar structural differences, habit of
erowth, periodicity, and so on.

25. It is believed that the attempt to utilise Ciassification
the presence or absence of trees as a character to separate different types of grassland, eee
such as steppe from savannah, must result in a purely artificial classification. There are
very few regions of the earth where grasses grow where some kind of woody plants cannot
exist and it is believed that the presence or absence of trees or shrubs in grassland is
usually, as it were, an accident, due to a peculiar combination of local conditions which
may, or may not, exist in what is obviously one and the same type of grassland. There is
no doubt that steppe can be produced directly from savannah and even from forest by
artificial means and on the other hand it is certain that trees may and often do occur in
typical meadow.

Areas of precisely the same type of grassland, e.g. composed chiefly of Saccharum
Munja, or of Saccharum Narenga, may be found sometimes with, and sometimes without,
scattered trees, according as the local conditions of frost, facilities for seed distribution
and so on are, or are not, favourable for the establishment of the parallel type of woodland.

Warming considers the meadow as a type of his mesophilous grassland while reed-
swamp and savannah are respectively dealt with as hydrophilous and xerophilous forms
and are placed by him accordingly in inferior classes. When we consider the gigantic
dimensions of the constituent plants and the dense stocking of areas carrying reed-swamp
and wet-savannah, it is impossible not to regard. these as far more highly developed cecolo-
gical types than the typical meadow. It must also be noted that the peculiar close carpet
formed by the vegetation and the moderate dimensions attained by the constituent species
(usually not over 3 feet) in what is usually termed meadow is, sometimes at least, an arti-
ficial character resulting from grass-cutting, firing or grazing. Any of these agencies can
seriously retard and influence the development of grasses. They frequently for instance
do not permit the culms to attain their full dimensions and less height growth consequently
results; the ground in the immediate neighbourhood of the plants is thus less shaded and
more culms are therefore able to develop and occupy the spaces between the individual
plants, a close sward being produced in consequence. On the other hand, however, meadow
does undoubtedly often owe its characteristics to the peculiarities of the constituent species
and in India it may arise as the result of a regressive succession, such as occurs for instance
when the firing of moist Saccharum Narenga savannah so dries the loamy soil as to make

[ 2 ]

Suggested
Scheme of
Ccological
Classification

for India.

26 INDIAN FOREST MEMOIRS.

the vigorous development of this species impossible and its place is taken by smaller species
such as Andropogon intermedius, Pollinia sp. and others.

For the present it seems advisable to disregard the presence or absence of scattered
trees as an important character and to classify grasslands, provisionally, on the basis of the
density of stocking combined with a consideration of the dimensions attained by the consti-
tuent plants. Disregarding for the present reed-swamp, suvannah would cover the more
strongly mesophilous and all the hygrophilous grasslands, meadow might be retained for
the less strongly mesophilous grasslands and steppe for all xerophilous grasslands.

So far as domestic animals are concerned, grassland consisting of xerophilous, or the less
marked mesophilous, species, as a rule produces the best fodder, hygrophilous fodder being
what is usually described as too coarse or rank to be palatable except to such animals as
buffaloes. At the same time the less vigorous and the younger the culms and foliage of
coarse species, the more palatable as a rule the fodder and hence, firing, which decreases
the vigour of the plants and lays bare to the attack of animals the young shoots and leaves
as they appear, by depriving these of the protection afforded by the dead leaves and culms,
is an expedient which has been resorted to by man from time immemorial to improve the

fodder yield of coarse mesophilous and hygrophilous species.
26. Based on the above ideas, the following

is suggested as a purely tentative and provisional scheme of cecological classification adapted
to the vegetation of India which will give some idea of the limits between which the
different types of vegetation vary and of the class of vegetation which may be taken as
fairly typical of the main groups. As noted above, it is obviously impossible at present,
in the absence of definite data regarding the weight of organic material produced per unit
of area, to draw a definite line between the various groups, or to define the latter correctly.
It is believed, however, that the scheme will facilitate the work of compiling and arranging
the descriptions of the various existing plant-communities, without causing confusion by
attempting to introduce too much detail; that it will in fact facilitate the collection of
further information and data which are essential before a sound ciassification can be

evolved.

Type of Vegetation. Types of Communities.

I.—Aquatic or Hydrophilous : Communities of Aquatic Plants.
II.—Terrestrial—

A. Xerophilous . 0 .| Grassland (=Steppe).—This is either ephemeral with meso-
philous structure and adaptations, or more persistent with
xerophilous adaptations. Examples of the former are the
poor grasslands constituted chiefly of ephemeral species of
Eragrostis common on soil-less places, or in deserts. This
type gradually passes through grasslands of such species
as Andropogon monticola and A. contortus into meadow.

[ 26 ]

HOLE: SOME INDIAN GRASSES AND THEIR GiCOLOGY. 27

Type ot Vegetation.

A. Xerophilous—conid.

B. Mesophilous

Types of Communities.

Open grasslands composed of such species as Aristida cyan-
antha and T'riraphis also probably come here, although
when densely stocked such types must be perhaps consi-
dered mesophilous.

Woodland.—This is also characterised by a short period of
vegetative activity, or by a longer period with more or
less marked xerophilous adaptations,

The dry open deciduous woodlands formed of such species as
Cocklospermum Gossypium, Sterculia urens, Boswellia
serrata, Euphorbia sp. and others come here, also many
woodlands of Acacia Catechu, the scrub or bush-growth
composed largely of such species as Nyctanthes, Zizyphus
and Acacia and the desert communities of Tamarix,
Prosopis spicigera, Capparis, Salvadora, Ephedra, Orthan-
thera and others.

Tt will be noticed that oniy the poorest type of evergreen
woodland comes here, consisting of such species as Salva-
dora oleoides and Tamarix while Zizyphus Jujuba is also
practically evergreen.

Shrubby growth of temperate species such as Arundinaria,
Rhododendron lepidotum and R. Anthopogon also comes
here, while among Conifers some Juniper forests, or
bushland, and possibly also some forests of Pinus, e.g. of
Pinus Gerardiana, should probably be included in this
eroup.

Grassland (=Meadow or Moist Savannah).—This group in-
cludes the Meadows of the Himalayas and of the so-called
Hill-Forests of India generally. Of these Duthie writes:
“The vast stretches of undulating meadows, known as
‘“maidans,’ and which extend from the upper limits of
the forests to the snow line, are composed of many of the
most nutritious grasses of the world, some of them belong-
ing to species well known in Europe for their good grazing
qualities.” Here also come the best fodder-yielding
erasslands of the deciduous forest areas, while the highest
development is probably attained in the Moist Savannah
formed of such species as Saccharum Munja and Saccha-
rum Narenga.

Woodland.—The majority of the so-called “‘ Deciduous-’’ and
“* Hill-’”’ Forests of India fall into this group, including
most of the Coniferous Forests and most of the Bamboo
areas composed of such species as Dendrocalamus strictus.

1 Fodder Grasses of Northern India by J. F. Duthie, 1888, p.ii.

[ 27 ]

28 INDIAN FOREST MEMOIRS.

Type of Vegetation. Types of Communities,

C. Hygrophilous : .| Grassland (= Wet Savannah and Reed Swamp).—The grass-
lands densely stocked with gigantic species such as L71-
anthus elephantinus and Saccharum arundinaceum must
be placed here and may be termed Wet Savannah. Gyrass-
lands of Erianthus Ravenne must be regarded either as
one of the most highly developed forms of mesophilous
savannah or as ene of the less strongly developed forms of
hygrophilous wet savannah.

Probably also this group would include the more vigorous
Reed-Swamps composed of gigantic species of Phragmites
and Arundo.

Woodland which is typically developed in the densely stocked
Tropical Evergreen Forest. The woodlands composed
mainly of gigantic Bamboos found in the evergreen zones
of Burma, Assam and Bengal probably come here.

The communities which have been roughly indicated above are merely intended to
serve as fairly intelligible types around which those communities which resemble them most
closely can be conveniently grouped, but it 1s at present impossible to describe these com-
munities in full detail, or to accurately define the limits of the groups under which they are
classified, for which far more detailed and accurate information is required than is at
present available.

27. The scheme of classification which has
been sketched above, very imperfect though it is, at all events indicates that, so far as India
is concerned, for each main type of grassland there exists throughout a more or less parallel
and corresponding type of woodland, both of these types existing under approximately the
same conditions of available moisture, as determined by the combined effects of climate,
soil, and temperature.

In the case of both grassland and woodland it will be noticed that the scheme com-
mences with the simplest or most xerophilous form and culminates in the most highly

valueof developed type, i.e. with the evergreen tropical forest for woodland, and with the wet-

colog. ay
Forest 6Savannah and reed-swamp for grassland.

Officers. 28. The above somewhat detailed sketch of
the scope and objects of the study known as (Ecology, together with a tentative skeleton
scheme of classification capable of adoption for the compilation and arrangement of the
cecological descriptions of the various plant-communities constituting the vegetation of
India has been given here, partly because this is required to justify the classification and
description of the types of vegetation dealt with in this paper, and partly because, for
Foresters at all events, cecology is probably the department of botany of most interest and
of greatest practical importance. In cecological study a Forester can utilise all the

r 2 1

HOLE: SOME INDIAN GRASSES AND THEIR CECOLOGY. 29

knowledge he possesses of Morphology, Anatomy, Systematic Botany, Physiology, and
Pathology to the greatest advantage in the solution of problems of practical importance
and the successful solution of which moreover constitutes an important part of his daily
duties.

Trained Foresters will find no difficulty in distinguishing and describing the various
communities of woodland, a task indeed which forms a part of the duties of every Working
Plans Officer, but this work requires to be undertaken systematically and it is obviously
advisable to commence the work with special reference to particular species of greatest
importance in all the localities where it occurs. Far more attention also must be paid to
the accurate determination and recording of the condition of the factors of the locality
which constitute the plant’s environment in various habitats. The more detailed and
-accurate this determination, and the greater the number of the localities in which the observ-
ations can be made, the more certainly will the conditions be revealed which are essential
for the healthy existence and development of a particular species, and the clearer will be
the indication as to the method of treatment which must be adopted if we wish to favour
its development and reproduction and to cope with its diseases. It must also be clearly
‘recognised that, in so far as such recorded observations of the condition of environmental
factors are correct, they are facts, and as such provide just as satisfactory a basis for a
logical conclusion as do experiments. In careful experiments we endeavour to simplify
the environment of a plant and in order to test the effect on it of a single factor the latter
alone is, as far as possible, allowed to vary and the effect of such variation
on the plant is watched. The practical drawback to experiments, however, lies partly in
the difficulty experienced in artificially arranging for the isolation of the various factors
and partly in the long period of time which is required to obtain accurate information
regarding the effect of all the factors of importance capable of effecting the development of
a particular plant. Information regarding a single factor alone is of very little value and
goes a very little way in explaining the facts observed in the field, inasmuch as in nature
several factors are in action together and while one factor may be of primary importance in
producing a certain result in one case, in another case its action is masked by the effect of a
different factor which, in its turn, has become of primary importance and responsible for
the result obtained.

In recording careful observations regarding the conditions to which a particular plant
or type of vegetation is exposed in nature, however, we are doing no more nor less than
attempting to read the result of experiments so-to-speak which have been arranged and
carried out for us—certain factors have been and are stil! in cperation—the result of this is
a certain type of vegetation—-and the problem for solution is to answer the query what
particular factor or combination of factors from among those in operation is responsible for
the result produced ?

Moreover, it must be remembered that, whereas experiments with woody plants usually
require long periods for the production of valuable results, the work of recording the details

eee dh

30 INDIAN FOREST MEMOIRS.

of ‘readymade’ experiments can be comparatively quickly done, and by comparing a
number of such cecological records it is frequently possible to find several cases in which all
important factors, with one exception, have varied very slightly or remained practically
constant and thus to directly obtain information regarding the effect of the factor which
varied considerably. Even when such comparisons do not at once enable a satisfactory
conclusion to be reached they indicate the lines on which an experiment should be organised
in order to give the necessary result. An idea frequently prevails that no really valuable
information can be obtained regarding such a factor as the quality of the soil, except by
detailed and skilled work such as by the chemical analysis of the soil. In the present
state of our knowledge of the nutrition of our principal species, however, a chemical analy-
sis alone can teach us practically nothing regarding its suitability for a particular species,
whereas careful comparative observations regarding the depth of soil, admixture of gravel,
sand, clay and humus, moisture content, and depth of water table can be made by anyone
and are as a rule, except in rare cases such as where an excess of salts is present, of primary
importance in deciding the suitability or otherwise of a soil for a particular species, inas-
much as these factors regulate the quantity of water and soluble salts and also of the
oxygen available for the roots.

More or less complete data of rainfall and temperature are usually available and the
other most important factors constituting the environment of our forest plants are probably
winds, light, associated plants, grazing and fire damage regarding all of which comparative
chservations of value are possible.

It should also be noted that it is by no means always necessary to extend observations
over a wide area before satisfactory conclusions can be obtained. Experience has shown
that if observations are made in different carefully selected habitats good results can be
obtained in a limited area, e.g. of a single district or division, where the total range of
intensity of the various environmental factors, within which the existence of a particular
plant is possible, can often be found to be operative, and it must be remembered that local
differences in soil and situation are capable of exerting an effect on the water content of
the soil equivalent to that caused by the most arid or humid climate.

While Forest Officers in India have few facilities or opportunities for carrying out and
recording the results of prolonged experiments, the work of carefully observing and record-
ing descriptions of the types of vegetation and of the environmental factors which exist in
the forests constantly traversed by them on tour is comparatively easy.

Forest Officers in India have a unique opportunity of adding to cecological knowledge
which besides being of great scientific value will enormously increase the efficiency of forest
management; they have in their charge every type of both grassland and woodland, ranging
from the lowest type of alpine or desert grassland to the luxuriance of the evergreen tropi-
cal forest.

It is hoped that Forest Officers, realising the possibilities of such work, will do their

utmost to systematically publish accurate descriptions and photographs of the chief types
[ 30 ]

HOLE : SOME INDIAN GRASSES AND THEIR GiCOLOGY. aL

of vegetation found in their forests and also so far as possible ascertain and record the
condition of the environmental factors existing in the various habitats. As regards the
cecology of the principal species, also, it is particularly desirable to record careful notes on
such points as the periods of vegetative activity, of flowering and fruiting, the agencies by
which flower pollination and seed distribution are effected, the time when the seed naturally
germinates and the factors which appear of special importance in effecting the development
of the young plants during the first few years of existence, as on all these points the main-
tenance or extension of the area occupied by a particular species, or type of vegetation,
largely depends.

29. It is a remarkable fact that, although the
value of the study of mycology and entomology in helping us to secure the healthy develop-
ment of our valuable plants is fully recognised, the dominating importance of physiology
and cecology is frequently lost sight of and the view is not infrequently held that most plant-
diseases of importance are caused primarily by either fungi or insects.

Every plant-physiologist however is aware of the vital importance to plant-life of such
primary factors as the available moisture supply, the degree of soil aeration, the tempera-
ture, light and competition of other plants and knows that even slight variations in these
factors are able to cause disease and death. Notwithstanding this, we have, as yet, no
precise knowledge (in the case of any single Indian tree of importance) as to the degree of
intensity at which such factors first begin to cause disease and when they must accordingly
be considered injurious.

It is important to note, also, that an accurate knowledge of such factors offers every
prospect of producing practical results of value, seeing that the intensity of these factors
can be altered within considerable limits by the ordinary operations of practical forestry
and thus those conditions may be produced which are necessary for the healthy development
of particular plants in cases where such conditions do not exist, or have been destroyed by
careless management. Thus the water content of a sandy soil may be increased and its
aeration diminished by introducing a species with considerable power of humus production
such as Adhatoda Vasica, the aeration and texture of a heavy soil may be improved by
introducing a species capable of withstanding poor soil-aeration and of producing a good
humus supply, temperature can be altered by a judicious use of shelter-woods and by
arranging fellings so as to secure gocd circulation of air, while plant competition may be
controlled by introducing factors such as periodic fires which are unfavourable to the com-
peting plants. On the other hand it is a remarkable fact that, although mycology and
entomology are able in many cases to suggest effective remedial and preventive measures
which are applicable in agriculture, horticulture and arboriculture, such measures can
rarely be adopted in practical forestry, especially in countries where the general value of
the produce is small and the forest management not intensive. Frequently the only remedy
is to cut out the affected trees and in bad cases to change the species, which is often a
lengthy and difficult operation, while the new species is frequently less valuable than the

[3r]

Importance
of Ecology
in Study of
Plant-
Diseases.

32 _ INDIAN FOREST MEMOIRS.

one it has replaced. Yet even in agriculture the impossibility of adopting effective direct
remedial measures is leading more and more to the adoption of such indirect methods as
the production of new and resistant varieties. Measures such as these, however, are
obviously out of place in practical forestry; the forester, as a rule, must take his species as
he finds them in nature and if he wants to favour their healthy growth and development he
must keep as closely as possible to those conditions under which they grow best in nature
and under the influence of which the species have been gradually evolved.

Moreover to carefully study the requirements and patiently elaborate the economic
treatment of our indigenous species, which we know to be suited to the local conditions of
environment, is surely, as a rule, a more rational policy for practical forestry than to try
haphazard experiments with exotics on which money has frequently been lavished in the:
past with little practical result. Such experiments indeed as a general rule merely
emphasize the fact that exotics are usually more liable to diseases than are the local species.
It is a well-known fact that the most injurious insect and fungal-attacks tend to occur
where unnatural conditions of culture have been adopted, such as massing in pure woods
species naturally occurring chiefly in mixed woods, and the adoption of mixed crops (i.e. a
return to normal conditions of existence) is one of the most practical preventive measures
recommended alike by mycologists and entomologists. There is moreover good reason to
believe that the expert mycologist, or entomologist, with his ideas concentrated on his special
branch of study, frequently fails to realise the dominant importance of the primary factors.
of plant physiology. He may thus be led to believe that a particular fungus, or insect, is
the primary cause of a diseased state for which in reality the unfavourable condition of
some primary factor, such as available soil-moisture, is really responsible, the fungus or
insect being merely a secondary factor and the natural consequence of a sickly condition.
In such a case the real remedy should consist in attacking the dominant factor and not the
fungus or insect.

In the treatment and prevention of diseases generally the great importance of main-
taining the general health and resistant-power of the organism is now usually admitted.
and this can best be done by studying the effects of those primary factors which we know
to be capable of directly causing disease and health. This study is capable of giving us
the knowledge necessary to manage our forests in such a way that our valuable species will
be exposed to the conditions most favourable for their development in nature. Considera-
tions such as the above are gradually forcing practical Forest Officers to the conclusion that
physiology and cecology at present constitute the richest field for research work which aims
at improving the economic management of our forests.

When the post of Forest Botanist was created at the Dehra Dun Forest Research
Institute the principal duty prescribed for the incumbent of that post was that of studying
the diseases of our important forest species. With the object of attaining the most valu-
able results in this direction, for the reasons briefly outlined above, the Botanist undertook
the systematic study of the physiology and cecology of forest plants and has commenced.

fe 32) i)

HOLE: SOME INDIAN GRASSES AND THEIR (COLOGY. 30

work with a single species, viz. the Sal tree. It is hoped that, so far as possible, other
Forest Officers will do what they can, on the lines which have been briefly indicated in the
last paragraph, to start similar work for other important species.

30. The principal object of the study of eae
Indian Forest Grasses with which the present paper is immediately concerned is to acquire, nore
as soon as possible, information which will indicate the best treatment to apply to forest
grasslands for the purpose of (@) improving the yield of fodder and (b) accelerating their
afforestation.

It was recognised that the information required might be obtained by carrying out Uulity of
careful experiments. veriments,

Forest experiments, however, as noted in paragraph 28 above, usually require
long periods for their completion and they cannot therefore supply information of imme-
diate practical utility. In the second place anyone who has attempted to practically carry
cut a forest experiment knows that it is exceedingly difficult to obtain a result which can
with certainty be ascribed to the influence of a single factor, owing to the co-existence of
other factors capable of affecting the resuits; in other words that it is very difficult to
obtain accurate information regarding the effect of only a single factor, the way in which
it acts and the conditions capable of accentuating or diminishing its action.

In the third place a plant in nature is subject to the influence of not one but a number
of factors, each of which, under certain conditions, may be of dominant importance in
rendering the existence of the plant, or at all events its vigorous development, impossible.

It is therefore obvious that a few experiments designed with the object of obtaining
information regarding the effect of a single factor, such as fire damage, when that factor is
of dominant importance, can only give results of very limited practical utility and which
are not found to hold good under other conditions, when some other factor has become of
dominant importance.

It is a frequent experience that the results obtained from carefully conducted experi-
ments in one locality, while apparently accounting for the facts observed in that locality,
obviously fail to explain the facts existing elsewhere.

Such apparently contradictory results are clearly of little practical value and they are
the natural consequence of an insufficient knowledge of all the factors capable of vitally
affecting the welfare of a plant in nature. It is only when a good knowledge of all such
factors has been obtained, of the way in which they act and of the conditions capable of
accentuating or diminishing their action that we can hope to recognise at once which of
these factors is of dominant importance in any particular case and what is the best practical
treatment to adopt under each set of conditions.

It will therefore be seen that, instead of a few casual experiments, an extensive series
of experiments is required in the case of each plant and that such experiments cannot
possibly be successfully organised until a preliminary knowledge has been obtained as to
what factors appear to be of most importance in regulating the natural distribution of the

[ 33 }

Importance
of careful
Field Obser-
vations.

Od INDIAN FOREST MEMOIRS.

plant in nature, the way in which they appear to act and the conditions capable of affecting
their action. The necessity of acquiring a preliminary knowledge of the factors at work
in each case, before attempting to organise experiments, or to interpret experimental
results, will be more clearly understood perhaps from an example.

In the preliminary work which has been carried out with forest grasses it has been
ascertained that while Saccharum Munja may be killed out by repeated early firing on
sand, early firing on loam is practically innocuous. Also that although Andropogon
monticola may be more or less injured by an early fire on sand, repeated early firing on
such soil may result in this species becoming dominant on areas formerly occupied by a more
vigorous species Saccharum Munja, owing to the fact that the latter is liable to suffer more
severely by fires. If such points had not been ascertained by preliminary field work, it
is practically certain that experiments would have given incomplete and apparently contra-
dictory results, whereas when such points have been ascertained from field-work they can
subsequently be checked and confirmed by suitably organised experiments as opportunity
sccurs, when this is considered desirable.

The view which has been emphasized above then consists in this that forest experiments
cannot, as a rule, be properly organised in such a way as to give results of practical value
and of wide application and the results obtained from such experiments cannot as a rule
be correctly interpreted until careful and extended field observations have first indicated the
principal factors which are at work and the way in which they ct.

In addition to the above considerations also, owing to the recent organisation of the
Research Institute, the Botanist had not the staff required for the organisation and conduct

of extensive field experiments.
31. While the capacity of forest experiments

(on the scale on which they are usually conducted) to supply satisfactory evidence regarding
the effect and importance of various factors is frequently over-estimated, the importance
of careful and systematic field observations on the other hand is often not sufficiently
appreciated.

It has been pointed out in paragraph 28 above that satisfactory evidence can often be
obtained by careful and systematic field observations. In a wide series of observations in
which all important factors are constant, or vary only within very narrow limits, with the
only exception of a single important factor which varies considerably, if the growth and
vigour of a particular species is invariably found to vary in direct proportion to the
variations of that single factor, it is logical and permissible to conclude that such a varia-
tion of growth has been caused by the factor in question and such a conclusion is obviously
of great practical importance.

Difficulty is of course experienced in the case of factors which are closely correlated,
such as the intensity of ght and the amount of available moisture, when dealing with
species for which the possible range of intensity of both factors is very limited, seeing that
a given condition of growth may frequently be due as much to one of these factors as to
another. It is however perfectly possible to arrange a series of observations in the forest

Ege

HOLE: SOME INDIAN GRASSES AND THEIR (G:COLOGY. BD

where the water supply is ample on account of percolation or otherwise but where the
light varies, and similarly a series of observations where the light is obviously ample but
where the water supply varies, and thus to assess separately the effect of each factor. An
interesting point arose in this connection with the observations carried out during the
present work regarding the effect of light on the development of Hrianthus Ravenne.

Several cases were noticed in coppice areas where the growth of this species was
obviously inferior and this was at first ascribed to the slight lateral shade existing in the
habitat. Further repeated observations showed that considerably denser shade than this
had no marked injurious effect provided ample water was available and the inferior growth
noticed was thus eventually shown to be mainly due to the drying of the soil by the numer-
ous superficial adventitious roots developed by the coppiced trees. Similarly it is quite
oossible to arrange for a series of observations in nature in localities where the hight
intensity varies greatly but where the temperature varies slightly and vice versa, although
the artificial isolation of these factors in experiments usually offers considerable practical
ditficulties.

While field observations can thus be arranged in such a way as to obtain information
regarding the individual action of each factor of importance, they offer this great advan-
tage over experiments that they enable information to be obtained regarding all the
factors of primary importance far more quickly than would be possible with experiments.

The way in which the distribution of a particular plant depends in nature on a com-
plex of factors all acting together, so that in one place one factor is of dominant importance
and in another spot an entirely different factor, has already been briefly alluded to in
paragraph 28 above and from this it will be clear that whereas cecological observations in
the field can give us information, often very little less accurate than that yielded by experi-
ments regarding the individual action of isolated factors, they give us an incomparably
better idea of—

(1) what factors are of primary importance in the case of any particular species,
(2) what is the approximate degree of intensity of each such factor which in itself
may suffice to cause obviously unhealthy or inferior growth of that species,

and they thus enable us to explain the facts of plant distribution, to suggest suitable
methods of treatment calculated to improve the development of various species, to prevent
their diseases, or to decide on what species can be grown with most success in a particular
locality, far more rapidly and satisfactorily than can the results of a few experiments
dealing with the effect of one or two factors.

32. It was therefore decided that the objects pines of
of the proposed work in connection with forest grasses could be most satisfactorily tdeee ae
attained by— Morest

: z j A : : : Grasslands.
(a) Selecting a few important species of grass which are widely distributed and
which tend to be dominant over large areas.

so]

36 INDIAN FOREST MEMOIRS.

(b) Studying the selected species carefully in the field in their various habitats and
compiling for each species a careful cecological record showing the conditions
under which it is found to grow vigorously and unsatisfactorily, respect-
ively, in nature, special attention being paid to the character of the soil, the
available moisture, light, vigour of associated species, grazing and fire.

(c) Cultivating the selected species during at least one entire season, noting the
normal habit, periodicity, development and growth of the culms in the experi-
mental garden and comparing this as far as possible with what was observed
to be the usual development in the forest. By examining these species care-
fully on areas which have been fired at different seasons and in different
habitats and by there noting the direct damage done by fire and its effect on
the normal development of the plant.

(d) Studying the conditions affecting the growth and development of seedlings of
the tree species with which it is desired to afforest erasslands as far as pos-
sible, 7.e. of the Sal.

It was believed that the above would quickly supply information which would be of
immediate utility—

(1) in indicating the best treatment to be applied to forest grasslands,

(2) in indicating the lines on which a comprehensive scheme of experiments could
be satisfactorily organised and carried through, for the purpose of testing
the correctness of the conclusions which had been based chiefly on field
observations and of supplying additional information which could not be
obtained satisfactorily from field observations.

The work done therefore up to date has been carried out on the above lines.

Owing to the recent organisation of the Research Institute and to the lack of the neces-
sary staff and applances it has been found impossible, up to date, to do more than carry
out a few rough preliminary experiments and control cultures, or to determine instrument-
ally in the field the exact intensity of such factors as available moisture, light, temperature
and the proportion of sand, clay and humus in the soil. The observations which have been
made, however, have it is believed heen sufficiently accurate to give results of practical
value.

The cecology of sal seedlings is a study which has been only just commenced and as
this study is continued it is hoped that with reference to it, at all events, instrumental
observations will in all cases be feasible, perhaps with the help of the students of the
Imperial Forest College, and that a more accurate determination and more precise defini-
tion of the intensity of the various factors will thus be possible in future work.

Sees 33. It was advisable as far as possible to
ar a select not only those species which are widely distributed and markedly gregarious, but

also those which are of economic value, so that their cecological study might also incident-

[ 3h

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 37

ally provide information of economic value on such points as the best treatment to adopt
in areas where a grass like Saccharum Munja is worked for munj fibre, or for paper mate-
rial, the best species to cultivate for the purpose of fixing sand or unstable soil, the kind of
locality in which a valuable fodder grass hike Andropogon monticola can be cultivated
with most prospect of success, methods to adopt with a view of eradicating troublesome
species which have taken possession of valuable agricultural lands, and so on.

Another point borne in mind also is that, so far as possible, when an important species
is being studied, other species of less importance, occurring in the same locality and likely
to be mistaken for it, should also be studied and fully described and illustrated, special
attention being paid to characters which can be easily recognised in the forest and which
serve to distinguish the important from the unimportant species.

For these reasons the following species were selected for study :—

Saccharum spontaneum,
Saccharum Munja,
Saccharum Narenga,
Erianthus Ravenna,
Imperata arundinacea,
Aristida cyanantha,
Triraphis madagascariensis,
Andropogon monticola.

In subsequent papers it is hoped to deal with the Bhabar grass (Ischemum angusti-
folium) and the valuable odoriferous oil-grasses.
34. The information collected regarding srrange-

ment of

each species has been arranged under the following heads :— teeseiintion
collected.

(A) Botanical Description.
(B) Taxonomy.
(C) Biological and CEécological Notes—
(a) Habitat.
(5) Development of Culms, Season of Vegetative Activity and Flowering.
* (c) Susceptibility to Fire Damage.
(zd) Utility as Fodder and best Treatment for Fodder Production.
(e) Relations of Grassland to Woodland.
(D) Economic Uses.

This paper has also been divided into three chapters, the first containing a brief des-
cription of the locality, the second containing the detailed information referring to each
species and the third containing a summary of the work done in the form of suggestions
regarding the practical treatment of grasslands.

. . 35. The identification of grasses from the mustrations.
brief written descriptions given in a work like the Flora of British India is often found

[: 3f J

References.

Acknowledg-
ment of
Assistance
received.

88 INDIAN FOREST MEMOIRS.

difficult and troublesome by the average Forest Officer and it is hoped that the stress which
has been laid on characters easily recognised in the forest and the drawings and photo-
graphs which have been given of each species will facilitate the work of identification. In
most cases a measuring rod, showing lengths of 6 inches, painted alternately black and
white, have been included in the photographs, so that the dimensions of the plants and

their various parts can be easily obtained.
36. References have, when possible, been

given for each species to the following works :—

Name of Book. Abbreviation used in present Paper.

1. Flora of British India by Sir J. D. Hooker : 6 || 185 13),, IL,

2. Andropogonew by HE. Hackel m De Candolles’ Monogr. | H.
Phanerog., Vol. VI, 1889. |

3. Bengal Plants by D. Prain : : : ° ues

4, Flora of Bombay by T. Cooke . : : : 5 || Ce

37. The writer's most cordiai thanks are due
to Colonel Prain, Director, and to Dr. Stapf, Curator, of the Royal Botanic Gardens, Kew;
also to Captain Gage, Director of the Botanical Survey of India, and to Mr. E. G. Baker
of the British Museum, who have with unfailing courtesy given him the most generous

assistance on all points referred to them.

The writer has also to acknowledge the great assistance received by him from the
splendid herbarium, mainly accumulated and arranged by Mr. J. F. Duthie and containing
valuable collections contributed by many Forest Officers, which is now in charge of the

Forest Botanist at Dehra Dun.

[ 38 ]

HOLE : SOME INDIAN GRASSES AND THEIR G@COLOGY. 39

CHAP DT Be.

DESCRIPTION OF LOCALITY.

38. The locality in which the species included TPesraphy.
in the present paper have been chiefly studied comprises the Dehra Dun valley and the
northern slopes of the Siwalik range which bound that valley on the south, these together
forming a portion of the Dehra Dun Civil District, and also the southern slopes of the
Siwaliks which are included in the Saharanpur Civil District. This tract lies between
Long. E. 77° 40’—78° 22’ and Lat. N. 29’ 55’—30° 32’, and the principal part of it is
shown in the map which forms Plate I.

The elevation above mean sea level varies from 900’ to 3,000’. The Dun valley is divided
into two approximately equal portions, the Eastern Dun lying to the south-east of Dehra
cantonment which drains into the Ganges, and the Western Dun lying to the north-west of
the cantonment which drains into the Jumna. The Asan River flowing close to the
foot of the northern slopes of the Siwaliks constitutes the main line of drainage in the
latter, while in the former the Suswa River occupies a corresponding position. The chief
tributaries of the former are the Tons, Suarna Rau, Chor Khala and Sitla Rau and of the
latter the Bindal, Rispana, Song and Jakhan Rau, ali from the Himalayas. The upper
slopes of the Siwaliks consist mainly of sandstone and conglomerates with some clay beds,
the lower slopes and the entire Dun valley are covered with the débris brought down from
the Himalayas, or from the Siwaliks themselves, consisting of boulders and sand, mixed
with which are clay beds of varying thickness.

Where the impermeable clay bands crop out, swamps are often found, but owing to
the great thickness of the pervious boulder and sand deposits the subsoil water level is
frequently at a great depth below the surface. In the Dun valley owing to the masses of
débris brought down from the Himalayas the greatest thickness is naturally in the north
of the valley near the foot of the Himalayas, and Fernandez notes : “ It is for this reason
that the permanent level of water is at such a great depth, the well at the cutcherry at
Dehra being 228 feet deep.”” Another striking feature of the locality consists in the pecu-
liar character of the streams by which it is drained.

The tributaries of the Asan and Suswa rivers which rise in the Himalayas and Siwa-
liks and the streams which drain the southern slopes of the Siwaliks are, with the excep-
tion of one or two of the largest, such as the Song, dry during the greater part of the year
and their exposed beds of boulders and sand, often of great width, are then a conspicuous
feature of the landscape.

1 Working-Plan of the Dehra Dun Forests by BE. E. TSEN ED, 1889, p. 10.
eS 3) a

40 INDIAN FOREST MEMOIRS.

Fernandez remarks :—

“This absence of water at the surface is characteristic of all the streams of the
Dun, and is due to the great depth of the loose mass of boulders and shingle
into which the water sinks, to reappear only where the underlying imperme-
able band of clay comes up near the surface. Thus even the Suswa and the
Song rivers become dry for several miles of their intermediate course.”*

These streams become mountain torrents in the rainy season, quickly rising and falling
after heavy rain, and then bring down masses of boulders, shingle and sand which they
deposit as they flow along.

These streams also continually change their course and, leaving behind their former
alluvial deposits to gradually become covered with vegetation, proceed to erode new chan-
nels and form fresh deposits elsewhere, in the course of which areas often carrying valu-
able forest may be entirely swept away and fertile fields overwhelmed with boulders and
sand.

This continual denudation and deposition is more or less characteristic of the entire
Sub-Himalayan tract which contains some of the most valuable Indian Sal forests, and in
the locality now dealt with, omitting the dry wind-swept higher slopes and ridges of the
Siwalik range which carry comparatively valueless stunted forest, three main types of
topography, each of which is capable of supporting a distinct type of vegetation, can be
more or less clearly distinguished as below :—

I. Areas where the soil, at least during some months of the year, tends to become
more or less water-logged and which in their extreme development become
perennially wet swamps. The soil as a rule contains a considerable propor-
tion of clay.

II. The dry alluvial deposits of sand, gravel and boulders chiefly found in and near
the stream beds, where the water table is usually at a considerable depth
below the surface. These possess little power of retaining water or of rais-
ing the subsoil water by capillarity to replace that lost at the surface by
evaporation. They are liable to be excessively heated by solar radiation in
the day and in the hot season and to be excessively cooled by terrestrial radia-
tion at night and in the winter, especially where the free circulation of air is
impeded.

III. Areas where the soil is characterised by containing a considerable admixture of
clay, or humus, or both, and which varies from light to fairly stiff loam.
These areas are to a considerable extent confined to the higher plateaus owing
partly to the stiff soil offering resistance to erosion. They are as a rule well
drained anc the soil therefore retains water well, while the water lost at the

DROS pacar

E40) il

HOLE : SOME INDIAN GRASSES AND THEIR GiCOLOGY. AL

surface is readily replaced by capillarity. Both as regards supply of mois-
ture, degree of soil aeration and temperature, therefore, this type is interme-
diate between I and II.

39. The above types have, as a rule, been
clearly distinguished by practical Forest Officers and, in Working-Plans, I and IT are
usually described under the name of the recent or low alluvium and III under the name of
the old or high alluvium.

From the explanations given in paragraph 7 above it will be seen that IT is a typically
xerophytic and III a mesophytic locality. I is probably on the whole mesophytic although
for some types of vegetation it is hygrophytic.

The terms old and recent are apt to encourage the deduction that there has been in
every case, as regards the cecological characteristics of the soil, what may be called a develop-
mental, or progressive, succession of the same type and which has gradually resulted in the
conversion of the recent, or low alluvium, into the old, or high, alluvium.

To a great extent this is probably true, but it is obvious that we must clearly distin-
2uish two main types of succession, viz. :—

(1) That in which water-logged soil, usually containing a large admixture of clay,
becomes converted into well-aerated moist loam by a lowering of the water-
table and addition of humus.

(2) That in which dry sand becomes converted into moist loam chiefly through the
addition of humus.

It is equally clear that we must recognise the possibility of a regressive succession,
such as might arise for instance through the clearing of the forest growth and cutting off
the supphes of humus.

Mr. Millward* in an interesting paper has emphasized the fact that, according to
their mode of deposition, in still or running water, soils differ ab initio essentially in their
quality and composition owing to the size of the constituent particles, on which the distinc-
tions between gravel, sand, loam and clay depend. Consequently bare unoccupied areas of
soil of the same age may become available for the first time for plant colonisation which
may vary from practically pure clay to coarse gravel, and the fact that the heavy soil in
this locality is now chiefly found on the higher slopes and plateaus is probably due to its
having resisted erosion better than the light sandy soils.

It is important to realise these points, inasmuch as changes in the cecological character-
istics of the soil are directly connected with the succession of different types of vegetation,
and it has already been explained in the footnote to paragraph 14 above that, as regards
the most important type of vegetation in the locality here dealt with (viz. Sal forest), we

1 Indian Forester, Vol. XXVIII, p. 411.

[ 41 ]

4? INDIAN FOREST MEMOIRS.

must recognise not one, but several, possible modes of origin, differing according to the
5 a Seles
moisture content of the soil and to its initial sandy or clayey nature.

Climate. 40. As regards the climate of the locality
dealt with, the details given below, which have been abstracted from the meteorological
records of Dehra Dun station, give some idea of the prevailing conditions. Dehra Dun,
as will be seen from the map, Plate I, is approximately in the centre of the Dehra Dun
valley and it is situated at an elevation of 2,233 feet above mean sea level.

The area lying to the south of the Siwaliks receives less rainfall, averaging from 30 to

KE ; co) O . .

50 inches only per annum, and also possesses a more severe climate with a uniformly

higher average shade maximum temperature throughout the year and with a lower average
oto)

minimum shade temperature in December and January.

| -
> a 3 "9 es) q iS Description of Statistics,
- n =| \ S tol =| .
2 5 < a : : B Sas ie 2 g 4
= Ei x iy tes g = oe 5, Hots S | 2
a = os} oy | oS 3 =] o | A tH
E ele Niobe Fc Wh Ol ae
} |
| Par | "5 8:7 811 | Average monthly and annual
| fs lve 0; 85:5 | 840] 843 | 823) 750 | 6 ge monthly
662 | 692 797 | 90°6 | 95:0 | 93:0 | | means of maximum shade
| | | temperature in degrees
| | | Fahrenheit from observa-

i tions for period 1875—
| | | 1899.

a oy | : | iy ‘ Ke 22-7 m0): 60°6 51:8 456 ; 609 | Average monthly and annual
44-8 47-0 55°C | 643 | 70:5 | 744 | Tal | 4 IO | : means of minimum shade
temperature.

x wey I Dae oy, 1 | 7047) 62: *3. | 70-1 | Average monthly and annual
774 | 8176 | 822 (Sy || toe sor geet | ae ey means of shade tempera-
| | | | ture corrected to true
diurnal means obtained
| from the above maximum
| and minimum data, a cor-
|

rection obtained from
| hourly observation data
| being applied.

— : alle Sa : -, oF: en | 083 O21 068 79°63 | Average monthly and annual
2:26 2°04 116| 065] 1-45) 9:09) 2598) 26°31 Bet | vainfall (in inches) caleu-

| lated from all available
| | data down to 1900.

Per cent./Per cent.) Per | Per | Per | Per | Per | Per |Per cent. Per |Per cent.|Per cent./Per cent.
| cent. | cent. | cent. | cent. | cent. | cent. cent.

|

aR. : = BI oe, on . 0 2: 6 | 2: 62:8 63°8 | Average monthly and annual

669 619 4rd | 37-4 | 40:5 | 621 | 85°5 | 906 coe bee oie means of Lea humidity

| | | corrected to true diurnal

| means, based on the data

| | available for the period
| | ; 1877—1902.

The highest absolute maximum shade temperature recorded at Dehra Dun previous to 1903 is 111° registered in June.

3) “9° uy a iW
lowest 5 minimum .,, Pr ie Pic fhe as 33°9 55 » February.

[ 2 J

3 »>

HOLE: SOME INDIAN GRASSES AND THEIR (@COLOGY. Aes

Of the greatest importance for vegetation also are the maximum temperatures recorded in
the sun’s rays and the minimum temperatures on the grass, regarding which no reliable
figures are available for years subsequent to 1890.

The following figures show the maximum temperature recorded in the sun each month
during the three years 1888—90 :—

Maximum reading in sun’s rays by Black Bulb Thermometer.

Year. | January. February. March. | April. | May. | June. | July, | August. | September.| October. | November. December.
| hs | a
| - 2 peat a ll |
1888 126°°7 132°°5 146°'9 | 153°5 155°°5 | 161°:0 156°:0 | 155°-4 151°°5 142°1 135°:9 126°:0
1889 131 °5 133 °5 1444) 155°3 159 °0 IBY / 933 |) TBS} 97/ 157 :0 154:0 | 142:9 1360 129 -1
| |
1890 132 8 136 “4 143-5 | 157°8 | 157-5 | 155°3| 159°6| 152°8 512 | 143°6 132 6 130 °6
| ee ee { | tee ee!
Mean 130 °3 134 +1 144°9 | 155°5 | 137 °3 1579 | 158 +1 | 155°1 152 -2 | 142 ‘9 134°8 128 °6

The highest temperature recorded being 161° registered in June 1888. These figures
indicate the temperature which may be reached in grasslands surrounded by forest where
there is little or no circulation of air.

The following figures show the minimum temperatures recorded on the grass for the
same period, as recorded at Dehra Meteorological Station :—

Minimum reading on grass taken at 9-39 4.m.

Year. January. | February. March. April. May. June. July. August, | September. | October. | November. | December.
ahs wee aes | | | |
| | | |

1888 Hl 31°*2 36°°2 AG°*2, | 46° 1 5o7:2 63°°0 68°°6 67°:0 61°°3 AT°3 39°°7 35°-4

1889 | 30°0 | 30°3 43 °8 | 49 °3 58 ‘0 66°9 68 ‘1 68°8 56 ‘4 AT +2, 38 °9 | 32 °4
| | |

1890 | 32°1 | 34:7 43:8} 50:2 59 °8 65 *4 69-7 66 °5 59 °4 | AT 9 | 40°3 37 °1

| | | |

ns ee : | | ! aks

Mean | 32°83 | 33-7 44:8| 485 | 57-°7| 65:1) 68:8| 67-4} 59:0 | 47:5| 39-6 | 35-0

In many of the forest grasslands of the locality, however, considerably lower tempera-
tures are undoubtedly reached than those recorded at Dehra Dun Station. Frost is
frequently observed in these grasslands during December and January and is in many cases
the principal factor responsible for the absence of tree growth.

The period of maximum vegetative activity for most of the grasses dealt with in this
paper is, as might have been expected, the rainy season from June to September.’

It is remarkable, however, that almost all these species show a certain degree of activity
throughout the year, with the exception of the months December—January. In these

In the ease of one species, Imperata arundinacea, it is remarkable that maximum growth usually takes place in the peviod Marech—May.

[ 43 ]

44. INDIAN FOREST MEMOIRS.

months a general cessation of activity is more or less marked and is apparently caused
mainly by the low temperature.

As regards an explanation of the periodicity shown by the various species it is highly
desirable that statistics should be obtained showing, for the different classes of soil, the
actual average water-content of the surface layers of bare unworked soil monthly throughout
the year and especially during the period October to May inclusive. Such statistics would
indicate the net result of the opposing factors of, on the one hand, rainfall, dew deposition
and the raising of the subsoil water by capillarity, and on the other hand of loss through
percolation and evaporation at the surface, and this net content is of extreme importance
for vegetation.

Unfortunately no statistics of this nature appear to be at present available in India,
but observations on this point will now be made at Dehra Dun in connection with the study
of the cecology of Sal.

A point which has been frequently noticed in connection with several of the species
dealt with in this paper is that, in areas where the grass has been cut or burnt, vegetative
activity commences earlier than it would otherwise have done and the growth of the young
shoots is far more vigorous, especially during the period February to May, than in areas
which have not been cut or burnt. Forest Officers are well acquainted with this phenomenon
and know that areas burnt in the cold season are covered with a good crop of green grass
in the ensuing hot weather, when there are few or no green shoots to be seen in protected
areas. The clearing away of the old grass and dead leaves by fire enables the dew and
rainfall to at once penetrate the soil instead of being obstructed by, and evaporated from,
the old grass, and thus prevented from finding its way into the soil and becoming available
for the grass roots, while it also removes the injurious shade which to a considerable
extent retards the development of the young shoots.

soe ice 41. In the area considered in this paper
tion. examples of almost all the main types of vegetation mentioned in paragraph 26 above, both

of grassland and of woodland, are to be found.

Hygrophilous grassland is represented by well-developed reed-swamp communities of
Phragmites and Arundo and perhaps also by the wet Savannah Erianthus, while a type
of woodland which should possibly be ranked as intermediate between mesophilous and
hygrophilous is represented by some remarkable swamp forests containing such species as—-

Carallia lucida.

Cyclostemon assamicus.

Bischofia javanica.

Calamus tenuis. -

Of the Indian Evergreen Forest tracts.

As well as such temperate species as Quercus incana, Acer oblongum and Hedera
Heliz.

These swamp areas however are of comparatively limited extent and are not dealt
with in this paper.
[i Aare

HOLE: SOME INDIAN GRASSES AND THEIR (ECOLOGY. 45

The most important and widely distributed types of grassland in this locality which
it is proposed to consider in detail are briefly defined in the accompanying statement which
also indicates the corresponding type of woodland :—

ee SS lst

GRASSLAND, Wooprann

Serial No, |= —,

Type of Vegetation, | Description of Community. Type of Vegetation, | Description of Community,

-

Andropogon monticola . : (8)

a ; {| Calotropis procera.
Aristida cyanantha s ; .| | Xerophiloas Orthanthera viminea.
Zizyphus Jujuba.

|
| |
{ Triraphis madagascariensis, .| p Miscellaneous 4 | Adhatoda Vasica.
|
U

_ Xerophilous

3 {| Steppe.

| Acacia eburnea.
|

Saccharum spontaneum (xerophilous| | Forest. | Acacia Catechn.
form). | Dalbergia Sissoo,
Saccharum Munja (xerophilous |J
form).

| (| Zizyphus Jujuba.

| Adhatoda Vasica.

| Acacia Catechu.
Dalbergia Sissoo.

| Bombax malabaricum.

| Garnga pinnata.

| Odina Wodier.

| Phyllanthus Emblica.

| gle Marmelos.

4
| Zizyphus xylopyvra.
I
K

Mesophilous

Dry Misceila-
neous

Mesophilous

f |
II .4 | Moist j Saccharum Munja dominant 4
|| |

L J L

Forest. Grewia oppositifolia.

| Moringa pterygosperma,
Bauhinia racemosa,

| Murraya Koenigii.
Holarrhena antidysenterica,
Kydia calycina.

| Ehretia levis.

Savannah.

(| Mesophilous
|
4 Shorea robusta dominant.

{ Mesophilous

7
|

Moist | Saccharum Narenga dominant
| Sal Forest. J
J

l Sayannah.

Dalbergia Sissoo.
Garuga pinnata.
Holarrhena antidysenterica.
Kydia calycina.
Ehretia levis.
Grewia elastica.
Bridelia retusa.
Butea frondosa.
Stereospermum suaveolens.
(( | Bombax malabaricum.
| Mallotus philippinensis,
Erianthus Ravenne dominant . | Moist Miscel- 4 | Anogeissus latifotia.
/ |
|
L
|
L

Sy

ieee ae a

Mesophilous |
or perhaps |
Mesophilous
Hygrophilous
|
)

laneous Adina cordifolia.
Terminalia tomentosa,
Terminalia belerica.
Terminalia Chebula,
Gmelina arborea.
Eugenia Jambolana,
Cordia Myxa.

Cedrela Toona.

Celtis australis.
Trewia nudiflora.

Savannah.

is

[ 45}

Intermediate

Forms,

4.6 INDIAN FOREST MEMOIRS.

For the reasons noted in paragraph 39 above itxis perhaps inadvisable to employ the
terms “ old ” and “ recent alluvium ” but Vegetative type III may be said to occur chiefly on
what is often known as the high alluvium and Nos. I, II and IV mainly on the low
alluvium. Considering the parallel grassland and woodland together as a compound com-
munity the above communities may be more briefly defined as below where are also approxi-
mately indicated the conditions of soil and moisture, which, in the climate of this locality,
enable the various species to here exist together and to form more or less clearly defined
communities.

Community. Soil and moisture characteristics of Habitat.

I. Aristida—Acacia . ‘ : .| The soil is very dry ; it consists chiefly of gravel and stones and
has very little capacity for retaining moisture or for raising
it by capillarity.

Il. Saccharum—Dalbergia—Bombax .| The soil differs from that of No. I in containing a greater
proportion of small particles and is therefore more sandy but
it is still essentially dry.

(JI. Saccharum—Shorea ; ; The soil is distinctly loamy and moist, but well aerated and is
usually characterised by a more or less considerable ad- -
mixture of humus.

TV. Erianthus—Mallotus—Terminalia .| The soil is moist to wet, less aerated than that of No. II] and
with a more or less considerable admixture of clay. Owing
to the large proportion of clay this soil is apt to become
slightly water-logged in the rains and also to suffer from
drought after the rains, owing to the capillary movements
of water being very sluggish.

42. It must of course be clearly understood
that there is no sharp line of distinction between the above communities and just as the
factors of the locality, such as the character of the soil, extent of soil aeration, quantity of
soll moisture, etc., frequently change very gradually and vary very slightly from place to
place, we get Habieae with intermediate characteristics and more or less intermediate
minor communities connecting the main types.

Thus considering grassland first, the coarse shingle and oionel thrown up by the most
rapidly running currents provide a habitat for the typical desert, or steppe, community
composed chiefly of Aristida, with often scattered tufts of Andropogon monticola. As
the proportion of small particles increases in the deposits left by less rapidly flowing water,
and the soil becomes more sandy, the following species appear; the narrow leaved most
xerophilous form of Saccharum spontaneum, Triraphis, and the narrow leaved, more or
less stunted, most xerophilous form of Saccharum Munja, while Andropogon monticola
becomes more vigorous and numerous. As the proportion of sand increases and the capacity
of the soil for retaining more moisture and raising it from the water-table by capillarity

[ 46 }

HOLE: SOME INDIAN GRASSES AND THEIR COLOGY. AQ

slightly increases, owing te a small admixture of clayey particles, or organic humus,
Saccharum Munja becomes the dominant species. While intermediates between these two
communities thus occur, there is a marked dominance of Aristida on the very dry coarse -
grained soil and of Saccharum on the relatively moist and fine-grained soil. Similarly in
the Saccharum Munja savannah, in places where the soil is distinctly wet, usually on
account of a considerable increase in the proportion of clay, Erianthus appears and
gradually connects the moist with the wet savannah. EHrianthus, while approaching swamp
very closely, does not itself actually enter into the typical swamp community found on
perennially water-logged soil, but Saccharum spontaneum var. nepalense and Imperata
arundinacea var. latifolia both find a congenial habitat in the soft, spongy, perennially
thoroughly saturated soil of the marsh; other forms of both these species are found in the
moist savannah, and Saccharum spontaneum also occurs in the steppe. |

The Saccharum Narenga savannah is usually a very well marked type, but here also
‘small areas may occasionally be found where this species occurs in company with Saccharum
Munja and also with Erianthus Ravenne. Such an admixture is usually brought about
in nature as the result of a regressive change in the soil. When sal forest on light loamy
soil, for instance, is felled, previded the area is sufficiently large, the proportion of humus,
and consequently the capacity of the soil for retaining water, rapidly decreases, while the
soil also becomes dried by exposure to the sun and air currents. In such conditions
Saccharum Munja and Saccharum Narenga may be found together, the former eventually
becoming dominant and ousting the latter, if frost or other factors prevent the re-establish-
ment of woodland.

Similarly if moist, mixed forest on heavy soil (especially if it contains a considerable
admixture of Sal) is felled, the resulting decrease in humus content tends to make the soil
denser and thus less porous, while the rapid movement of water by capillary action is
checked. The soil thus tends to become less aerated and more liable to become somewhat
water-logged during the rainy season.

In such conditions both Saccharum Narenga and Erianthus Ravenne may coexist for
a time and these species may thus be found together. If the cleared area is extensive,
however, the drying action of sun and wind soon makes the habitat too dry for Erianthus
and it may be replaced by the moist savannah form of Imperata and Saccharum spontaneum
and by less strongly mesophilous species of Andropogon and Pollinia.

In dealing with this question of intermediate forms, also, it must be remembered that,
in addition to the very gradual changes in the intensity of the factors of the locality, we
must allow for the undoubted fact that the constitution and adaptations of a particular
species frequently enable it to thrive in essentially different habitats and to enter into the
constitution of essentially different communities. Thus Saccharum spontaneum enters
into the composition of the steppe—A ristida community, it forms a part of a typical reed-
swamp community, and finally may be often found in mesophilous grasslands.

Similarly Imperata arundinacea is found locally in both moist and wet savannah.
usually on heavy clay, which is liable to become water-logged during rain and to become
fr 4i> |

cL

48 INDIAN FOREST MEMOIRS.

rapidly dried after rain owing to poor capillary action, the oxygen content being apparently
too small for Saccharum Munja and the water content too small for Erianthus Ravenne,
these species therefore being unable to oust the less vigorous plant. On the other hand
Imperata is able to thrive and attains its maximum dimensions in a typical swamp, where:
neither of the larger species can penetrate.

This has already been alluded to in paragraph 22 above, where attention was drawn
to the fact that one and the same species may frequently be found to thrive well on a
physically dry soil, such as a sand or dense clay, and also on a physiologically dry soil,
such as the saturated, water-logged soil cf a marsh. Notwithstanding the fact then that.
intermediate forms occur, partly on account of the gradual shading away of the environ-
mental factors and partly on account of the fact that one and the same species may occur in
different habitats and communities, field workers will, it is believed, readily admit that the
four main types of grassland indicated above can be easily distinguished and that they
are separated by real differences, both as regards the conditions of the habitat and the
characteristics of the vegetation.

43. Turning now to the woodland com-
munities, the Forest Officer will, it is believed, readily admit the importance
of selecting and describing types, even though these be connected more or less
completely by intermediate forms. Thus, although four of the seven species given
for community No. I are commonly found also in No. II, there is a real difference.
between the open crop of No. I with more or less isolated clumps of Zizyphus,
Adhatoda, Acacia and Dalbergia, with bare spaces occupied by the shrubs Calotropis and
Orthanthera and with scattered tufts of xerophilous grasses, on the one hand, and the more
or less close crop of the dry miscellaneous forest forming type No. II on the other hand, in
which a number of additional species have appeared.

Similarly there is an undoubted difference between Nos. II and IV although many of
the species found in the former also pass into the latter. In IV, however, there is an
obvious preponderance of species which require a larger quantity of available moisture and
which are not found in No. II.

Again, although many of the species characteristic of No. IV occur also associated with.
Sal, there is no difficulty in distinguishing No. III in which the Sal is always the dominant.
species as a distinct type. Just as in the case of grassland, so here also we have species like
Bombax malabaricum and Garuga pinnata which grow well in physically dry soil and also.
in heavy wet soils and species like #gle Marmelos which thrive on dry sandy soil and also.
on hard clay.

The Forest Officer has long recognised this fact that certain species occur in very
different habitats in distinguishing what he calls indicating species, or indicators, which
are, as a rule, only found in places where a particular soil occurs, where a certain quantity
of moisture is available, or where a particular intensity of light prevails, and which thus
indicate the existence of certain factors and their intensity, from those termed indifferent

Eas 1)

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 4Y

species, which give no such indication. At the same time probably no plant community
consists entirely of indicators, considering the term in the present narrow sense, and hence
in our descriptions cf communities we must record the constituent species as we find them,
whether they be indicators or indifferent.

Moreover it is only by studying these so-called indifferent species in their various
habitats that we get a correct idea of the conditions necessary for their development.

Plates II—XII inclusive will give an idea of the physiognomy of the various com-
munities of grassland and woodland which have been briefly indicated above.

44. As regards furnishing a complete ex-

planation as to why each of the individual species mentioned is, in nature, able to form a
constituent part of the various communities indicated above, under the conditions ruling in
this locality, it is perhaps hardly necessary to point out that this will only become possible
when the cecology of each individual species has been thoroughly worked out in detail, and
this has not yet been done.

50 INDIAN FOREST MEMOIRS.

CSHVAC Pie wel

(1) DETAILED DESCRIPTION OF SPECIES.

Saccharum spontaneum, Linn. (Plates III and XIJI—XVIII.)

References. Jie Bip Ea WEIS alike}: Jel, IIL3}.
Pear 1188: | C., II, 948
Distribution. India, Afghanistan, Burma, Ceylon, Arabia, Syria, Africa, China, Java, Philippines,

New Guinea, Australia.

A.—DESCRIPTION.

Vernacular Kans, kansi.
Name. D
Culm. Erect or decumbent at base. Has been measured locally 14’ 2” (excluding prostrate or

subterranean portion) long but is said to attain 20’ (fide F. B. 1).

On dry sand very slender often not exceeding 0-10 in. diameter, but in favourable
localities attaining a diameter of 0-6 in. Solid above, fistular below, terete, indistinctly
striate.

Usually pruinose when young, polished when old.

Silky below the panicle and minutely silky below the upper leaf insertions.

Glabrous or minutely pubescent below the lower leaf insertions.

Leaf-sheath. Longer than proper internode, often with reddish or purplish blotches.

Villous at mouth, often minutely pubescent at base, otherwise glabrous or with scattered
adpressed hairs, sulcate.

Lamina. Erect. Of uppermost leat of flowering culm usually long, but varying from 2” to 3'
in length.

Of lower leaves, attains a length of 4’ (Roxburgh gives 7’).

Usually very narrow, frequently not exceeding 0-05” in width and then consisting of
a very narrowly margined concayvo-convex mid-rib, but attains a width of 0-7 in.
Greatest width about the middle.

Colour glaucous, mid-rib white.

Apex long acuminate, base narrow consisting of the mid-rib

Margin scabrid.

Often villous above at base immediately behind the ligule.

Ligule. Ovate to deltoid, base often sub-auricled.

Membranous, sub-acute or sub-truncate, often fimbriate when old.

Up tc 0:25 in. high. Minutely silky dorsally and ciliate.

Tnflores. Spikelets in pairs, one pedicelled and one sessile on the capillary jointed branches and
branchlets of a terminal panicle.

fp AeGaa

HOLE: SOME INDIAN GRASSES AND THEIR CECOLOGY. 51

Primary rachis sulcate, silky with long white hairs,

Primary branches sub-verticillate, simple or compound.

Flowering panicle 6”—2 feet long, conical or lanceolate to oblong, branches horizon-
tally spreading, or slightly ascending, usually reddish or purplish in colour, with the callus
hairs closely adpressed to the panicle branches.

Fruiting panicle is woolly and silvery white with the long wide-spreading white callus
hairs and the panicle branches are erect or ascending.

The horizontally-spreading callus hairs of the fruiting spikelet forms an efficient
parachute which aids its distribution by wind. The hairs of neighbouring spikelets becom-
ing entangled together, characteristic flocculent masses of several spikelets are often seen
being carried by the wind or hanging on the adjacent vegetation.

Awnless, lanceolate, 0-08 to 0:2 in. long. Spikelets.
Sessile and pedicelled similar, each one-flowered and ¢.

Pedicelled fruiting spikelet falling from the pedicel, the sessile spikelet falling later
with the attached pedicel and joint of axis.

Joint of axis longer or shorter than sessile spikelet, villous on margins, cr cn margins
and dorsally.

Pedicel 4 to 14 sessile spikelet (but usually shorter than spikelet), glabrous or ciliate,
shorter than proper joint.*

Callus hairs white from 14 to 7 times as long as sessile spikelet.

Lanceolate, the basal third thickened becoming hard and polished in fruit and more éiune I.
or less brown in colour, the upper two-thirds membranous hyaline. With two lateral nerves
from which the margin is inflexed. Apex entire or minutely bi-dentate. Margin ciliate;
dorsally with the upper two-thirds minutely adpressed-pubescent.

Broad-lanceolate, similar to I, but sub-keeled with one central nerve. Glume II.

Apex sometimes mucronate.

Margin inflexed and long ciliate.

Hyaline, nerveless, shorter than II, ovate-lanceolate, long ciliate, minutely pubescent Glume mI.

above dorsally.

Hyaline, minute, linear, ciliate, sometimes absent. In all fresh flowering specimens eiume w.
examined this glume has been found, in dry fruiting specimens the dissection of the inner
parts of the spikelet is somewhat difficult and it is possible that this glume is sometimes
overlooked.

Ovate, minute, ciliate, often shorter than the lodicules. Pale.
Two, cuneate, glabrous or apex ciliate. Lodicules.
Three, yellow, turning brown, 0-06 to 0-09 in. long. Anthers.
Two, purple, 0:05 in. long. Stigmas.

1 Proper joint = joint to which the pedicel in question is attached.

Ein

52 INDIAN FOREST MEMOIRS.

es The narrow leaves and slender culms, the long callus hairs and the brown coriaceous
base of glumes I and II are good field characters.

B.—Taxonomy.

45. Hackel " classifies the Indian forms of
this species under two sub-species which he defines as follows :—

Sub-species a. indicum.—Slender. Culm 3—4 mm. in diameter. Lamina narrow
linear (2—6 mm. wide). Ligule ovate, sub-acute, often sub-auricled at base.
Racemes slender, few-flowered. Spikelets 3—4 mm. long, callus-hairs white,
very soft, 4—6 times longer than spikelets.

Sub-species b. egyptiacum.—Robust. Culm 4—6 mm. in diameter. Lamina broad
linear (6-—14 mm. wide). Ligule truncate. Racemes stout, densely flowered.
Spikelets 46 mm. long, callus-hairs twice as long as spikelets (or a little
more),

The only Indian form placed by Hackel under egyptiacum is the variety nepalense which
he describes as having leaves up to 6—10 mm. wide, a linear-oblong panicle, spikelets 4 mm.
long and callus-hairs somewhat stiff, white, 24 times longer than the spikelets.

The present species varies greatly according to its habitat, as will be seen from the
biological and cecological notes given below, and in India the three following cecological
forms can be distinguished in the field although, connected as they are by numerous inter-
mediate forms, their separate definition under different names as sub-species or varieties
does not seem advisable :—

(1) The most xerophilous form found on dry sandy soil. Culms slender, usually less
than 0:2” in diameter, erect and tufted. Leaves exceedingly narrow, some-
times only 0-05” wide, callus-hairs not less than 34 times the spikelet. This
may for the present be distinguished as the sand-form.

(2) The most hygrophilous form found in swamps and marshes where there is an
abundance of available moisture more or less throughout the year. Culms
stout, 0:2” to 0-6” in diameter, usually decumbent at base and not tufted,
leaves broad attaining a width of 0-7", callus-hairs 14—3% times as long as
spikelets.

The fruiting panicle is elongate-elliptic to oblong with its branches usually
more persistent than in other forms.

This includes the var. neyalense of Hackel and may be distinguished as the
swamp-form.

(3) A form intermediate between (1) and (2), usually found on loam, with culms
more or less decumbent at base and not tufted, but less robust and with longer
callus-hairs than (2). This may be called the loam-form.

1 In Monograph. Phaneroyom. of DeCandolle, Vol. VI, 1889, p. 114.

HOLE : SOME INDIAN GRASSES AND THEIR (ECOLOGY. D8
Whatever characters are taken, however, whether the habit of growth, dimensions of culms,
width of leaves, length of spikelets, or of the callus-hairs, numerous intermediates connect-
ing the above forms can be found and observations in the field indicate that the characters
in question vary directly according to the quality and stability of the soil, liability to the
“ laying ” action of wind and water, and the quantity of available moisture in the habitat.

Plants of the above three forms of this species are now being cultivated in the Debra
Dun Experimental Garden with the object of testing the effect of soil and varying quantities
of available moisture on the characteristics of the plants. Some tufted plants taken from
dry sand and planted in loam have already, in a single growing season, produced some wide-
spreading robust culms which resemble those of the decumbent loam-form.

The African forms placed by Hackel under his sub-species wgyptiacum, variety
agyptiacum, difier from the Indian plants examined by the writer chiefly by their slightly
larger spikelets. This difference however is very slight and fails in the case of some African
specimens. Considering the great variability of the species in India it seems possible that
a more complete knowledge of the African plant will prove egyptiacum to be merely one of
several cecological forms which are defined by inconstant characters and which are con-
nected by numerous intermediates.*

Hackel himself notes that intermediates do occur between his sub-species indicum and
egyptiacum (l.c., p. 116). .

Roxburgh’s species Saccharum spontaneum, S. semidecumbens and S. canaliculatum*
are undoubtedly all referable to the single species described as S. spontaneum in this paper.
It is however important to note that in his descriptions of all these three plants Roxburgh
states that the corol is one-valved. Now glume IV is frequently present in this species,
although as a rule it is minute and very narrow, while the pale is also present, although
the latter is minute and often shorter than the lodicules. Hence when Roxburgh speaks
of a “ one-valved corol ” in his descriptions it must not be assumed that glume IV and the
pale are absent, but rather that these parts are small and inconspicuous. This is im-
portant, since Hackel (/.c., p. 119) rejects Roxburgh’s name of Saccharum Munja on the
ground that the description of that species as having a “ two-valved corol ” necessarily
implies the absence of the pale.

Roxburgh’s Saccharum eaaltatum is a doubtful plant. The sheet from Roxburgh’s
collection representing this species in the Herbarium of the British Museum is only named
in pencil and is believed to have been named by Solander. The plant on this sheet is
undoubtedly that described as Saccharum Munja in this paper.

1 The view taken in this paper regarding the limitation and definition of plant groups in descriptive botany is that it is neither advis -
able on the one hand to combine together distinct species or snb-species between which only isolated intermediates of probably hybrid
origin exist and which can therefore be distinguished in the field as distinct groups, nor, on the other hand, to separately describe and name
groups which are defined by inconstant characters varying with the cecological factors of the habitat (available moisture, soil, light, tem-
“perature, etc.) and which are connected in the field by numerous intermediates. Groups of the latter description can be sufficiently indicated
by such terms as swamp-forrm, sand-form, shade-form, ete.

2 Flora Indica, Wd. Carey, 1832, Vol. I, pp. 235, 236, 246.

fr RS.
Le

Habitat.-

5A INDIAN FOREST MEMOIRS.

Hackel refers this species of Roxburgh doubtfully to Saccharum arundinaceum Retz.
It is possible that Roxburgh’s remark in his description of this species “compare with
Saccharum arundinaceum ” (i.c., p. 245) is to some extent responsible for this identification.
It is not clear what Roxburgh meant by this, for there is no doubt that the Saccharwm
arundinaceum of Retz is the Saccharum procerum of Roxburgh. The unpublished plate of
Saccharum exaltatum, while clearly showing that Roxburgh’s plant is not Saccharwm
arundinaceum, is a very fair representation of Saccharum Narenga with which the culm,
leaf, narrow brownish panicle, length of joints of panicle branches, the brown colour of
glumes I and II and other characters agree well.

On the other hand the plant figured in Rhede’s Hortus malabaricus, XII, t. 46, is
almost certainly Saccharum spontaneum and Roxburgh himself considers Rhede’s plant to
be probably his Saccharum exaltatum (l.c., p. 245).

Against this identification with Saccharum spontaneum however is the fact that
glumes I and IT of Saccharum exaltatum are described and drawn as “ covered with much
long, cream-coloured soft hair” (/.c., p. 245). Roxburgh’s description “ root-creeping ”
while applying to forms of both Saccharum spontaneum and Saccharum Narenga does not
apply to either Saccharum arundinaceum or Saccharum Munja. On the whole therefore
it seems probable that Saccharum exaltatum Roxb.= Saccharum Narenga Wall. although it
is remarkable that Roxburgh makes no mention of the bearded leaf-insertions and hirsute
leaf-sheaths. It must however be remembered that the hairs on the sheaths of this species:
are deciduous.

C.— BIOLOGICAL AND CEcoLocicaL NotTEs

46. This is on the whole the most xerophilous:
species of Saccharum dealt with in this paper, a fact which is indicated by its exceedingly
narrow, hard and glaucous leaves. It is often found on dry sand associated with Triraphis,.
Andropogon monticola and stunted Saccharum Munja.

At the same time it is not sensitive as regards soil-aeration and like Imperata it can
thrive on stiff clay, while its maximum dimensions are attained (by variety nepalense) in
the water-logged soil of the marsh.

Its occurrence indicates that 1t requires even less moisture than Imperata.

It is frequently seen on loam which is too dry for a vigorous development of Imperata,
or which is too dry or too water-logged for Saccharum Munja, S. Narenga or Erianthus
Ravenne and it is there often associated with Vetiveria zizaninides Stapf (= Andropogon
squarrosus).

It has some capacity for standing shade and culms produced in the shade of trees and
shrubs show exceptional length and have been measured in this locality up to 12 feet 8:
inches.

Eee]

HOLE : SOME INDIAN GRASSES AND THEIR CCOLOGY. 55

47. This plant is perennial and usually Pevelopment
/ = ” of Culms,

eregarious. The culms originate from the basal nodes of older culms either above or below ae
the ground surface. A culm which originates below the ground surface may either at once Activity and
assume an erect habit of growth, in which case the young culms arise in the immediate ot
neighbourhood of the older culms and a more or less clearly defined tuft or clump results, or

else the young culm may grow for some time in a horizontal or oblique direction below the
vround surface, before its apex eventually turns upwards and assumes an erect habit of
erowth. In the iatter case the culms are widely scattered and no tuft or clump is found.

In this respect this species differs from the other species of Saccharum and Erianthus dealt

with in this paper in which each culm at first only grows outwards in a more or less hori-

zontal direction at the base for a very short and constant distance before eventually turning
upwards and becoming erect and in which a new bud at once arises and continues this
horizontal growth, as the older culm becomes erect. In these species therefore the growth
resembles that of a typical rhizome, whereas in S. spontaneum there is no well-defined
rhizome.

In the other species of Saccharum and Erianthus dealt with, also, the culms usually
arise close together and form well-marked clumps. In S. Narenga, however, a tendency
towards a spreading habit and the consequent production of ill-defined clumps is some-
times seen.

The present species in this locality is often found on sand in and near the beds of
water-courses and in such places the plant is lable to be covered by masses of sand and
débris brought down by floods. It is only under these conditions that the tufted habit
prevails and this appears to be an obvious adaptation which facilitates the rapid produc-
tion of aerial culms from the ground surface and prevents the plant being overwhelmed and
smothered by accumulations of débris and sand. In relatively stable soil, such as the loamy
soii of cultivated areas, the spreading habit prevails and enables the plant to extend and
occupy large areas far more rapidly than would be possible with the tufted habit of
growth.

In places where the plant is directly exposed to the rush of a water current, the spread-
ing habit is also common and this is to a great extent caused by the young culms being
“laid” by the mechanical action of the water, just as are often cereals by heavy wind.
Culms which are thus laid prostrate are often covered more or less deeply with sand and
root at the nodes and they then look exactly lke the ordinary subterranean rhizome
‘ranches. The more luxuriant the growth, the taller and larger the culms and the larger
the leaves. the more lable is the plant to be laid in this way.

The luxuriant form of the species (which has been named nepalense by some botanists),
therefore, which is often found in or close to the beds of streams is particularly liable to
this action and considerable areas have been seen where all the culms of this plant have been
laid perfectly flat and more or less covered with sand. There is little doubt therefore that
the spreading habit exhibited by this form is to a great extent caused in this way.

[ 55 J

56 INDIAN FOREST MEMOIRS.

The xerophilous form of the species, on the other hand, which shows much inferior
height growth, very slender, stiff and almost wire-like culms and very narrow leaves, as a.
rule resists the laying action of water well. At the same time isolated culms of this form
have been seen laid by water and rooting to a distance of 2—4 feet from the parent plant.

The above remarks on habit may be summarised as follows :—

(1) The species shows a markedly xerophilous form frequent on dry sand, a hygro-
philous form where an abundance of water is available and a form inter-
mediate between these two which is common on agricultural lands, wet
pastures, ete.

(2) The xerophilous form which often occurs on the dry sand deposits brought down
by torrents in the rains, is liable to be covered by accumulations of water-borne
(and possibly to some extent wind-carried) sand and débris, and it usually
shows a marked tufted habit which is advantageous in enabling the plant to
avoid being smothered. The stiff slender culms and small very narrow leaves
of this form enable it to resist the mechanical action of wind and water and
it is rarely “laid” in consequence. See Plate XIV.

(3) The intermediate form, common on agricultural lands and usually on loam, with
a more or less constant surface level, shows a spreading habit which enables
the plant to rapidly extend and occupy large areas. See Plate XIII.

(4) The hygrophilous form also shows a spreading habit and this 1s to some extent
at least caused by its liability to be “laid” by water currents and possibly
also to some extent by wind. See Plates III and XV.

Immature leafy culms of this species were cut over in the Dehra Dun garden and the
effect of the cutting on their growth carefully watched. The cutting was done in such a
way as not to injure the apical bud of the culm itself but to more or less destroy the green
leaves existing on the culm when the cutting was done. The first thing noticed was that
the destruction of the green leaves resulted in greatly diminishing the length of the inter-
nodes of the culm. See Plate XVIII, figs. 7 and 8. A series of abnormally short inter-
nodes preceded and succeeded by normal long internodes are frequently seen in the culms
of this and other species and these, as a rule, indicate that the leaves of such culms have
been more or less destroyed by the grazing of cattle, grass-cutting, or fire.

In order to study the susceptibility of any grass to fire-damage at any particular season
it is obviously necessary to be acquainted with the life-history of the culms and to know
what their usual state of development would be at that season. The development of the
culms cannot be always satisfactorily studied in the Experimental Garden, as it often tends.
to there become more or less abnormal. Direct observation in the forest throughout the
year is often not practicable and is in any case tedious. If, therefore, a method could be
discovered by which the age of a culm could be approximately calculated at sight, and if
its rate of development were known, it is obvious that a single observation would enable us

[ 3 J

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 57

to determine its size and state of development in any previous month (provided that growth
had been continuous) and this would greatly facilitate the inquiry.

The culms of most grasses are annual and if such an annual culm is examined it will,
as a rule, exhibit two or more very short internodes at the extreme base which can be clearly
distinguished from the subsequent much longer internodes. In other words such a culm
exhibits more or less distinctly two periods of growth—

(a) a preparatory period of slow growth;

(b) a subsequent period of vigorous growth.

The short internodes produced during the preparatory period carry only scales, or else
small and obviously under-sized leaves. The internodes produced during the period of
vigorous growth, on the other hand, carry well-developed green leaves. These internodes
are also, as a rule, much longer than those of the preparatory period, but in some cases
(especially in species like Saccharum Munja, the culms of which are biennial) several of
them remain as short as those of the preparatory stage. A culm in the preparatory stage
of growth is obviously merely a bud which possesses as yet no well-developed leaves and
for practical purposes the life-cycle of a culm may be considered to commence with the
period of vigorous growth, i.¢., with the first production of well-developed leaves which is
often (but not always) correlated also with the first production of long internodes and the
commencement of a marked increase in length.

As regards the development of the culm during the period of vigorous growth, obsery-
ations made at Dehra Dun on the sand-form of Saccharum spontaneum showed that in this
plant, in this locality :—

(1) The period of development of an internode carrying a well-developed leaf at its
apex approximately commences with the first appearance of the leaf at its
apex and terminates with the complete withering of that leaf (including the
sheath). This period can therefore be determined by noting respectively the
date of appearance and of the withering of the leaf at its apex. In this
species the period observed was five months. In one month therefore’ one
internode with the leaf at its apex completes }th of its life-cycle.

(2) The total number of internodes of the culm carrying well-developed leaves and
which are in active growth at one and the same time is 5 and therefore the
erowth of the whole culm during the period of vigorous activity proceeds at
the rate of one complete internode per month.

(3) The period between the commencement of the growth of one internode and of
the internode which immediately succeeds it is one month.

If we assume therefore that all the internodes carrying well-developed leaves require
the same period for their development and if we consider the life-cycle of the culm to
commence from the time when the terminal tuft of 5 green leaves each carrying an obvious
green lamina has become visible, it is clear that at the end of the first month the first leafy

Oe al

58 INDIAN FOREST MEMOIRS.

internode will be mature, at the end of the second month the second leafy internode will
be mature and so on, the terminal length carrying the panicle being four months old, when
the long internode immediately below it has become mature, at which time the erain is
usually ripe. Whether this explanation, which depends on the assumption that each in-
ternode carrying a well-developed leaf at its apex requires the same period for its develop-
ment, really explains the process and rate of development of the culm in Saccharum spon-
caneum, or any other species, can of course be only satisfactorily proved by observing the
exact time at which each individual internode commences and terminates its growth
respectively throughout the life of the culm, In the case of each species also such observ-
ations would have to be made on a considerable number of culms in order to obtain an
average result. Up to date no opportunity has been available for making such detailed
observations satisfactorily, but pending more precise investigation it was considered that,
for the purposes of practical work, the following principle might be accepted unless it was
found to be obviously at variance with the facts actually observed in the field : the number
of mature internodes, cach of which carries, or has carried, a well-developed leaf at its apex,
produced by a culm during its life-cycle is approximately equal to the number of months
constituting its period of vigorous growth, ie. to the approximate age of the culm in
months.

For the species dealt with in this paper, therefore, the average number of internodes in

a mature culm carrying, or which have carried, well-developed leaves! has been calculated
as follows :—

(a) The total number of internodes visible on the mature culm is first counted.

(5) By an examination of the young culms, the number of short basal internodes
carrying small scale-like leaves with little or no lamina is determined,

(c) Then (a)—(6)=the total number of internodes carrying more or lesg well-
developed leaves.

So far as possible the number of internodes obtained for each species under (c) was then
compared with the number of months of vigorous vegetative activity of that species, as
actually observed in the field in the locality from which the culms were obtained, and in all
cases where this could be done satisfactorily the agreement between the two numbers was
remarkable, while in no species was the period of activity obtained from the calculation
found to be in obvious disagreement with the facts observed in the field. Tt must be noted
that. in every case, culms which appeared to be of normal development were selected for
observation, i.¢. culms obtained from plants which had not been damaged by fire, grazing,
or grass-cutting and which had not developed in a locality where the ground-surface level
had altered, all of which factors are capable of causing abnormal development.

* Leaves are usually regarded as “ well-developed ” when they have a large green lamina, but it must be noted that the upper
leaves of the flowering culm (especially the leaf next below the panicle) often have a very minute lamina. Such leaves however aut
have a long sheath which to some extent functions ay a lamina, and, with reference to the point now under discussion, all such anynas eee,
have been considered to he « well-developed ” in this paper. Q

[ 58 9

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 59

The results regarding this point which have been obtained, however, up to date can only
be regarded as approximations for the following reasons :—

(1) The correct total annual period of vigorous growth for the majority of the culms
of a species in a particular locality can only be determined by observations
extending over several years, as this period fluctuates to some extent with the
seasons.

(2) The period required for the individual development of leaves and internodes
also depends to some extent on the character of the season and a correct
average can only be obtained from observations extending over several years.
Thus in years when the rains stop abruptly and the soil dries rapidly, the
normal period of development of certain internodes would be cut short.

As opportunity permits observations are now being made at Dehra Dun with the object
of determining more precisely than has been possible up to date the correct average period
of vigorous growth for each species in selected localities and the data thus obtained will
indicate how far the approximation which has been provisionally adopted as correct can

be relied on.

An average flowering culm of the form of this species usually seen in wet pastures
and agricultural lands shows about 9 (7—12) very short internodes at the base aggregating
about an inch in length and then 11 long internodes excluding the terminal segment which
terminates in the panicle. The leaves on the basal six short internodes have no green
lamina, which gives 14 internodes with green leaves and indicates a period of 14 months’
vegetative activity for the production of the culm, which may therefore be described as
annual. This form flowers in August—September. The most vigorous growth takes
place in the rains from June—September, but there is no marked season of rest and vege-
tative activity proceeds more or less throughout the year, at all events in the locality now
dealt with. The average culm therefore would start its deviopment at the beginning of
the rains in July and complete its cycle of 14 months at the end of August or beginning of
September.

The xerophilous form of the species common on dry sand has a slightly shorter period
of vegetative activity. Flowering takes place in August and the young culms commence
their development chiefly towards the close of the rains in August—September.

The average number of internodes bearing green leaves on the culms examined of this
form is 12, thus indicating a period of 12 months’ vegetative activity.

The flowering culms either die back to the ground after flowering, or their basal inter-
nodes persist and in the latter case secondary culms are usually developed from their

nodes.
fF 8% jl

Susceptibil-
ity to Fire
Damage.

Utility as
Fodder and
best Treat-
ment for
Fodder

Production.

60 INDIAN FOREST MEMOIRS.

48. The robust swamp form of this species
is not as a rule subject to fire damage and excluding this form the plants of this grass, say
about January, will usually be found to consist of —

(1) The old flowering culms which flowered in the preceding August—September,
and which are as a rule more or less dead and dry, with the exception perhaps
of a few of the lower internodes.

(2) The immature young leafy culms most of which vary in size from the just
developing buds to culms of 1—2 feet in length. The latter show four or
five mature long internodes and are seven or eight months old. These culms
still bear the persistent dead leaves on their lower internodes and terminate
in a tuft of five green leaves inserted on still young and immature nodes,
which are immediately below the apical bud of the culm.

The old flowering culms and the older of the young leafy culms which carry dry leaves
on their lower internodes are more or less inflammable, but the culms are usually slender
and the leaves very narrow so that the total quantity of inflammable material is compara-
tively small.

The damage done by fire consists in the destruction of the basal internodes of the old
flowering culms which usually remain alive and from the nodes of which young culms might
be developed and in the destruction of the immature leafy culms which would in due season
produce flower and seed.

In loamy soil the damage actually done is as a rule very slight for the following
reasons :—

(1) The plant there shows a spreading habit of growth in consequence of which the
culms are scattered and the intensity of heat developed at any one point is
slight.

(2) The soil retains moisture well and conducts heat badly so that the subterranean
culm-internodes, buds and young culms are well protected.

On the other hand, the tufted xerophilous form of the plant found on sand is liable to
somewhat severe injury. An early fire, while the soil is still a little moist, does little
damage, but a fire occurring late in the hot season may penetrate into the soil and destroy
not only the aerial part of the plant but also the tufted subterranean culms and thus prove
fatal.

Fig. 2, Plate XIV, shows a plant which was almost killed in this way by a forest fire
in May 1908, and other plants have been seen which were completely killed.

49. This plant is a favourite buffaloe-fodder,
especially the hygrophilous form with comparatively broad leaves common near streams,
and in such places firing is usually impossible. In drier places at all events. occasional
early firing would be probably beneficial by cleaning the clumps of débris, by destroying the

Eau

HOLE: SOME INDIAN GRASSES AND THEIR (COLOGY. 61

old flowering culms and encouraging the production of vigorous young shoots from the
eround. It has frequently been noticed that the secondary culms which tend to spring
fzom the basal aerial nodes of the old flowering culms are of inferior vigour to those which
originate at or below the ground surface. Duthie writes as follows regarding this plant :
“In the Jhang Settlement Report it is stated to be found in the moistest portions of lands
adjoining the rivers, where it affords most valuable pasturage for buffaloes. The zamindars
of those parts say that if there were no Kanh (local vernacular for ans) there would be no

01

buffaloes, and they consider it too valuable to be used for thatching.

D.—Economic Usss.

50. The leaves of this species are much used
for thatching.

This plant possesses considerable value on account of its capacity for fixing shifting
sand and unstable soil. Any node of a culm which is covered with soil as a rule at once
develops vigorous roots and usually also young culms, so that there is no danger of the
plant being smothered and killed by accumulations of sand and débris.

In places where it is subjected to the force of a heavy current of water also, although
the aerial culms are frequently laid flat, these rapidly root at the nodes and produce new
shoots, so that the plant is able to hold the soil with great tenacity and to resist erosion

well.

A question connected with this species which is of the greatest economic importance is
that of its eradication from agricultural lands. In the Central Provinces this species is
responsible for the fact that large areas of valuable soil are at present out of cultivation.

Agricultural soil is usually more or less loamy in which the plant assumes a spreading
habit and consequently firing in such cases would be of little value.

The only possible way of attacking the plant appears to consist in some remedy which
will starve the subterranean portions of the culms and thus prevent the appearance of
vigorous new shoots.

The following plan might in some cases be efficacious :—

(1) After flowering protect the areas from damage by fire or grazing and allow the
young culms to develop as vigorously as possible.

(2) About the middle of the following July the areas would ordinarily be covered
with a very heavy growth of grass including a proportion of very young
culms which commenced vigorous growth at the beginning of the rains. The
production of these young culms would tend to diminish the reserve materials
in the subterranean portions of the culms available for the production of new
culms. The grass should at this period be cut over as close to the ground as
possible and the cut grass should be left lying on the area.

} Fodder Grasses of Northern iets by J. F. Duthie, p. 25.
Lo

References.

Distribution.

Vernacular
Name.
Culm.

Leaf-sheath,

62 INDIAN FOREST MEMOIRS.

(3) This covering of cut grass by interfering with the access of light should to a

considerable extent prevent the production of vigorous young culms.
If leafy branches of trees and shrubs could be strewn over the cut grass the

latter would be kept in position and the shade increased.

(4) When the cut grass and branches become thoroughly dry in the cold or begin-
‘ning of the hot season they should be fired and the area ploughed if possible.

(5) If the grass was still found to be vigorous, it should be again allowed to develop:
and be again cut over in July before flowering but when a fairly vigorous
regrowth of young leafy culms had taken place and the same treatment
repeated.

This treatment would undoubtedly to a great extent prevent flowering and the spread
of the grass by seed, and as the production and development of leafy culms would he
practically prevented during half the normal period of vegetative activity annually, the
treatment should have a progressively weakening effect on the plant.

It is possible that in some cases firing in May followed by heavy and continued erazing
by buffaloes might be effective. In some localities it has been suggested that flooding the
fields to a depth of 2—3 feet might smother and kill the grass, but a possible result of such

treatment, which moreover is not easy to put into practice, would be to produce the most
vigorous form of the species, viz. variety nepalense.

saccharum Munja, Rovb. (Plates III and XIX—XXT_)

F. B. I., VII, 119 (under S. arundinaceum in part only).
H., 118 (under S. ciliare Anderss. excl. vars. 8 and y); P., II,.1189.
Chiefly found in Northern India, in the Punjab and Upper Gangetic Plain.

A.—DESCRIPTION.

Munj, munja, sar, sarkara, ekar (Dehra Dun).

Erect, attains a height of 18 ft. and dia. 0-5 in.

In dry unfavourable localities the height is much less, often not exceeding 6 or 7 ft.
Pale straw-coloured, smooth, striate and solid.

Shortly silky at extreme base, otherwise quite smooth, striate, pale straw-coloured.

Villous on margins at apex with long white hairs. Longer and usually much longer
than proper internode, uppermost sheath sometimes extending beyond the base of the
panicle.

[ 62 ]

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 63

Upper leaf of flowering culm 9”—23 ft. long, flat, tapering from base, long acumin- Lamina.
ate, width at base 0-2 in. to 0-4 in.

Lower leaves attaining a length of 7 ft. (8 ft. fide Roxburgh) and width of 1 in.,
but as a rule width not exceeding 0-75 in. and in unfavourable localities sometimes not
exceeding 0:13 in. wide.

Greatest width usually about the middle. Apex long acuminate and narrowed to-
wards the base. In basal leaves* the concave mid-rib occupies 4 or more of width of lamina
at base (often the entire width of the leaf).

Colour glaucous, mid-rib white. ‘The glaucous colour is caused by a thin covering of
wax.

Margin scabrid and also scabrid, often only on one side of the leaf, on 1 or more intra-
marginal nerves below, otherwise smooth but densely white villous at base behind the
ligule.
Truncate, usually a narrow membranous rim not exceeding 0:07 in. high. Of upper tigule.
leaves longer attaining 0-13 in.

Minutely silky dorsally and ciliate.

Spikelets in pairs, one pedicelled and one sessile on the capillary jointed branches and tforescence.
branchlets of a terminal panicle.

Primary rachis glabrous, sulcate, more or less scabrid on ridges.

Primary branches sub-verticillate, compound. Ultimate branchlets triquetrous, more
or less villous with long white hairs on angles and on two faces.
Flowering panicle 1—3 ft. long, usually lanceolate, pale cream-coloured to dark red-
dish-purple, branches spreading.
Fruiting panicle oblong, branches adpressed to axis, white to greyish-white in colour.
Awnless, lanceolate, 0:15 to 0:2 in. long. Sessile and pedicelled similar, each one spikelets.
flowered and ¢.
Pedicelled fruiting spikelet falling from the pedicel, the sessile spikelet falling later
with the attached pedicel and joint of axis.
Joint of axis triquetrous, + to subequal sessile spikelet but usually shorter than spike-
let, villous on two faces and on margins, villi overtop joint by once to twice length of joint.
Pedicels triquetrous 3 to 3 sessile spikelet. Majority of pedicels shorter than proper
joint, rarely subequal proper joint. Pedicels villous with long white hairs on two faces
aud on angles.
Callus hairs shorter than to subequal the spikelet.
Lanceolate, chartaceous, with 2 strong lateral nerves and usually 1—4 more or less Sessile
distinct, but fainter, additional nerves. Ga
Dorsally long villous on basal half or two-thirds, the hairs overtopping the glume hy
about the glume’s length.

1 Basal leaves are those which mature on the immature 1-year culms,

[ 63 }

Glume II.

Glume III.

Glume IV.

Pale.

Pedicelled
Spikelet.

Lodicules.

Anthers.
Stigmas.

Field
Characters,

Anatomy.

64 INDIAN FOREST MEMOIRS.

Scabrid dorsally on keels.

Margin inflexed, sparsely ciliate above.

Apex minutely bi-dentate to entire.

Subequal to I. Lanceolate, chartaceous, keeled. With one strong central nerve and
usually 2—4 more or less distinct, but fainter, additional nerves.

Glabrous dorsally or minutely pubescent towards apex, scabrid dorsally on keel—
margins incurved, ciliate above.

Apex usually shortly mucronate.

Oblong-lanceolate, hyaline-membranous, a little shorter than II, 1—3 nerved.

Margins incurved, ciliate.

Apex acute or short mucronate. !

Broad lanceolate to elliptic. Shorter than or subequal IT.

Hyaline, 1—3 nerved. Mucronate. Ciliate.

Mucro short to 0-05 in. long, but not exserted beyond the apex of the spikelet.

Ovate, hyaline, ciliate, from 4 to 2 IV.

Similar, but glumes I and II are both dorsally long villous and usually with 3—5
strong nerves and occasionally two additional fainter nerves.

Two, cuneate, glabrous, 0-02 in. long.

Three, pale yellow to purple, 0-08 to 0-10 in. long.

Yellow, often tinted with purple, 0-04 in. to 0-06 in. long.

The glaucous narrow leaves, awnless spikelets and smooth leaf-sheaths at once distin-

guish this species from Erianthus Ravenne, with which it is sometimes confused in the
field.

The striking contrast, as regards colour, afforded by the glaucous leaves of S. Munja
and the dark green leaves of S. Narenga, suggested that there might be considerable ana-
tomical differences in the leaves of these species. Examination, however, showed that
there were no anatomical differences which would account for this contrast, the glaucous
colour of the leaves of S. Munja being due to a superficial waxy deposit which is especially
copious on the under-surface. The general structure of the leaf and the disposition of the
chlorenchyma in these two species is similar and resembles that of Imperata described
below. The chlorenchyma is arranged in more or less complete rings around the vascular
bundles and is mainly disposed near the lower epidermis, the chlorenchyma strips flanking
the larger nerves, alone, reaching the upper epidermis, while a band, 4—6 cells wide, of
colourless bulliform cells is situated over the smaller nerves and the chlorenchyma which
surrounds them.

[ 64 J

HOLE: SOME INDIAN GRASSES AND THEIR C2COLOGY. 65

B.—Taxonomy.

51. Hackel (l.c., p. 117) distinguishes two
species mainly on the following characters :—

Saccharum arundinaceum Retz—Leaves broad linear. Width 2—3 cm. (0:8’—
1:2”). Mid-rib at base of lamina much narrower than width of lamina.
Joints of panicle usually longer than spikelet. Sessile spikelet 3-5 to 4 mm.
(0-:14” to 0:16”) long.

Saccharum ciliare Anderss. (=Saccharum Munja Roxb.).—Leaves narrow linear.
Width 3 to 6 mm. (0:12” to 0:24"). Mid-rib at base of lamina more than half
width of lamina. Joints of panicle shorter than spikelet. Spikelets
5—7 mm. (0:2”—0-28") long.

Both the above plants have been combined in the Flora of British India (VII, p. 119) in one
species named Saccharum arundinaceum Retz., but a comparison of Plate XIX with fig. 1,
Plate XXII, will indicate how distinct these two plants really are, and there can be little
doubt that they must be regarded as distinct species. Fig. 1, Plate XXII, is a photograph
of a plant of Saccharum arundinaceum Retz. growing in a garden at Dehra Dun which was
kindly identified in the first place by Dr. Otto Stapf and which the writer was subsequently
able to compare with a co-type in the Herbarium of the British Museum. As indicated in
the photograph Saccharum arundinaceum is a plant with strongly hygrophilous characters
and it attains gigantic dimensions. The writer has measured culms 24 ft. in length and
0-75 in. in diameter and these dimensions are probably exceeded in favourable localities.
The leaves are dark green and attain a length of 6 ft. and a width of 2 in. whereas those
of Saccharum Munja are glaucous and have never been found to exceed 1 in. in width.
The immature leafy culms of Saccharum arundinaceum show a solid mature, basal, piece
of culm of 5—6 ft. in length terminating in the apical tuft of green leaves, whereas in
Saccharum Munja the mature basal portion of the immature culms does not as a rule exceed.
2 in. The great length of the immature culms gives Saccharum arundinaceum a very
distinctive and characteristic appearance, totally unlike that of any other Indian species
of Saccharum with the sole exception of the cultivated Sugar-cane, Saccharum officinarum.
Saccharum arundinaceum is undoubtedly identical with the Saccharum procerum of Rox-
burgh and the latter was clearly acquainted with the characteristic habit of the plant
since he describes it as coming “nearest in appearance to Saccharum officinarum”
(l.c., p. 243). A peculiarity of this plant is that the spikelets (in the specimens seen by the
writer the pedicelled spikelets) are sometimes (?/ only in cultivated plants) 2—3 flowered
and inside glumes III and IV there are in such cases 1—2 additional hyaline, paleate,
glumes. Retzius (Obs. bot., fasc. IV, p. 14, and fasc. V, p. 16) says that his Saccharum
arundinaceum and Saccharum bengalense are practically identical except that the “ corolia
of the former is often 3-valued and of the latter 2-valved”” and there is little doubt that
[, Ga

66 INDIAN FOREST MEMOIRS.

these plants belong to the same species. Saccharum arundinaceum is a native of the
Evergreen Zone of India characterised by a rainfall exceeding 70 in., but it is not infre-
quently cultivated in gardens throughout India. The type of Retzius was probably a culti-
vated plant and he notes (J.c.) that it is “ colitur juxta sepes et ad stagnorum margines prope
Tranquebar.” It is indigenous in Bengal, Assam, and Burma, and extends to China. As
regards the specific value of Saccharum arundinaceum and Saccharum Munja respectively,
and as indicating that these plants are not cecological forms of the same species which owe
their peculiarities to variations in the available water supply, it is interesting to note that
the rainfall of Dehra Dun is 80 in., that Saccharum arundinaceum grows well there in
gardens not liberally supplied with water but is not found wild in the locality, that on the
other hand Saccharum Munja is indigenous and very common in the locality but that
although the latter has been found in habitats with great differences in the available mois-
ture supply and on soil varying from practically waterless sand to moist loam (which
results in a variation of the leaf-width of this plant from 0-13 in. to 1-0 in.) no indication
has been found of a tendency in Saccharum Munja to assume the characteristics of Sac-
charum arundinaceum.

As regards Saccharum ciliare Anderss. there is no doubt that this includes the plants
described by Roxburgh as Saccharum Sara and Saccharum Munja.*

Roxburgh’s drawing of Saccharum Sara at Calcutta represents a plant with an un-
usually dense, compact fruiting panicle. The density of the panicle however in this genus
is in itself not a character of specific importance and has been found to vary in one and the
same undoubted species. It depends largely on the age of the panicle and the way in which
it has dried or matured. There is moreover no authentic specimen of this plant named by
Roxburgh. There is a sheet from Roxburgh’s collection in the British Museum which has
been named Saccharum Sara in pencil, probably by Solander, and this is typical Saccharum
Munja Roxb. On the other hand the name Saccharum Munja is supported not only by the
description in Flora Indica and by an excellent drawing, but also by a specimen in the
British Museum named by Roxburgh. Hackel (l.c., p. 119) gives Andersson’s name pre-
cedence over the Saccharum Munja of Roxburgh on the ground that the description of the
latter plant as having a “two-valved corol ” necessarily implies the absence of the pale.

Tt has however been shown on p. 53 above that this inference is not justified. The
type also has been examined by the writer and there is no doubt that the pale is present both
in the sessile and pedicelled spikelets. Hence the name Saccharum Munja Roxb. must
stand and it is believed that Indian botanists will appreciate the restoration of this well-
known and appropriate namte. In his Saccharum ciliare Hackel includes a plant which he
describes as variety Griffithii and which is characterised by having glume II of the sessile
spikelet dorsally villous and not glabrous as in Saccharum M unja proper. In the Flora of
British India (p. 122) a new species is established under the name of Erianthus Griffithii

* Flora Indica, Kd. Carey, 1832, pp. 244, 46.

roe

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY, 67

Hook f. which includes Hackel’s var. Griffithii. The establishment of this species is the
result of including a specimen collected by Thomson at Rondu, Tibet, which has spikelets
with exvserted awns. This specimen however undoubtedly belongs to Hrianthus Ravenne
(a species which varies considerably as regards its dimensions and the width of its leaves
according to the amount of moisture available in the habitat) and consequently the species
Erianthus Griffithii cannot be maintained.*

Hackel’s variety Griffithit is identical with Saccharum Griffithii Munro, accepting the
plant, which was actually given this name by General Munro himself, as this species (vide
1193 Griffith and 666 Herb. Falconer in Herb. Kew).

The first valid publication of this name however appears to be in Boissier’s Fl. Or., V,
453 (1884), and this author, while unfortunately assigning specimens (Aitchison’s 467, 546),
which are undoubtedly conspecific with the numbers quoted above, to the entirely different
plant Saccharum Sara Roxb., applies the name Saccharum Griffithii Munro to a sheet in
his own herbarium (No. 507) which was apparently collected by Stocks (fide Hackel Mon.
Androp., 119), and which has not been seen by the writer.

It is possible that Boissier’s 507 is really referable to the Saccharum Griffithii of Munro
but its spikelets are longer (8 mm., fide Boiss., 7 mm., fide Hackel) than in any specimen of
the latter plant seen by the writer.

If Boissier’s plant is referable to a hitherto unpublished species his plant should,
under a strict application of the rules of priority, be called Saccharum Griffithii Munro,
but, under the peculiar circumstances of this case, it appears reasonable to keep this name
for the plant to which Munro undoubtedly gave it.

Under this view, if Boissier’s plant proves to be a different species, the date of valid
publication of Saccharum Griffithii will be that of the present paper.

A brief Latin diagnosis of this plant is accordingly given below.

Saccharum Griffithii is closely allied to Saccharum Munja and exhibits marked xero-
philous characters, but, although the writer has found Saccharum Munja in waterless
stations with leaves only 0-13 in. wide, no tendency to assume the villous glume II of the
sessile spikelet and other characters of Saccharum Griffithvi has been seen.

Seeing, therefore, that the plant, considered a variety by Hackel, is distinguished by
important and well-defined characters and has a distinct geographical range, it seems
preferable to deal with it as a distinct species under the name of Saccharum Griffith

Munro.

1 In F. B. I. (VII, 121) it is rightly pointed out that the separation of Saccharum from Erianthus solely on the character of the
exserted awn is unsatisfactory and that a careful study of the species constituting these genera is advisable with a view to testing their
validity. The plants concerned are not always easy to discriminate in the herbarium and a careful study of representative species in the
field is essential before a sound reclassification is attempted. It is hoped that the present paper will serve as asmall contribution to that
end. Whatever tho result of such study may be, the name Erianthus (@riffithii, which was based on an error (viz. the assumption that the
spikelets are awned), should obviously not be maintained.

It should be noted that, so far as Hrianthus Ravenne is concerned, the writer has found the exserted awn a reliable and constant
character, clearly separating it from all the species at present classified as Saccharums,

Cie

68 INDIAN FOREST MEMOIRS.

The following classification, therefore, of the forms now under consideration appears
to be the best :—
1. Saccharum arundinaceum Retz. (Obs. bot., fasc. IV, p. 14, 1786).
Syn.—S. bengalense Retz.
S. procerum Roxb.
Ref—Roxburgh, l.c., p. 243.
Hackel, l.c., p. 117, excl. syn. S. exaltatum.
Vern.—Ramsar (Dehra Dun), Teng (Bengal).
Habitat—Bengal (Darjeeling 3,500’, C. B. Clarke; Valleys of Sikkim 1—
4,000’, J. D. Hooker; Chittagong, 0—1,000’, Hooker and T hompson),
Assam (Griffith; Khasia Hills 2—4,000’, Hooker and Thompson),
Burma (Banks of Salween River), Southern India (Tanjore; culti-
vated or indigenous ‘).
Distribution —China.
Illustrations—See Plate XXII.

Notes—This plant is commonly cultivated in India in gardens and near
villages, as it is of some medicinal value. |

II. Saccharum Munja Rovzb. (1.c., p. 246, 1832).
Syn.—sS. Sara Roxb.

_ §. ciliare Anderss. in GEfvers K. Vet. Akad. Forhand. Stockh., p. 155,
1855. ;

Ref—Roxburgh, l.c., pp. 246, 244.

Hackel, J.c., p. 118, evel. vars. B Griffithu and y Boissiert.
Vern.—Munj, munja, sar, sarkara, ekar (Dehra Dun), Sara (Bengal).
Habitat.—Northern India in the Punjab and Upper Gangetic Plain, extends

into Sindh (fide Hackel). Is occasionally cultivated, e.g. near
Calcutta.’

Filustrations.—See Plates III and XIX—XXI.
III. Saccharum Griffithii Munro.

Syn.—sS. Sara, Aitchison, in Journ. Linn. Soc., XIX, p. 191.
S. Sara Boiss., Fl. Or., V, 453.
2 §. Griffithii Boiss., Lc.
S. ciliare var. Griffithii Hackel, l.c., 119.
Erianthus Griffithii Hook f. (in part), Fo By Eb Villy 122:

sah RU cation of the Districts of HughlimHowrah and the 24-Pergunnahs by D. Pyrain in ‘Records Botanical Sanne of Tina, Vol.
» No. 2, 5. I
as - |

HOLE : SOME INDIAN GRASSES AND THEIR CGCOLOGY. 69

Ref —Grithth, /tinerary Notes, 316.

Description —Cespitosum. Culmi 2 m., vel paulo altiores, farcti. Vagine
non hirsute, nodis imberbibus, laminis glaucis anguste linearibus
usque ad 8 mm. latis. Racemorum rhachis fragilis. Spicule ad
quemvis rhacheos nodum bine, altera sessilis cum articulo aecum-
bente demum decidens, altera pedicellata, a pedicello demum soluta,
utraque uniflora, ¢.

Spiculze 4—6 mm. longze, muticee.
Pedicelli articulique spiculis breviores.

Pedicelli plerumque articulos proprios subeequantes vel eis
longiores.

Callus villis gilvis spiculas subzquantibus vel eis brevioribus
dense vestitus. .

Gluma I* et II* et sessilis et pedicellatz spicule chartaceo-mem-
branaceze pilis glumas non, vel vix, superantibus dorso. villose.

Gluma III* dorso nunc glabra nunc parce hirsuta.

Gluma IV* mucronata. Mucro usque ad 1°5 mm. longus non ab
apice spiculz exsertus.

Afghanistan: Griffith, 1193.
Aitchison, 271, 467, 546.

Baluchistan : Stocks, 1209.
Punjab: Attock, Herb. Falconer, 666.
Sindh : fide Hackel, l.c., 119. |

Notes—Aitchison (J.c., 143) writes as follows regarding this species : “ Along
the edges of the dry water-courses, and on the higher island-like
plots of ground in the beds of these dried-up streams, Saccharum
Griffithii, a great coarse stiff grass, occurring in large tussocks, is
very striking, owing to the absence of other vegetation generally
than to any peculiarity of its own.”

; From what has been said above it will be clear that there has been some confusion
between’ S. arundinaceum, S. Munja, S. Griffithii and Erianthus Ravenne. As S. Munja
is of considerable economic importance it is obviously desirable that the characters distin-
guishing it from the other less valuable species should be clearly defined.

ea ee.

70

INDIAN FOREST MEMOIRS.

For this purpose and with the object of indicating at a glance most of the important
characters of the species of Saccharum and Erianthus dealt with in this paper the follow-

ing comparative table is given :—
|

Saccharom sponta-
neum (Indian form).

Saccharum
arundinaceum.

Saccharum Munja.

Saccharum Griffithii.

| Saccharum Narenga.

Erianthus Ravennz
(Indian form).

Frequently not form-
ing defined tufts.

Culms annual

Flowering culm solid
above, fistular below,
attains 20 ft. and
dia. 0°6 in.

Lamina glaucous, at-
tains 4ft.in length
(Roxburgh gives 7
ft.) and 0°7 in. in
width.

Midrib at base usu-
ally occupies more
than 3 width of
lamina.

Leaf-sheath not hir-
Bute.

Leaf-insertions
bearded.

not
Spikelets 0°08—0'2 in.
long (2—5 mm.).

Pedicel®’ ;—14 sossile
spikelet.

Usually shorter than |

sessile spikelet,

Joint longer or
shorter than sessile
spikelet,.

Pedicel shorter than
proper joint.’

Callus hairs white,
13—7 times as long
as sessile spilelet.

Forms well-defined

tufts.

Culms biennial (? or
triennial). Mature
basal portion of 1-
year (? or 2-year)
culms attains 6 feet
in length.

Flowering culm solid,
attains 24 ft. and
dia. 0°75 in.

Lamina dark green,
attains 6 ft. and
width 2 in.

In basal leaves!

midrib at base occu-
| pies 4 or less of the
width of the lamina.

Leaf-sheath not hir-
sute.

Leaf-insertions
bearded.

not

Spikelets 0°.1—0°15 in.
long (25—4 mm.).

Pedicel 4 to equal
sessile spikelet.

Joint usually longer

(% to 18
spikelet).

sessile

|

| Majority of pedicels

| shorter than proper
joint.

'Callus hairs pale,
shorter than to sub-
equal spikelet.

than sessile spikelet |

Forms well-defined

tufts.

| Culms biennial. Ma-
ture basal portion
of 1-year culms
| usually ‘not exceed-
ing 2 in,

| Flowering culm solid,
attains 18 ft. and
dia. 0°5 in.

Lamina glaucous, at-
tains 7 ft. in length |
and 1 in. width.

In basal leaves?
midrib at base occu-
pies 4 or more of
width of lamina.

| Leaf-sheath not hir-
sute.

| Leaf-insertions
| bearded.

not

| Spikelets 0°15—0°2 in.
long (4—5 mm.),

Pedicel 4—3 sessile
spikelet.

Joint usually shorter
than sessile spikelet |
(t to subequal |
sessile spikelet).

| Majority of pedicels |
shorter than proper |
joint (rarely sub-|

equal to proper
joint).
Callus hair pale, often
purplish, shorter
than to subequal
spikelet.

Forms well-defined

tufts.

Duration of culms not
known.

Flowering culm solid,
is said to attain 7
ft.

Lamina glaucous, nar-
tow. Has been seen
0°3 in. wide in her-
barium specimens,
but probably attains
a greater width.

Midrib at base usu-
ally occupies 34 or
more of width of
lamina.

Leaf-sheath not hir-
gute.

Leaf-insertions
bearded.

not

Spikelets 0°15-—0°24 in.

ong (4—6 mm.).

Pedicel 4--3 sessile
spikelet.
Joint 4—2 sessile
spikelet.

Majority of pedicels
subequal to longer
than proper joint.

Callus hairs yellow,
shorter than to sub-
equal spikelet.

Usually forms well-
defined tufts, but
sometimes spread-
ing.

| Culms biennial. Ma-
| ture basal portion
of 1-year culms
usually not exceed-
ing 2 in.

Flowering culm solid,
attains 16 ft. and
dia. 0°5 in.

Lamina dark green,
attains 6 ft. in
length and 1 in.
width.

In basal _ leaves?
midrib at base occu-
pies 4 or less of the
width of lamina.

Leaf-sheath hirsute
with bulbous-based
hairs.

Leaf-insertions long-
bearded.

long (2—33 mm.).

Pedicel 4—# sessile
spikelet.
| Joint 3—1} sessile

spikelet, but usually
shorter than to sub-
equal spikelet.

Majority of pedicels
shorter than to sub-
equal proper joint.

Callus hairs usually
purple, subequal
spikelet.

Spikelets 0:08—0:13 in,}

Forms

well-defined
tufts. ganed

Culms biennial. Ma--
ture basal portion
of 1-year eculms-
usually not exceed-
ing 2 in.

Flowering culm solid,
attains 19 ft. and 0°7~
in. dia.

Lamina dark green,.
attains 6 ft. in
length and width.
1°5 in.

In basal _ leaves?
midrib at base occu--
ples } or more of
width of lamina.

Leaf-sheath hirsute -
with bulbous-based
hairs.

Leaf-insertions short-
iy and finely beard--
ed.

Spikelets 0°13—0°16 in. .
long (83—4 mm.).

Pedicel 4—3 sessile -
spikelet.
Joint 4-3} sessile-
spikelet.

Majority of pedicels -
longer than proper-
joint.

Callus hairs purplish
or brownish, shorter -
than to subequal
spikelet.

SS SS SSS
? Basal leaves are those which mature on the immature 1-year (? 2-year) culms.
? Basal leaves are those which mature on the immature 1-year culms.
‘Jn recording the lengths of pedicels and joints of all species the basal pedicel and joint of each panicle-branch have been omitted as-
these often vary irregularly.

ww

HOLE : SOME INDIAN GRASSES AND THEIR CGCOLOGY.

-

il

se eR RR SR RS SR RR

Saccharum sponta-
neum (Indian form).

Hairs of joint overtop
joint by 2 to 3 times

length of joint.

Saccharum
arundinaceun.

joint by less than to
ii times’ joint’s
length.

|
|

|
|

Saccharum Munja.

| Saccharum Griffithii.

joint by once to)
twice length of joint.

| Hairs of joint overtop Hairs of joint overtop | Hairs of joint overtop

joint by once to

twice length of joint.

~Glume I sessile spike- | Glume I sessile spike-| Glume I sessile spike- | Glume I sessile’spike-

| Saccharum Narenga.

Hairs of joint overtop
joint by less than
joint’s length.

Glume I sessile spike- |

let coriaceous. Vil-
lous or not villous

dorsally. Villi not |

| overtopping the
glume.
Glume II sessile

spikelet coriaceous.

let coriaceous in let chartaceous. let chartaceous. | let chartaceous.
basal third. Not} Dorsally sparsely) Dorsally densely) Dorsally — densely |
villous dorsally. | villous. Villi over-| villous (in basal }—| villous (in basal
| topping glume by) #3). Villi overtop-| 4). Villi not
about 14 glume’s| ping by about overtopping glume,
length. glume’s length (1— | or overtopping by)
1} glume’s length). | less than } glume’s |
| length.
~GlumeII sessile spike-|Glume II _ sessile Glume II of sessile Glume II sessile |
let coriaceous in| spikelet chartaceous.) spikelet charta-| spikelet charta- |
basal third. Not| Not villousdorsally.| ceous. Not villous) ceons. Dorsally vil- |

villous dorsally.

~Glume III of sessile

spikelet not villous

dorsally.

Glume IV muticous

Pale ciliate.

-Glumes I and II pedi-
celled spikelet not

villous dorsally.

“Spikelet 1 flowered. | Spikelet

No additional
glumes inside III
and 1Y.

Glume III of sessile
spikelet not villous
dorsally.

Mucro of glume IV
0:03—0°05 in.
exserted beyond
apex of spikelet.

Pale ciliate.

Giumes I and II pedi-
celled spikelet dor-
sally villous.

Villi |

|
|
|
|

Not |

|

dorsally. |

|

Glume III of sessile
spikelet not villous |
dorsally.

Mucro of glume IV |
very short to (05 in. |
Not exserted beyond
apex of spikelet.

Pale ciliate.

lous (in basal 4 to
4). Villi
overtopping or over-
topping by less than
4 glume’s length.

sally.
Mucro of glume 1V
| apex of spikelet.

|
|

| Pale ciliate.

| Glumes I and I pedi- | Glumes I and II pedi-

overtopping spikelet |
by 1—1} times spike- |

let’s length.

sometimes
2—3 flowered with
1—2 additional pale-

ate, hyaline, mucro- }

nate glumes inside
TII and IV.

celled spikelet dor-
sally villous. Villi
overtopping spikelet
by about spikelet’s
length (—1; length
of spikelet).

celled spikelet dor-

sally villous (in
basal 34 to #).
Villi not overtop- |

not |

Glume III of sessile |
spikelet sometimes |
sparsely villous dor- |

very short to 0°06 in, |
Not exserted beyond |

Villous or not vil-
lous dorsally. Villi

glume.

Glume III of sessile
spikelet not villous
dorsally.

| Glume IV muticous.

| Pale ciliate.

| Glumes I and II pedi-
celled spikelet vil-
lous or not villous
dorsally. Villi not

ping or overtopping |
by less than} spiko- |

let’s length.

Spikelet 1 flowered. Spikelet 1 flowered. Spikelet 1 flowered.

No additional glumes
inside III and IV.

No additional glumes

inside III and IV.

No additional
glumes inside III
and IV.

not overtopping the

overtopping spikelet.

|

Erianthus Rayennz
(Indian form).

—_—-

Hairs of joint overtop
joint by twice lengt
of joint.

Glume I sessile spike-
let chartaceous.
Villous or not vyil-
lous dorsally, Villi
not overtopping
glume or overtopping
by less than 4 glume’s
length.

Glume II sessile spike-
let chartaceous. Vil-
lous or not villous
dorsally. Villi not
overtopping glume
or overtopping by
Jess than } glume’s
length.

Glume III of sessile
spikelet not villous
dorsally.

Mucro of glume IV
01 to 0:23 in. long.
Clearly exserted
from spikelet,

Pale glabrous,

Glumes I and II pedi-
celled spikelet vil-
lous dorsally. Villi
not overtopping or
overtopping by less

than 4 spikelet’s
length.
|Spikelet 1 flowered.

No additional glumes
inside III and IY.

xerophilous.

C.—BIoLocIcAL AND CEcoLocicaL NOTEs.

52. The comparatively narrow, hard, glau-
cous leaves of this species show marked xerophilous adaptations and although the plant
attains its maximum development on moist sand, it can exist in stations which are typically

_ Scattered individuals are thus often found on the shingle banks where Aristida is
dominant and on dry sand with the xerophilous form of S. spontaneum, but in such local-
ities the plants are more or less stunted and of inferior dimensions.

r

, 4

Ee

Habitat:

Development
of Culms,
Season of
Vegetative
Activity and
Flowering,

72 INDIAN FOREST MEMOIRS.

On sandy soil, however, which is moist, at least during the greater part of the rainy
season, this species is markedly dominant and it probably occupies a larger area than any
other savannah grass in Northern India.

It is found also on well-drained loam.

It appears to be decidedly sensitive as regards the aeration of the soil and plants taken
from sandy soil and cultivated in well-drained loam showed decidedly inferior develop-
ment, a fact which was not noticed in the case of T’riraphis for example.

It has less capacity for standing shade than either S. Narenga or Hrianthus Ravenne,
but still it can stand a certain degree of shade, as is shown by the fact that culms are fre-
quently produced under the shade of Zizyphus bushes on grazing grounds. The rapidity
with which this grass is killed out by Acacia Catechu is believed to be due not entirely to
shade but partly to the drying of the surface soil by the tree roots.

This species is here found at its best on the alluvial sandy deposits in the immediate
neighbourhood of the larger streams where the scil is not water-logged and where it can
establish itself on unoccupied areas, free from the injurious competition in respect of light
and moisture of earlier arrivals.

53. This plant is perennial and as a rule
markedly gregarious.

The plant consists of a much branched rhizome which constantly extends centrifugally
outwards in all directions.

The aerial culms are biennial and are either produced from the apex of the rhizome
branches or they arise as axillary shoots from the basal nodes of older culms, close to or
below the ground surface.

The culms are situated close together, the entire plant thus forming a well-defined
clump or tuft, with the oldest culms in the interior and the youngest at the periphery. The
latter as already noted is constantly extending outwards as the young culms continually
develop from the buds of the rhizome or from the base of older culms.'

The period of comparatively vigorous vegetative activity is approximately 10 months
from February to November. It usually flowers in October and the grain ripens in
November.

If a clump of this plant is examined in December—January it will as a rule be found
to contain—

(¢) A number of old dead culms which completed their life-cycle in the preceding
October—November.

(77) A number of immature culms bearing green leaves in which the culm proper is
as yet very short.

Owing to the very short internodes on these immature culms and to the leaf-insertions
being in consequence crowded close together each such culm resembles a tuft of leaves.

1 De ing the cold, or hot season, in areas which have been recently burnt the position of the youngest culms at the periphery can be

very clearly seen, as a ring of young green shoots then surrounds the blackened stumps of the old culms in the centre of the clumps
which haye been burnt.
eae

HOLE : SOME INDIAN GRASSES AND THEIR GECOLOGY. 13

These culms vary in size from the largest which have had the benefit of a complete season
of vegetative activity to the young buds which commenced their development towards the
close of the season of vigorous growth in September—November.

If one of the larger of these shoots is looked at more closely it will be found to consist
of a short hard basal portion of mature culm measuring usually 1—2 inches in length and
with as a rule 12 (9—16) short internodes which are clearly defined by the insertions of the
leaves. The leaves inserted on the nodes of this piece of culm are more or less dead and
withered and can be readily stripped from the nodes of the hard culm on which they are
inserted. Inside these we find one or two not yet fully developed green leaves and finally in
the centre is the terminal growing point of the culm situated about 0-25 inch about the
upper node of the mature basal portion of the culm.

The first two leaves produced at the extreme base of the culm are scales, or at all
events, have no obvious green lamina and omitting these it will be seen that the short piece
of culm matured during the first season’s growth carries, as a rule, 10 leaves. Growth is
resumed with more or less vigour in February and towards the end of that month young
culms may often be found in which the apical bud of the culm has been raised some 3 inches
above the surface of the ground owing to the growth and elongation of the upper immature
internodes. Towards the end of April culms have been collected in which all the inter-
nodes of the flowering culm can even then be clearly distinguished and in which the apical
bud has been raised 8—9 inches above the ground by the elongation of these internodes.
It is interesting to note also that this vigorous growth is taking place at a period when the
rainfall is very scanty.

The total number of long internodes developed in the flowering culm is usually 10,
excluding the apical segment which terminates in the panicle. Thus we have here each
year an apparently constant average rate of growth corresponding to a single leaf for each
month of the season of vigorous growth from February—November.

As an average culm which dies after flowering thus persists through two growing
seasons the culms may be described as biennial. No case has been seen in which a flower-
ing culm has been produced in one season and the time of flowering is as a rule very con-
stant. At the same time the exact period which an individual culm takes to complete its
cycle varies, owing to the fact that young culms may commence their development at any
time during the period of vegetative activity. For this reason also if a clump is examined
in the season of rest, the immature culms are found to vary in age and size from young
buds which have recently commenced development to the culms of one full season’s growth.

An exceptional type of development is very frequently scen in this species as follows.

After the usual number of short basal internodes which as a rule immediately precede
the production of the long internodes of the flowering culm, only a few long internodes are
found and then a second series of short internodes, after which follow the long internodes

1 Rare cases of culms of this species and also of Hrianthus Ravenne flowering in November, December and January have keen
met with which indicate that under exceptional circumstances these species are capable of vigorous growth practically throughout the year.

Ee

74: INDIAN FOREST MEMOIRS.

of the flowering culm proper. The basal series of short internodes in a normal culm is pro-
duced as a rule close to the ground surface, the initial long internodes of the flowering culm
being well above the ground. In the exceptional case noted above, however, the first series
of short internodes is deep below the ground, the second series is just below the surface,
while vigorous roots are developed from this second series and from the long internodes
below it.

The clumps of this species, which is usually found on alluvial soil in, or close to, the
beds of water-courses, are particularly liable to be more or less covered by masses of sand,
cr débris, brought down by floods in the rains and this is apparently the cause of the abnor-
mal development noted above.

At the beginning of the rains, the basal long internodes of the flowering culm have, as
a rule, already completed their growth, but when these are covered with soil or débris,
vigorous roots are developed from their nodes and this appears to check the development
of the flowering culm and to result in the production of a second series of several short
internodes near the surface of the ground, after which the normal long internedes of the
flowering culm may be produced the same season, or this may be deferred to the next growing
season. In the latter case the culm persists for three seasons.

If, when the change of soil level takes place, the flowering culm has so far developed
that some mature long internodes are still left uncovered, although roots are produced by
the submerged nodes, there is no development of a second series of short internodes, or
check in the growth of the flowering culm as noted above.

Exceptional development is also frequently caused by the fact that the leaves of this
species are largeiy cut and used for thatching. For this purpose the immature leafy culms
of the first season are cut over at a height of 1 to 2 feet above the ground. This operation
does not damage the low-seated apex of the culm itself, but results in the more or less
complete destruction of the upper green leaves which, in ordinary course, would have
nourished the long internodes of the flowering culm next season. Instead of long inter-
nodes and the flowering panicle being developed next season therefore, such a damaged
culm only produces a second series of some 10 short internodes, while the production of the
long internodes of the flowering culm is deferred to the third season. In such cases there-
fore the culm persists for three instead of two seasons and the base of the flowering culm
shows about 22 short internodes at the base, aggregating 2—4 inches in length, instead of
ihe usual 12 short internodes aggregating 1—2 inches. In areas where this grass is cut
for thatching purposes it has been noticed that there is frequently a marked periodicity in
‘lowering, an unusually large number of flowering culms being produced in one year, while
the next year very few or no culms flower. The cutting is usually done in October—Janu-
ary and it is clear that during the following rains each clump will contain (1) the culms
which were cut over and which are developing a second series of short basal internodes in
consequence, (2) the young culms which are in their first season. None of these will ordi-
narily flower that year but next year both (1) and (2), if they are not further damaged or

reed

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 75

interfered with, will flower and the clump will thus contain double the usual number
of flowering culms. A natural result of the presence of this unusual number of large old
culms is that less light is available for the healthy development of young culms of the
current season, of which therefore an unusually small number are produced. The natural
result is again an unusually small proportion of flowering culms the following year.

As the commercial munj is only obtained from the upper leaf-sheaths of the flowering
culm, it would appear from the above that the areas worked for thatching grass and munj7
should be kept distinct, in the former the grass should be cut over annually towards the
close of the season of vigorous growth and in the latter, damage to the leaves, by cutting or
otherwise, should be avoided as far as possible.

The mode of growth and development of the culms in Hrianthus Ravenne is practi-
cally identical to that described above except that the basal scales are usually four in
number. In old clumps of Erianthus, however, it has been noticed that the culms which
originate from the buds of the rhizome frequently show an average of 20 short basal inter-
nodes measuring 2—4 inches and such culms apparently persist for three years, whereas
the axillary culms which originate at a higher level show the normal development.

It is believed that such shoots commence their development at the break of the rains
in June, whereas the average shoot starts vigorous growth in February. This is possibly
on account of insufficient light being available, until the old dead leaves in the clump have
been beaten down by the rain and are more or less decayed. During the first season such
a shoot would be expected to show ten short internodes (four with scales and six with leaves
for the period June—November). During the second year, owing to the upper leaves reach-
ing a considerable height in the clump, activity might commence in February and an addi-
tional 10 short internodes might result for the period February—November. The flower-
ing culms eventually produced in the third season from these long persistent bases usually
show unusual vigour both as regards diameter, length and number of long internodes. It
is believed that similar development occurs in S. Munja.

54. The remarks given under this head for Susceptibil-

S. Narenga below apply also to this species as regards the kind of damage done by fire. Uamaees
The present species however is far more liable to severe injury than S. Narenga from both
early and late fires for the following reasons :—

(a) It usually occurs on sandy soil. In areas which are burnt, the rain washes away
such soil from the base of the clumps, more or less exposing the base of the
culms and the rhizome branches from which the young culms are developed.
Such soil also rapidly dries after the rains and the base of the clumps, which
as noted above are more or less exposed to the sun and wind, also become dry,
as do the dead leaves and débris contained in the clump. There is therefore
a far greater danger of the fire penctrating to the base of the clump and des-
troying not only the first year culms, but also the rhizome itself. On loam

Eto Tt

76 INDIAN FOREST MEMOIRS.

fire damage is greatly diminished and an early fire is often almost innocuous
on such soil.

(b) Owing to vegetative activity being fairly vigorous in February, the erowing
points of the young culms, commencing their second season's growth, are often
2-3 inches above the ground surface in that month, whereas in S. Narenga
they are as a rule still below the surface.

As in S. Narenga, the shoots which are most subject to fire damage are the one year
culms which, in the ordinary course, would produce the long, flowering culm and panicle the
following season. Thai early fires on sandy soil are capable of destroying these culms and
thus of greatly diminishing the production of flowering culms (and consequently of the
commercial munj fibre) has been observed again and again. Clumps recently burnt have
been examined in the field and the terminal buds of the one year culms frequently found to
be brown, discoloured and obviously damaged, while unburnt areas can often be at once
distinguished in the forest from adjacent burnt areas owing to the far more numerous
flowering culms in the former. This is most marked in November—-December before the
grass is fired and when the green areas which are annually burnt are sharply defined from
the adjacent unburnt tracts which resemble a white sea on account of the multitudes of
fluffy panicles.

Cases however do occur where occasional firing is apparently beneficial.

Mention has been made above of the fact that where the leaves of this grass are cut
for thatching purposes, the clumps, which are occasionally and not regularly cut, frequently
show congested growth and a very poor production of vigorous young culms, owing to the
presence of an unusually large number of flowering culms, the leaves of which prevent the
young culms at the base of the clump obtaining sufficient light for their development. The
burning of such clumps which contain an excess of old flowering culms, results in more or
less effectually cleaning the clump of the dead leaves, old culms and débris and is frequently
followed by a very vigorous production of young culms during the next season. In this
way occasional burning in areas worked for thatching grass is probably beneficial, but the
most satisfactory treatment for such areas probably consists in endeavouring to prevent
the production of flowering culms and to obviate the necessity for firing by cutting over
the immature leafy culms regularly every year. One year culms which are cut over usually
produce only leaves and short internodes again during the next season and by continuing
to cut these at a suitable height it should be possible to prevent the production of flowering
culms.

Intentional burning should be carried out early in the season before the clumps get too
dry, as otherwise there is a danger of the rhizome being destroyed by the intense heat.

Any factor which interferes with the natural outward growth of the clumps and
causes the young culms to appear inside the clump and under the shade of older culms,
instead of just outside the clump at the periphery, as is naturally the case, also results in
a similar congestion of growth and inferior production of vigorous young culms. Such

[ 7 J

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 77

factors may for instance consist in the presence of large rocks in the soil, the proximity of
other plants of the same or other species, the hardening of the soil around the clumps by
heavy grazing and trampling of cattle and the destruction of the young culms developed
from the external rhizome branches by the trampling or browsing of cattle. It is probable
that in all such cases of congestion temporary relief would be secured by firing, but this
would be of no permanent utility.

It is obvious that a clump which has been well burnt one year would be cleared of
debris and would probably have most of the one year culms also destroyed. Next year
therefore we should expect the clump to consist chiefly of immature leafy culms and the
only inflammable material in the clump would consist of the old leaves at the base of these
culms, while there would be a quantity of green leaves. Such a clump therefore would not
burn well and it is a frequent experience in burnt areas that a year in which the clumps
burn well is followed by one in which they burn with difficulty. The greater the accumu-
lation of dry flowering culms the more injurious the fire is likely to be and it is probable
that on fire lines the plants would be most injured and weakened if they were cut over for
thatching grass every third year. The second year after such cutting there would ordinar-
ily be twice the usual number of dead flowering culms in the clumps.

55. The above remarks on fire damage as
regards this species may be summarised as below :—

(a) Fire damage results chiefly in the destruction of the one year culms and in
diminishing the production of the commercial munj fibre. It is however
often sufficiently severe to damage the rhizome branches and may thus kill
the plants entirely.

(b) The damage done is undoubtedly most severe on sandy soil and on loam the
damage done by an early fire is as a rule insignificant.

(c) The later in the season the fire occurs the more severe the damage, owing to the
greater dryness of the dead leaves and débris in the clump.

(d) Area: required for the production of mwnj should be carefully protected from
fire especially if on sandy soil, and also from damage to the leaves by grass-
cutting, grazing or otherwise.

(e) Areas worked for thatching grass should be kept distinct from those required

for mun7.
In such areas the grass should be cut annually to diminish as far as possible
the production of flowering culms and if heavy flowering does occasionally
occur the clumps should be burnt early in the season. The soil around the
clumps should be kept Icose to allow of the free growth outwards of the
rhizome and to prevent the young culms and ends of the rhizome branches
being situated close to the inflammable material inside the clump.’

1 The mode of development of the clumps and culms and the occurrence of congested growth in this and other closely allied species
-of Saccharum and Hrianthus have many points in common with what is seen to obtain in tufted species of Bamboo, as will be seen by a
reference to a most interesting paper by Mr. W. H. Lovegrove on Derdrocalanus st-ictus in the Indian Forester, Volume XXVI,

p. 433.
eae |

Utility as
Fodder and
best Treat-
ment for
Fodder
Production.
Relations of
Grassland to
Woodland.

INDIAN FOREST MEMOIRS.

~I
16 0)

56. This plant is as a rule not much utilised
for fodder and cattle only eat the young and immature leaves. Grasslands required for
orazing should be fired so that the young leaves may become available in the hot season.

57. On areas where this grass is dominant
the soil and moisture conditions are as a rule only suitable for dry miscellaneous forest, as
a fair representative of which we may take the woodlands of khair, Acacia Catechu. The
latter has been seen to obtain a footing and effectually oust S. Munja from areas where the
latter had first occupied the ground and become the dominant species.

A plant which frequentiy occurs with this grass is Zizyphus Jujuba and on areas
which are regularly fired and heavily grazed throughout the year, the latter tends to become
dominant. In such areas the Saccharum is unable to develop, the young shoots being
vrazed down by the cattle with the exception of the few culms which are able to develop
under the protection of the Zizyphus. The latter however is able to persist although the
repeated grazing and fire damage prevents the plant becoming more than a shrub. The
seed of the Zizyphus is largely distributed by jackals and Mr. P. H. Clutterbuck has
pointed out that the latter tend to frequent grazing areas, they being followers of the
larger beasts of prey and also attracted by the carcases of cattle which happen to die, and
this circumstance probably favours the rapid extension of the Zizyphus in such areas.

D.—Economic USES.

58. This species is of great commercial value
and is probably most widely known on account of the valuable fibre (called munj) extracted
from the upper leaf-sheaths of the flowering culm, which is used for cordage, mats, etc.
For this purpose, as a rule, only the two uppermost leaves of the culm are utilised, as these
have the longest sheaths. In an average culm measuring 17’ 3”, the topmost sheath
measured 3/ 10”, the next below was 2’ 11” and the next 2’ 1” which indicates how the length
of the lower sheaths diminishes.

Watt writes as follows regarding this fibre:—“ The much prized mun] is strong,
elastic and has a wonderful power of enduring moisture without decaying. It is exten-
sively employed in the manufacture of cordage, ropes, the famed Delhi mats, and in the
preparation of baskets, etc. Munj mats * * are largely produced in Allahabad, Agra,
Delhi, and are traded in all over India, and within recent years have begun to find their
way to Europe.”

The thin upper portion of the flowering culm is locally called sirki and is used for
making winnowing trays, cart-covers, etc. The thick lower portion of the culm is locally
called bind and is used in place of split bamboos for making screens (tatties) and in
thatching.

1 Commercial Prcducts of Irdia by Siy George Watt, 1£08, p. 930.°

[ eal

HOLE: SOME INDIAN GRASSES ANI) THEIR G@COLOGY. 79

The leaves are largely used for thatching and also constitute 2 valuable paper material.

Mr. William Raitt, the paper expert, in an article dealing with the relative value of
possible paper materials alludes to this grass as follows :—

* It is impossible to deal at length with all the fibrous grasses known, but as types of
the classes most worthy of attention we may refer to threc species which have already
proved of value for paper-making.

Muriz (Saccharum Sara)—a reed-like grass found on chur and waste lands in
Northern India. Very gregarious in habit, growing in dense thicket-like masses, which
can be cut and collected at low cost. Being generally found in the neighbourhood of rivers,
the economic radius of collection is widened by the possibilities of water transport.
Yields an excellent, easily bleached pulp, similar to that of wheat straw.”

This species also has considerable value on account of its power of fixing unstable soil
and of preventing the drifting of sand. It is not infrequentiv found in stream beds where
it has successfully resisted the force of the water, each clump forming a nucleus for the
accumulation of sand and débris.

Duthie writes as follows on this point: “ At Jeypur it is extensively used as a sand-
binding plant; and the experiments which have been undertaken there for arresting the
movement of sand prove it to be very suitable for the purpose.”

Saccharum Narenga, Wallich ex Hackel in Monogr. Androp. (1889), p. 119. (Plates XII
and XXIII—XXV.)

Syn. Saccharum porphyrocomum, Hackel, l.c., p. 120.
? Saccharum exaltatum, Roxb., l.c., 245.
12, 18), JL, WUE 120s sean: References.
JP, INE, wes.
The Sub-Himalayan tract from Dehra Dun to Assam, Bengal, Central India and Distribution.
Burma. Extends to China.

A.—DESCRIPTION.

Kanwal (Debra Dun), Bata (Goalpara, Assam), Barota (Sibsagar, Assam). OEE
Erect, attains a height of 16 ft. and diameter of 0-5 in. Solid. Culm.

Quite smooth and polished except immediately below the leaf insertions and the base
of the panicle, where it is densely bearded with long adpressed white hairs.

Covered with bulbous based hairs, especially towards the apex; the hairs easily break Leaf-sheath.
off and old sheaths are often rough with their persistent bases.

1 Tropical Agriculturist, Vol. XXXIII, July 1909, pp. 10--12.
2 Fodd2r Grasses of Northern India by J. F. Duthie, p. 25.

[yi |

80 INDIAN FOREST MEMOIRS.

Lower sheaths are longer, upper sheaths are shorter than the proper internodes.

Lamina. Of uppermost leaf of flowering culm from mucroniform, 0-25 in. long, to linear, 14 in.
long and 0-25 in. wide.

Of lower leaves, attains a length of 6 ft. and width of 1 in. Colour dark green, midrib
white.

Greatest width usually about the middle. Apex long acuminate, base narrow ana flat.

In basal leaves midrib occupies 4, or less, of the width of lamina at base.

Margin scabrid except at extreme base.

Bulbous hairs scattered chiefly on upper surface towards the apex.

Densely bearded above immediately behind the ligule.

Ligule. Membranous, rounded, up to 0-25 in. high.
Ciliate and minutely silky dorsally.
yallores cence: Spikelets in pairs, one pedicelled and one sessile on the capillary jointed branches and
branchlets of a terminal panicle.

Primary rachis yellow, sulcate, hairy on ridges, bearded at nodes.

Primary branches sub-verticillate.

Flowering panicle 9 in. to 2 ft. long, narrow, lanceolate to sub-cylindric; branches
horizontal or slightly ascending, simple, or sometimes branched at base; colour purplish-
brown.

Fruiting panicle purple, branches closely pre to rachis and very fragile.

Spikelets. Awnless, oblong, 0:08 to 0:13 in. long.
Sessile and pedicelled similar, each one-flowered and ¢ .

Pedicelled spikelet at length falling from the pedicel, sessile falling with the attached
pedicel and joint of axis. The pedicelled spikelets usually fall first and the sessile follow

later.
Joint of axis 3-gonous, sparsely hairy on angles, 4 to 14 the sessile spikelet but usually

shorter than to subequal the spikelet.
Pedicels glabrous or hairy on margins, 4 to 3 sessile spikelet. Majority of pedicels
shorter than to subequal the proper joint.
Callus hairs stiff, usually purple, shorter or a little longer than ne spikelet.
Glume I.—Oblong to oblong-lanceolate, brown, coriac2ous, apex sub-acute, obtuse or
minutely bi-dentate.
With two lateral keels; 2—7 nerved.
Margins inflexed, ciliate.
Dorsally flat between the keels, hairy or smooth below, puberulous above.
Villi not overtopping the glume.

[ 80 ]

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. SL

Glume II —Subequal I. Oblong to oblong-lanceolate, brown, thinly coriaceous, apex
membranous, sub-acute or obtuse.
A central keel, 3—5 nerved.
Margins inflexed, ciliate.
Dorsally glabrous or hairy below, puberulous above. Villi not overtop-
ping the glume.
Glume III—Subequal I. Oblong, hyaline, enerved, or with one or more partial
basal nerves, ciliate.
Glume IV.—Similar to III, but slightly shorter, enerved or with a partial median
nerve.
Pale—+ to 4 IV, ovate to oblong, hyaline, enerved, ciliate.
Lodicules, two, cuneate, glabrous, 0-02 in. .
Anthers, three, 0:05—0-09 in. long, yellow or more or less spotted with purple.
Stigmas, two, 0-03—0-04 in. long, dark purple.
The hairy leaf-sheaths, the culms bearded at the leaf insertions, the narrow, purple, pie enarac-

very fragile, fruiting panicle and the dark green leaves readily distinguish this species in ‘*™*
the field.

B.— Taxonomy.

59. Hackel divides the genus Saccharum
into four sub-genera, two of which contain all the Indian species.
These two sub-genera (sections) have been adopted in the F. B. I. as follows :—

I. Eusaccharum.—Rachis of spike (Hack. raceme) fragile. Spikelets of each pair
subequal, sessile and pedicelled, both fertile (Hack. ¢). Stem solid.

II. Sclerostachya—Rachis of spike tenacious. Both spikelets of each pair pedi-
celled. Stem fistular.

Hackel places only one Indian species in the latter, vig —

S. fuscum Roxb. This is adopted in F. B. I. where a second Malayan species is also
added, viz. S. Ridleyi which Hackel described after the publication of his monograph.

The characters of the culm being fistular or solid and the rachis tenacious or fragile
are therefore seen to be of sectional rather than specific value.

The flowering culms of the present species are very subject to damage by insects and
fungi. Culms which have been thus injured usually show a small perforation which is
apparently caused by some sucking insect, while the tissue of the culm in the neighbourhood
of the hole becomes discoloured and usually reddish giving the appearance of the “ red-
rot” of the Sugarcane.

Specimens of the damaged culms were submitted to Dr. Butler, Imperial Mycologist at
Pusa, but although fungal hyphz were found in the discoloured ti:sue these apparently
belonged to a saprophytic species, as no parasitic fungus could be isolated on incubation.

f 9

$2 _ INDIAN FOREST MEMOIRS.

The question as to whether, or not, a sucking insect is invariably the primary cause of the
injury however requires further investigation.

The injury is most frequently found at the leaf insertions where the insect, or fungus,
probably intercepts the food materials on their way from the leaf into the culm. This
iaterference with the normal nutrition of the culm results in the internode next above the
attacked leaf-insertion becoming fistular instead of solid and being unusually small, weak
and more or less imperfectly nourished. When the internode is much weakened in this
way it is frequently unable to withstand the action of the wind and the upper portion of
the culm, with the terminal panicle, then bends over, but usually still remains attached to
the plant. Such a culm looks as if it had been half-broken by a passer-by. The panicle
branches also of culms injured in this way are remarkably tenacious and the spikelets are
persistent. Such a panicle therefore forms a striking contrast to a normally matured
panicle and has a characteristic and distinctive appearance both when seen in the forest
and in the herbarium. The normally-matured panicle shows a persistent main rachis
with closely adpressed fragile branches which rapidly break up and fall off, the pedicelled
spikelets falling first from their pedicels and the sessile spikelets following with the joints
of the branches and branchlets, while, so Jong as the spikelets remain, the callus hairs give
the panicle a purple colour. The abnormal panicle, on the other hand, shows persistent
and often spreading branches and persistent spikelets, while the callus hairs are frequently
pale to almost white and thus give the panicle a pale, or brownish, feathery appearance.
See Plate XXIII, fig. 3, and compare with the normal panicles shown in fig. 2.

In these abnormal flowering culms of this species therefore we have a combination of
two remarkable and important characters, viz. the fistular culm and tenacious rachis of the
panicle branches, and the writer has seen panicles of this description placed under S. fuscum
Roxb." in herbaria. The latter species, however, apart from other characters, can be at once
distinguished by the fact that both spikelets at each node are clearly pedicelled and the
spikelets eventually fall off from the tips of the pedicels, leaving the persistent panicle
branches intact, a state of affairs which is not seen in the abnormal panicles noted above.
See Plate XXV, fig. 11.

Whereas the spikelets of a normally-matured panicle of the present species contain
the normal fully-formed fruit (grain), those of the abnormal panicles either contain a very
small and imperfectly developed fruit, or else the entire unexpanded flower with ovary and
stamens, thus indicating that the development of the flower and fruit has been prematurely
checked.

The glumes of these abnormal panicles show no marked peculiarity. Saccharwn
porphyrocomum, Hackel, is founded on No. 19285 Herbarium H. F. Hance. The type is in
the Herbarium of the British Museum and has been examined by the writer. It consists

1 In F. B. 1. Saccharum fuscum Roxb. is said to occur inthe upper and lower Gangetic Plains, and Duthie (Fcdder Grasses of
Northern India, 1888, p. 23), says that it occurs “in the plains of Northern India * * * usually in damp spots.?? The writer hag at present
no personal knowledge of this species in the field, and the only specimens in Herbarium Dehra are from Assam. Specimens and information
regarding this grass will be welcome at Dehra Dun,

[ 82 }

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 83

of abnormal panicles of Saccharum Narenga which owe their peculiarities of a fistular
culm and persistent panicle branches to some injury similar to that described above. This
species therefore cannot be maintained.

It is noticeable that the injurious action of the insect, or fungus, or both, described
above can to some extent be imitated artificially. On a plant growing in the experimental
garden at Dehra Dun one of the flowering culms was sharply bent over and left hanging on
the plant on October 20th, 1909. It was thought that this might interfere to some extent
with the passage of water and food materials to the panicle and thus more or less produce
the results mentioned above. The panicle which was treated in this way did show the
persistent branches and general appearance of those already described but the culm did not
become fistular, probably owing to the central tissue having been fulty developed when the
culm was bent.

Plate XXIII, fig. 2, shows the panicle treated as above described and three other normal
panicles, all four of these having been picked from the same plant on November 20th, 1909.
The panicle from the bent culm clearly shows the persistent branches.

It must be noted that culms are sometimes found in the forest which have been more
or less damaged by insects, or fungi, or both, but which do not show the abnormal characters
noted above, and the extent to which the culm and panicle are affected appears to depend
on the state of development of the culm when it is attacked and on the severity of the attack.
In no case, however, has an abnormal flowering culm, exhibiting the characters described
above, been found which had not been more or less severely damaged, while both abnormal
and normal culms have frequently been collected from the same plant.

It must also be noted that the flowering culms of this species and also those of
Saccharum Munja and Erianthus Ravenne are frequently attacked by larvee which hollow
out the culms and this also often results in the premature drying of the panicle but this
kind of damage obviously differs from that described above in which no obvious excreta
or other signs of larval damage are seen. Hackel divides the present species into two
varieties as below :—

a genuinum Glumes I and II of sessile spikelet obtuse, glabrous dorsally.

I sub-enerved with the exception of the keel nerves.

Sessile spikelet 2-5 to 3 mm. (0:1”’—0-12") long.

Tropical Himalaya.

B Khasianum Glumes I and II of sessile spikelet sub-acute, dorsally hairy below.
I faintly 3 nervid in addition to the keel nerves.

Spikelets larger 3-5 mm. (0:14”) long. Khasi Hills.

The writer finds it impossible to follow this classification. In the first place it must be
noted that in dried specimens the hairs easily break off from the hard coriaceous glumes,
especially in mature spikelets, and leave no obvious scar. It is not therefore always easy
to determine the degree of hairiness of the glumes from herbarium specimens. In all the
fresh flowering spikelets collected by the writer in Dehra Dun, however, there is an obvious

fh sa oy

84. INDIAN FOREST MEMOIRS.

difference in the hairiness of glumes I and II respectively. Thus in the sessile spikelet,
while glume I was more or less densely hairy dorsally, glume II was glabrous, and while
glume II of the pedicelled spikelet was often dorsally hairy it was usually less hairy than
I. At the same time the hairiness of even glume II of sessile spikelet appears to have
little constancy, geographical or otherwise.

In fourteen panicles, for instance, which were kindly sent me by Rai Sahib Kanjilal
trom Sibsagar, Assam, only a single panicle (with spikelets 0-13 in. long) showed the hairy
glume II of Khasianum and on this panicle glume II of sessile spikelet varied from glabrous
to fairly densely hairy dorsally.

So far as herbarium specimens are concerned the local Dehra Dun plant shows glume
I of sessile spikelet and glumes I and II of pedicelled spikelet varying from glabrous to
densely hairy dorsally and a similar variation has been found in panicles received from
Sibsagar. This variation also has been observed on one and the same panicle.

Apart, however, from the inconstancy of the hairiness of the glumes, the other charac-
ters selected by Hackel for the definition of his varieties are not constant and are not
necessarily correlated. Thus, on one and the same panicle, glume I of sessile spikelet has
been found to vary as regards its apex from sub-acute to obtuse, and as regards its nerva-
tion from sub-enerved to clearly 1—38 nerved in addition to the keels. The length of the
sessile spikelet also has varied from 0-1 in. to 0-13 in. on one and the same panicle, while
spikelets 0:13 in. long have shown the hairy glume II of Khasianum.

C.—BIOLOGICAL AND CECoLoGIcCAL NOTES.

Habitat. 60. This is probably the commonest and most
widely distributed savannah grass of Sal forests and is usually the dominant species, at all
events at first, in areas from which the Sal forest has been cleared.

It is found in this locality on well drained loam, usually with a considerable admixture
ef humus. Like Erianthus Ravenne it has considerable power of withstanding shade, but
requires considerably less moisture than that species.

Compared with Saccharum Munja, this species is a much stronger shade-bearer and
slightly less sensitive as regards good soil aeration. It requires for its vigorous develop-
ment a soil with a greater capacity for retaining moisture and not so subject to rapid
drying, as compared with that which suffices for healthy development of S. Munja.

As already noted it is usually found on soils which have been long under forest and
which have a considerable admixture of humus.

In this locality the species is found at its best in small shady blanks in Sal forest, and
it is here a valuable indicator cf soil and moisture conditions suitable for Sal.

Development 61. Perennial, usually gregarious.

of Culms, . -

Season of The plant consists of a much branched rhizome which constantly extends outwards on
Vegetative

Activity and all sides in a centrifugal direction. The aerial culms are biennial and are either the direct
Flowering. [ 84. i

HOLE: SOME INDIAN GRASSES. AND THEIR (ECOLOGY. 85

continuation of the growing points of the rhizome, or they arise as axillary shoots from the
basal nodes of older culms.

The culms are situated close together, the entire plant thus forming a clump or tuft,
the oldest culms being in the interior of the clump while the youngest culms are situated at
the periphery which constantly extends outwards. In some cases the tufts are not clearly
defined and the plant shows a somewhat spreading habit.

This grass occurs in comparatively moist localities but its usual period of vigorous
vegetative activity is about 6 months from June to November, it flowers in October—
November.

At the end of the first season’s growth the culm is found to consist of a short, hard,
mature, basal portion measuring 1—2 inches in length and consisting of 8—12 (usually 10)
very short internodes, which are clearly defined by the insertions of the outer older leaves.

These outer leaves are more or less dead and dry, while within and above them is
situated the still soft immature apical portion of the culm, carrying one or two undeveloped
green leaves, and still younger leaf rudiments, terminating in the apical growing point of
the culm.

Such a culm therefore appears to be merely a tuft of large leaves.

During the second season of vigorous growth the immature apical portion of the culm
develops with great rapidity and produces 5—7 (usually 6) long internodes and finally an
apical segment which terminates in the flowering panicle. Thus the entire length of the
culm, which may attain a height of 16 feet, with the exception of the basal length of some
2 inches, is developed in a single growing season.

It should be noticed that as a rule the 3 or 4 leaves which are first produced at the
extreme base of a young culm are mere scales with a very minute, if any, lamina and,
omitting these, the mature piece of culm usually developed in the first season carries 6
leaves. This again is the number of leaves carried on the 6 long internodes of the flowering
culm, so that we have in each year an apparently constant rate of growth corresponding
to a single leaf for each month of the vegetative season June to November. The culm dies
after producing flowers and seed and as it thus persists through two complete growing
“seasons, it may be described as biennial, but it must be understood that a culm does not
necessarily persist for exactly 2 years.

The young culms may commence their growth at any time during the period of
vegetative activity and a culm which first appears in June 1908 will have completed its
first year in June 1909, and will flower and complete its cycle in October—November 1909,
i.e. in about 14 years. Another culm which originates, say, in September 1908, would
ordinarily flower in October—November 1910, 7.¢. would complete its cycle in about 2
years and 2 months. Again a culm which originates in August 1908, if growing under
favourable circumstances, might flower in November 1909, 7.e. in 16 months. In no case,
however, has a flowering culm been found to be produced in a single growing season and the

f 8 }

Susceptibil-
ity to Fire
Damage.

86 INDIAN FOREST MEMOIRS.

normal rate of growth is, as noted above, 10 short internodes the first season, with 6 long

internodes and the panicle the second season. The season of flowering is also very constant.

Exceptions of course occur and some flowering culms may be found with 20, and some-
times even more, short basal internodes, thus apparently indicating that they have persisted
for at least 3 seasons.

Such cases however appear to be undoubtedly abnormal and are probably due to
exceptionally unfavourable conditions of light, competition by neighbouring shoots,
damage by insects, fungi, and soon. In some of these cases periods of normal development
are found to be succeeded by periods of exceptionally feeble growth and these are almost
certainly caused by damage to the leaves by grazing, fire, or grass-cutting.

62. During the cold and beginning of the hot
season, when the forest grasslands are chiefly exposed to fire damage, the aerial portion of
this species consists of —

(a) The dead flowering culms.

(b) The immature short leafy culms which vary in size from those which have
enjoyed a full season of vegetative activity to the young buds which have only
recently commenced development.

The flowering culms are, as already noted, situated in the interior of the clump and

the young leafy culms at the periphery. In old plants, also, there is usually a more or less.

considerable hollow space in the centre caused by the disappearance of previous generatious
of flowering culms and in fire-protected areas this space is more or less filled with dead
leaves and inflammable débris.

The shoots which are most exposed to damage are the one year culms which carry on
their lower internodes a number of large, more or less dry and inflammable, leaves and
which are situated in the immediate neighbourhood of the dead flowering culms and any

débris which may have collected in the interior of the clump. The youngest culms on the

other hand carry far less inflammable material and are situated on the exterior of the clump,
while the terminal buds of the branches of the rhizome are on the extreme outer periphery,
protected by hard scale-like leaves and as a rule below the growth surface. See Plate XXV,
fig. 15.

The principal cause responsible for mitigating the damage done by fire to this species
in this locality appears to be the fact that it occurs in comparatively moist soil with a
considerable admixture of clay. Owing to its compact nature the soil resists the denuding
action of rain and effectually protects the buds of the rhizome branches and also to a

ereat extent the base of the one year culms, the apex of which is usually situated below the

surface.

An early fire, therefore, as a rule does very little direct damage and is generally not
sufficient to prevent the production of flower and seed, as would be the case if the apical
bud of the one year culms was destroyed. On fire-lines and small grasslands where the

soil is more or less sheltered by the adjacent trees and thus kept moist, and also in large

[ 86 ]

i ee eee

HOLE : SOME INDIAN GRASSES AND THEIR C&COLOGY. 87

grasslands where the soil is well supplied with moisture, the damage done by fire is practi-
cally nil. f

In exposed grasslands, such as Gola Tappar (see Plate XL), however, which are
regularly fired and where the soil is comparatively dry, this species, although flowering
fairly freely, shows inferior dimensions, but this appears to be due rather to the want of
sufficient moisture in the soil for vigorous growth, caused mainly by the destruction of all
humus and organic débris by the fire, rather than to any direct injury by fire. No case has
yet been seen where fire alone has sufficed to exterminate this species.

As a general rule the later the fire occurs in the season and the drier the locality, the
more severe will the damage be, and a fire occurring in an area long protected from fire
will be more injurious than a fire in an area periodically burnt, owing to the accumulation
of débris in the clumps.

63. This plant is generally considered of Utility as
little or no value for fodder and cattle only eat the young and immature leaves. Grass- Bat nae
lands required for grazing therefore should be fired, so that the young leaves may become Poeeee
available for fodder in the hot season. age

On areas which are annually burnt and then grazed the production of flowering culms
is to some extent diminished.

64. As noted above, this species is a valuable Relations of

° : 5 6 ee ‘ Grassland t
indicator of soil and moisture conditions suitable for Sal. Woodlaiids!

D.—Economic USEs.

65. The culms and leaves are used for thatch-
ing, for making tatties (=screens), etc.

Erianthus Ravennae, Beauv. (Plates VIII, XXVIL.)

1a, 18% is Wat IDALS 13L 139. References,
C., Il, 949.
Western Himalaya, Punjab, Upper Gangetic Plain, Sindh. Extends westwards to the Distrivution,

Mediterranean.
A.—DESCRIPTION.

Dolsar, dolu (Dehra Dun). Vernacular

Erect, attains a height of 19 ft. and diameter 0-7 in. Calm

Solid, often slightly fistular just below the panicle. Smooth and polished, striate.

Shortly and finely bearded at the leaf-insertions.

Hirsute with bulbous-based hairs, the latter varying in colour from white to yellow or Leaf-sheath.
-rown.

[ st }

Lamina.

Ligule.

Inflorescence.

Spikelets.

88 INDIAN FOREST MEMOIRS.

The hairs are more or less deciduous and old sheaths are often rough with the persistent:
bulbous bases.

Upper sheaths glabrescent.

Always longer than the proper internode.

Long ciliate on margins towards the apex.

Of uppermost leaf of flowering culm from 8 in. long and 0-25 in. wide, linear and
tapering from base, to 24 ft. long and 0-65 in. wide with greatest width about the middle.

Lower leaves attain a length of 6 ft. and width of 1} in. Are usually 4—5 ft. long and
1 in. wide.

Greatest width about the middle, sometimes in upper third.

Dark green, midrib white.

Apex acuminate, narrowed towards the base.

In basal leaves the concave midrib occupies $ or more of width of lamina at base (often
the entire width of the leaf).

Densely villous above towards the base with bulbous-based hairs, more or less scaberu-
lous along nerves, margins scabrid.

A narrow membranous rim not exceeding 0-07 in. high. Entire and rounded or deeply
2-lobed, patently hairy dorsally with stiff white hairs.

Ciliate.

Spikelets in pairs, one pedicelled and one sessile on the capillary jointed branchlets
of an ample terminal panicle.

Primary rachis sulcate, scabrid on ridges. Primary branches compound sub-
verticillate.

Flowering panicle, 1—3 ft. long, lanceolate, brownish red in colour with spreading
branches. Fruiting panicle oblong, dirty white or brown in colour, branches closely
adpressed to rachis. ,

Distinctly awned, lanceolate acuminate.

0-13 to 0-16 in. long. Exserted part of awn usually about equal to length of spikelet
(from 4 to 14 times the spikelet).

Sessile and pedicelled similar, each one flowered and ¢.

Pedicelled spikelet at length falling from the pedicel, sessile falling with the attached
pedicel and joint of axis. Joints and pedicels villous with long pale purplish or brownish
airs. Hairs of joint overtop joint by about twice the length of the joint.

Joints 1 to 2 sessile spikelet. Pedicels + to $ sessile spikelet. Majority of pedicels.
longer than proper joint.

Callus hairs shorter than to subequal the spikelet, purplish or brownish in colour

i 88h

HOLE: SOME, INDIAN GRASSES AND THEIR GCOLOGY. 89

Sessile Spikelet.
Glume I.—Lanceolate, 2 lateral keels, dorsally flat or depressed between the keels.

Apex 2-mucronulate.

One or both margins incurved.

Dorsally scabrid on keels, otherwise glabrous, or more or less villous
dorsally.

Villi not overtopping the glume, or overtopping by less than 4 glume’s
length.

Two nerved, sometimes with 1—2 additional faint nerves between the
keels.

Glume IIT.—Subequal to I. A central keel.

Mucronate.

Margin incurved ciliate.

Dorsally scabrid on keel, otherwise glabrous or more or less villous
dorsally.

Villi not overtopping the glume, or overtopping by less than } length of
glume.

One nerved and sometimes one or two partial lateral nerves.

Glume III.—Slightly shorter than IT.

Oblong-lanceolate, hyaline. Apex mucronate or acute.

Glabrous dorsally.

Margin incurved, ciliate above.

One—three nerved.

Glume IV —Usually 3 III.
Ovate-lanceolate, hyaline.
Margin incurved, ciliate.
Long-awned. Awn 0:10 to 0-23 in. long.
Three nerved.
Pale—About 2 IV, ovate-lanceolate, hyaline, glabrous, enerved.
Lodicules, two, cuneate, glabrous.
Anthers, three, -05 to -07 in. long (0-08 to 0:10 in. fide Hackel) yellow, streaked with
purple.
Stigmas -03 to ‘06 in. long. Yellow.

Pedicelled spikelet is similar except that glumes I and II are often strongly three

nerved and both glumes I and II are more densely villous dorsally. Villi not overtopping,
or overtopping by less than 4 spikelet’s length.

guished by its distinctly awned spikelets, the broader dark green leaves and hairy leaf-

This species is not infrequently mistaken for Saccharum Munja, but is easily distin- Field Charac
ters,

sheaths. The fruiting panicle also is usually browner in colour.

[ee

90 INDIAN FOREST MEMOIRS.
B.—TAXoNomy.

66. Hackel separates the common Mediter-
ranean form of this plant as a sub-variety (genwinus) with larger spikelets (0-2 to 0-25 in.
long) and with glume IV usually glabrous.

The Indian plant is referred to sub-variety purpurascens.

C.—BIoOLOGICAL AND (EcoLocicaL Notes.

Habitat. 67. This plant, while undoubtedly requiring
a liberal supply of available moisture for vigorous development, does not extend into the
water-logged soil of the marsh. |

It is thus apparently more sensitive as regards soil aeration than Imperata, while it
requires far more moisture than either S. Munja or S. Narenga.

It is found on both sandy and clayey loam. Its broad, very thin, dark green leaves are
markedly mesophytic, if not indeed hygrophytic in their characteristics, but the plant does
not attain the dimensions of the truly hygrophilous species S. arundinaceum.

Warming’ states that this species sometimes enters into the composition of reed-
swamp in Mediterranean countries, accompanied by Phragmites and Arundo Donaz.

The plant is a strong shade-bearer, but its capacity in this respect is often masked by
the fact that the roots of neighbouring trees and shrubs frequently cause the available
moisture supply to fall below that necessary for its vigorous development.

In localities where ample water is available by percolation or otherwise, this plant is
seen to be a good shade-bearer.

In this locality it is found at its best on the banks of irrigation channels and water-
courses, where ample water is available by percolation and capillarity, but where the soil
in contact with its roots is not actually water-logged except for short periods. It usually
indicates a soil with the permanent water-table near the surface.

Development 68. The remarks given under this head for
of Culms,

Season of S$. Munja above apply, generally, also to this species. The culms are biennial. The usual
Vegetative i . 0.6 . + : °
Activity and period of vegetative activity is from February to November. Flowering takes place in
~ October—November, although panicles are occasionally met with also in December—
January, thus indicating that activity may in some cases continue practically throughout
the year.
During the first season’s growth from 1—2 inches of culm is produced with an average
of 14 short internodes and carrying 10 green leaves.
During the second season the long internodes (usually 10) of the flowering culm, carry-
ing 10 leaves and terminating in the panicle, are produced. Unlike the Saccharum, how-

ever, the leaves of this species are not cut for thatching purposes.

1 Kcology of Plants, 1909, p. 189.

205i

HOLE: SOME INDIAN GRASSES AND THEIR CECOLOGY. 91

69. The damage done by fire is similar to that Suscentibil-
described above for S. Munja and S. Narenga, but the species suffers much less than S. damage.
Munja owing to the fact that it usually occurs on loam. Growth during February also is as
a rule not so vigorous as in S. Munja and the growing points of the young clums are usually
situated close to the surface of the ground at the end of that month.

70. The leaves of this species are eaten by Utility as

. : : Fodder and

buffaloes to some extent, especially when young. Grasslands required for grazing should vest Treat-
c : ment for

be fired so that the young leaves may become available in the hot season. Fodder Pro-
duction.

71. On areas where this grass is dominant the Relations
: : O51,0 ° ° : of Grassland

soil and moisture conditions are as a rule suitable for moist mixed forest. Such representa- to Woodland.
tive species as Cedrela Toona and Mallotus philippinensis have been seen to be able to
naturally establish themselves in areas occupied by this grass without artificial assistance.

The large tufted species of Saccharum, such as S. Munja and S. Narenga, have con-
siderable power of improving the soil by the addition of humus.

Not only do the decaying culms and leaves of these plants themselves add considerably
to the humus content of the soil, but the central hollow of the clumps, surrounded | as it is
by a barricade of culms, acts as a veritable humus-trap in which the leaves and débris from
neighbouring trees and plants is caught, stored, allowed to gradually decay and find its way
into the soil.

Erianthus Ravenne, being of more robust growth than either of the two species named,
possesses this power to an exceptional degree and the writer has removed 11 handfuls of
decaying leaves and débris from the centre of a single clump.

D.—Economic USEs.

72. The leaves of this species quickly decay
and are not therefore used for thatching, a point of interest in connection with the
hygrophytic character of the species.

The culms are used for making screens, etc.

Imperata arundinacea, Cyrill. (Plates XXVIII to XXX.)

iP, 18. IL, WAL, 10s 181, CP

P., If, 1188; C., 1, 946.

The hotter parts of India and ascending in the Himalayas to 6,500 ft. and possibly Distribution.
higher.

Extending to the Mediterranean region, Africa, Java, Japan, China, Australia.

References.

A.—DESCRIPTION.
Siru, sirhu (Dehra Dun).

Vernacular

[ 91 1 Name.

culm.

Leaf-sheath.

Lamina.

Ligule.

Inflorescence.

Spikelets.

92 INDIAN FOREST MEMOIRS.

Erect, simple, slender, from 5 in. in height and almost filiform, in poorly nourished
individuals, to 91 ft. high and 0-3 in. dia.

Solid, slightly fistular at base, smooth.

Leaf insertions tumid, glabrous, or densely bearded with erect white hairs.

Glabrous or glabrescent, ciliate or glabrous along the margin towards the apex.

Usually longer than proper internode.

Of uppermost leaf of flowering culm from mucroniform and 0-05 in. long to 6 in. long
and 0:25 in. wide with greatest width in middle.

Of lower leaves erect or arcuate and attains a length of 4 ft. and width of 1-1 in.

Greatest width about the middle.

Dark green, midrib white.

Apex acuminate, narrowed towards the base where the midrib occupies almost the
entire width of the leaf.

Smooth, but scabrid on margin and on one or more sub-marginal nerves above, especi-
ally towards the apex.

White villous above on margins towards the base and behind the ligule.

Rounded, truncate or two lobed. Attaining a height of 0:13 in. Ciliate. Dorsally
silky.

Spikelets in pairs, both pedicelled (one pedicel long and one short), on the simple, or
compound, subverticillate, capillary branches of a narrow, spiciform, terminal, panicle.
The upper spikelets in the racemes are sometimes solitary.

Branches are tenacious and not jointed.

Primary rachis sulcate, glabrous or scaberulous, sparsely villous at nodes with long
white hairs.

Pedicels glabrous, scaberulous or pubescent, and with a few long white hairs towards
vhe base.

Panicle from a little over 1 in. (in depauperate individuals) to 20 in. in length, not
exceeding 1 in. in width.

Flowering panicle purple with the exserted stigmas, the callus-hairs being dose
adpressed to the axis, fruiting panicle silvery white with the wide-spreading callus-hairs.

Not awned, lanceolate, 0-13 to 0-15 in. long (attaining 0-2 in. fide Hackel).

Both spikelets of each pair similar, each one-flowered and ¢, and at length falling from
the pedicel.

Callus-hairs soft, white, 2—8 times as long as spikelet.

Glume I.—-Lanceolate, membranous, slightly thickened towards the hase, apex
hyaline, 3—9 nerved, none of the nerves extending to apex of
glume, margins incurved, ciliate above.

Dorsally villous with soft white hairs overtopping the glume by 14 to 3
times the length of the glume.
L |

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 93

Glume II.—Similar and subequal to I, but sometimes sub-keeled with mid-nerve
extending almost to apex.

Glume III.—Oblong, hyaline, enerved, apex acute or sub-truncate and laciniate or
denticulate, ciliate.
= +to# Il.

Glume IV.—Subequal to 3 III, ovate-lanceolate, hyaline, enerved, apex acuminate,
acute, or obtuse and laciniate or denticulate, minutely ciliate or
glabrous.

Pale. —-Quaarate, rectangular, or sub-pentagonal, hyaline, enerved, apex denticulate
or unequally laciniate, glabrous or ciliate.

4 to subequal IV.

Lodicules.—None, or very minute.
Anthers.—Two, 0:12 in. long, yellow. The filaments are sometimes connate below.

Stigmas.—Two, 0-12 in. long, purple.

The ordinary savannah form of this plant is about 3 ft. high and is strongly gregarious. Fiela Charace
When in leaf it is somewhat liable to be mistaken for Eragrostis cynosuroides (known in “*™
Dehra Dun by the vernacular name of dab) with which it is frequently associated and
which has a very similar habit.

The leaves of dab however are broader at the base and glaucous, not dark green like
those of sirhu. The old leaves of siriu also turn reddish or brownish yellow, those of dal

are much paler.

The somewhat succulent white stolons of sirhu, also, covered with soft membranous
sheaths, are at once distinguished from the hard, polished, yellow stolons of dab, which are
‘covered with hard shining sheaths.

73. The leaf of the ordinary savannah form Anatomy.
of this species exhibits considerable xerophilous adaptations.

The cuticle is strongly developed, particularly in the lower epidermis.

Each longitudinal nerve is provided with two girder-like groups of sclerenchymatous
fibres. The girders of the principal nerves are strongly developed and are situated, respect-
‘ively, close below the upper and above the lower epidermis, the nerve thus being connected
with the epidermis above and below. Each nerve is flanked laterally with a narrow band
-of chlorenchyma, these bands curving inwards and almost meeting around the nerve inside
the girders. The chlorenchyma thus gives the appearance of more or less complete rings
around the vascular strands. The epidermis situated immediately above the chlorenchyina
is provided with stomata which are particularly numerous on the upper surface of the leaf.
_Alternating with, and parallel to, the principal nerves on the upper surface of the leaf are
longitudinal bands of the remarkable, large, thin-walled, colourless, cells which are called

Pees |i

94, INDIAN FOREST MEMOIRS.

motor-cells by some writers’ and bulliform-cells by others.” Beneath these motor-cells, and
therefore alternating with the principal longitudinal nerves, are smaller nerves, also
provided with girders and chlorenchyma rings, similar to those of the principal nerves, but
smaller. Immediately above the midrib is a thick, cushion-like, band of thin-walled colour-
less cells which is believed to function mainly as water-storing tissue. It is to be noticed.
that this tissue is protected by a layer of sclerenchyma, one to two cells thick, which lies
immediately beneath the upper epidermis, while the bands of motor-cells are provided with.
no such protection.

When the leaf dries, the blade becomes convolutely folded inwards towards the upper
suriace. There seems little doubt that this folding is effected mainly by the shrinking of
the motor cells and in a section which has been allowed to dry and fold under the micro-
scope, the leaf tissue on each side of a band of motor cells can be distinctly seen to have been.
pulled inwards towards the motor cells. This convolute folding of the blade obviously re-
sults in the protection of the numerous stomata on the upper surface of the leaf and must
therefore considerably reduce transpiration, while the exposed lower surface is protected.
by the strongly developed cuticle.

In the marsh form of this plant, the stomata are chiefly on the lower surface of the
leaf, thus indicating less necessity for diminishing transpiration. The fresh leaves are
thicker than those of the ordinary savannah form and have a somewhat fleshy, almost.
spongy, texture. It is remarkable that this is caused mainly by an increase in the size and
number of the water conducting vessels of the xylem, annular, pitted and reticulately
thickened vessels, with large lumina, being noticeable in the nerves, while there is no.
marked development of special aerating arrangements, such as large intercellular spaces.

The stolons of the ordinary savannah form show in cross-section, commencing at the
exterior, (1) Epidermis with thick cuticle, (2) One to two layers of sub-epidermal scleren-
chymatous fibres, (3) A thick sheath of parenchyma, the walls of which turn purple with
chlor-zine-iodine, (4) A small central woody cylinder containing the vascular bundles. This:
cylinder is protected by an outer layer of thick sclerenchyma, while the bundles are also more
or less completely surrounded by sclerenchyma. The walls of the tissue of this inner
eylinder all turn dark yellow with chlor-zinc-iodine, with the exception of those of small
areas of parenchyma found chiefly in the centre, which turn purple.’ The stolons of the
swamp form show an essentially similar structure with the remarkable exception that large
air spaces are found in the outer parenchymatous sheath, thus indicating the necessity for
special aeration-devices in the subterranean stolons of the swamp form to enable the roots,
growing in poorly-aerated water-logged soil, to keep pace with the increased transpiration
caused by the enlarged leaf-surface and by the diminished protection of the stomata.

1 See Grasses by H. Marshall Ward, 1901, pp. 25, 62.

2 See Brandis in Trans. Linn. Soc. London, Vol. VU, p. 78.

_ 3 The soft outer parenchymatous sheath (3) is often destroyed by vermin and stolons are often seen in which this sheath has been
entirely eaten away, leaving only the central woody cylinder (4) intact.

[ oe

HOLE : SOME INDIAN GRASSES AND THEIR (ECOLOGY. 95

B—Taxonomy.
74. According to published descriptions,
Imperata exaltata, Brongn, the area of distribution of which, to some extent, overlaps
that of the present species, is distinguished from the latter mainly by the following

-characteristics :—
(1) Panicle thyrsiform, 4—5 inches broad at base (F. B. I., VII, 107).

(2) Stamen one.

(3) Upper spikelets in racemes solitary.

(4) Culm fistular.

(5) Glumes I and II three nerved with mid-nerve reaching, or almost reaching, the
apex of glume.

It is therefore interesting to note that, in the local specimens of Imperata arundinacea,
Cyrill, the culms are slightly fistular at the base, glumes I and II have 3—6 nerves and the
upper spikelets in the racemes are sometimes solitary.

70. Of Imperata arundinacea three forms can
be more or less clearly distinguished locally as under :—

(a) The depauperate form common on lawns or areas where the grass is continually
cut or grazed, with minute, almost filiform, culms and small leaves. Leaf-
insertions usually long-bearded. Glume IV and pale usually glabrous.

(b) The ordinary savannah form which usually attains a height of about 3 ft. with
leaves up to 0-7 in. wide. Leaf-insertions bearded, or glabrous. Pale and
glume IV ciliate.

(c) A robust form found in swamps or marshy soil where there is an abundance of
available mnoisture more or less throughout the year. This plant attains a
height of 91 ft. and probably more. Leaves up to 1-1 in. wide, leaf-insertions
glabrous. Pale and glume IV ciliate. (This plant is var. latifolia Hook f.)

The mode of occurrence of these different forms in their natural habitats and the fact
that numerous intermediates are met with, as the conditions vary, indicate that such
characters as the vigour of growth (indicated by the diameter and height of the culms,
length of the panicles, length and width of the leaves), the hairiness of the nodes, of glume
IV and of the pale, are directly dependent on the nutrition of the plant and it is believed
that form (a) owes its characteristics to the starving effect of the artificial interference
with its normal growth and development, while the peculiarities of form (c) are due to the
exceptionally favourable conditions of available moisture under which it grows.

Plants of all these forms are now being cultivated at Dehra Dun with the object of
testing the effect of protectior on (a) and of a diminished water supply on (c).

Hackel has subdivided Imperata arundinacea into three varieties of which he further
distinguishes sub-varieties, depending mainly on such characters as the hairiness of the leaf-
insertions, width of the leaves and height of the ligule. The height and shape of the ligule.
however, is more or less correlated with the width of the lamina, while the other characters
appear to vary with the locality as indicated above and do not define forms of any constancy.

ENS i

Habitat.

Development
of Culms,
Season of
Vegetative
Activity aud
Flowering.

96 INDIAN FOREST MEMOIRS.

C.—BIOLOGICAL AND CEcoLtogicaL Notes

76. This plant is here chiefly found in the:
same localities as Saccharum Munja and Erianthus Ravenne, both of which are stronger
and more vigorous species and its distribution appears to depend mainly on two facts :—

(1) It is not sensitive as regards soil aeration and can thrive on stiff clay, while its
maximum dimensions are attained (by the swamp form) in the perennially
saturated water-logged soil of the marsh.

(2) The ordinary savannah form requires far less moisture for its vigorous develop-
ment than does Hvianthus Ravenne.

{ft 1s thus able to occupy stations into which its stronger competitors cannot penetrate.
It occurs on sandy soil with Saccharum Munja, but is most common on heavy soil with a
more or less considerable admixture of clay and in the latter case its most frequent associate
is Eragrostis cynosuroides.

The species has considerable power of withstanding shade.

A perennial and usually markedly gregarious species.

The aerial culms spring from somewhat succulent, white, cylindrical, subterranean
stolons. These stolons are at first covered and protected by membranous sheaths (= leaves)
which eventually fall off, their circular insertions then clearly defining the internodes of
the smooth white stolon.

The culm may be the direct continuation of the growing point of the stolon, or it may
arise as an axillary shoot from a node of the stolon, or of the basal portion of an aerial
culm.

When a stolon is about to produce an aerial culm, its apex turns towards the surface
and a few short internodes are developed from the nodes of which new stolons are sent out,
the function of which is to provide for the continued growth and extension of the plant
below ground.

The production of the true aerial culm is then commenced, a number of short internodes.
being first formed, then a few long internodes and finally the flowering panicle. The leaves.
produced on the first internodes are mere scales with no green lamina, on the higher inter-
nodes the green lamina progressively increases to its maximum and then again rapidly
decreases in size in the leaves of the long uppermost internodes.

In many cases the short internodes of the stolon proper are more or less sharply defined,
but in others the change from stolon to culm is imperceptible and it becomes impossible to:
say whether a particular internode is to be considered a part of the stolon, or of the aerial
culm.

Taking the ordinary savaynah form of this grass, however, an average mature culm
usually shows a basal portion of about an inch in length, consisting of 14 short internodes
(including the short internodes of the stolon where these can be distinguished. The exact

L 25

HOLE; SOME INDIAN GRASSES AND THEIR GCOLOGY. 97

number may vary from 8—18), then four long internodes and finally the apical length which
terminates in the panicle.

The leaves on the basal nine internodes are usually scales with no green lamina, so
that the average culm produces nine leaves, with a more or less perfectly developed green
lamina.

The average period of vigorous vegetative activity is believed to extend from February
to October and the majority of the young culms appear to commence their growth at the
beginning of the rains in June.

During November——-January there is, as a rule, a well-marked period of rest and if an
average culm is examined during this period it will be found to consist of a short hard
mature basal portion oz about 18 short internodes, the upper four or five of which carry
more or less dead and witheréd large leaves.

Within and above these are one or two not yet fully developed green leaves and finally
in the centre at the apex of the culm is the terminal growing point situated close above the
upper node of the mature basal portion.

During the season November—January, therefore, the local grasslands of this species
show a multitude of short immature culms each of which looks like a tuft of leaves.

The blades of the majority of the leaves of these culms are mature, spreading, fully
expanded, and reddish, or brownish, yellow in colour, while only one or two interior and
apical leaves are still immature, with their blades convolutely folded and green in colour.
This is the cause of the characteristic reddish brown colour of the grasslands in the cold
season where this species is dominant. Vigorous vegetative activity is usually resumed
the following February and the flowering panicles are produced in May—June.*

As the aerial culm dies after flowering, its average duration is about 12 months and it
may be described as annual. |

It will be noticed that the nuraber of green leaves produced on the culm, viz. 9, corre-
sponds to the number of months’ vigorous vegetative activity, viz. June—October in the first
year and February to May in the following year, while the culm completes its cycle of
development in nine months of vigorous growth.

The above represents what is believed to be the average development of the savannah
form in the forest, but individual culms of course vary to some extent in the period of their
life cycle and in a plant of this form which was cultivated at Dehra Dun, on moist soil,
where activity was not checked at the close of the rains, a culm, regarding which dates
were recorded, actually completed its cycle in seven months from June—December.

In the dwarf form characteristic of areas regularly grazed or mown, and also in the
marsh form, the number of green leaves produced on the culm is usually greater, i.e. from
10—12 which is perhaps the result of vegetative activity being possible during a greater
number of months in the year.

1 In areas where the grass has been cut, grazed, or burnt, vegetative activity commences earlier than usual and flowering usually
takes place in March—April. The swamp-form of this species also usually flowers at the close of the raius, or in the cold se:son.

[ 97 }

98 INDIAN FOREST MEMOIRS.

Another irregularity of growth which has been particularly noticed in the case of the
marsh form is that a length of long stolen-internodes frequently succeeds a length of very
short internodes, which as a rule immediately precedes the aerial culm.

This is believed to be caused by an alteration in the ground-surface level and to the
growing points of young culms being covered with fresh deposits of soil and water-brought
deébris.

The marsh form, also, instead of having only four long internodes below the panicle,
usually produces 7 and the immature culms often show a height of 2 to 3 feet and probably
more, instead of about 1 inch, at the end of the first rainy season.

This form, however, is comparatively rare, of little value as fodder and, owing to the
conditions of its habitat, little subject to fire damage, and in the following notes it is left
out of consideration.

It must be noted that young culms may commence their development at any time during
the period of vigorous vegetative activity and culms which commence their growth towards
the end of the rains, or exceptionally early in the following hot weather, may complete
their cycle and flower in September—October. In this way the sporadic flowering in the
cold season which is characteristic of the species is probably brought about.

On lawns, or grazed areas, where vegetative activity is in progress for about 10 months
in the year, the time of flowering is also irregular for the same reason.

The course of development indicated above, however, which results in gregarious flower-
ing towards the close of the hot season is believed to be the rule, at all events in Upper
India, and Roxburgh nctes “ particularly common over Bengal, where the fields are white
with it when in flower, after the first rains in April and May.”’

Susceptibil-

Geen 77. During the cold and beginning of the hot
Damage. season when the sirhu areas are chiefly exposed to fire damage, the aerial portion of the

plant consists mainly of the immature short leafy culms, with a few flowering culms and a
few young culms which are just commencing their aerial development. The dead and dry
aerial portion of the flowering culms is of course liable to be burnt. The growing points of
the stolons and of the youngest culms are as a rule situated below the ground surface and
are therefore not liable to injury.

The most extensive damage is liable to occur in the case of the immature leafy culms
which commenced growth at the beginning of the preceding rains and which in ordinary
course would produce flowering panicles towards the close of the hot season. The mature
leaves of these culms are, as already noted, more or less dead and dry and the burning of this
inflammable material may result in the death of the growing apex of the culm and the non-
production of a flowering panicle. In this way fires may diminish and interfere with the
usual gregarious flowering in April—May. The extent of the damage done depends largely

1 Ley p. 284,

fr 98 |

HOLE: SOME INDIAN GRASSES AND THEIR GiCOLOGY. 99

on the quantity of dead dry leaves on the culm and this to some extent depends on the mois-
ture in the soil; where sufficient moisture is available, vegetative activity is prolonged and
the leaves persist and remain green well into December, while in drier localities they dry
off in November. The period of activity also to some extent depends on the nature of the
soil, as does the damage done by fire. The plant occurs both on sandy and loamy soil, the
former dries more rapidly than the latter which results in shortening the period of vegeta-
tive activity and increasing the number of dead inflammable leaves, while at the same time
there is more danger of the fire penetrating below the ground surface. Thus, as a general
rule, the drier the locality and the later the fire occurs in the season, the more serious is the
damage. An early fire frequently does very little damage and prolific flowering may often
be seen in May on areas burnt early in the season.
The following facts all tend to reduce damage done by fire to this species :—
(1) It is usually found on clayey soil which retains moisture tenaciously and conducts
heat badly.
(2) The stolons and their growing points from which the aerial culms arise, are
usually situated at a considerable depth below the ground surface.

(3) During the seasor when the grasslands are usually burnt, there are a very few,
old, dry and inflammable, flowering culms.

(4) The aerial culms are scattered and at considerable distances apart, instead of
being aggregated in tufts, so that the quantity of inflammable material and
consequently the intensity of the fire at any one point is small.

(5) One or more buds are usually developed from the basal nodes of the aerial culms,
or from the nodes of the stolon below it, which as a rule escape being damaged
by fire. These develop into aerial culms and are thus able to replace any culm
which is destroyed.

On the whole, therefore, the damage done by fires to this plant is slight and fire-damage
alone is not likely to be sufficiently severe to cause the destruction of the species.

78. The young shoots and leaves of this plant vtitity as
undoubtedly provide good fodder, but the mature leaves of a normally developed culm are het trate,
too coarse to be relished by cattle. If sirhu areas which have been allowed to develop with- radieeee
out interference during the rains are fired in the cold season, or beginning of the hot *t’™
weather, a number of young leafy aerial culms soon make their appearance, while the des-
truction of the oid dry leaves exposes clearly to view the young inner green leaves of the
older culms as they develop from the uninjured apical buds. These young shoots are
usually appreciated by cattle. It must be noted that when the upper green portion of a
young culm is bitten off by a grazing animal the result is not to damage the low-seated apex
of the culm itself, but only to more or less completely destroy the upper portion of the
leaves, the outer, more fully developed leaves being most injured, their basal portions often
alone escaping, while the inner and upper immature leaves are progressively less and less

[ 99 J

Relations of
Grassland to
Woodland.

Miscellane-
ous

100 INDIAN FOREST MEMOIRS.

damaged. The naturai result of this injury is that only the interior and upper smaller
leaves are able to develop, while owing to the more or less complete destruction of the outer
and lower leaves, the cuim 1s poorly nourished and shows exceptionally feeble development,
both as regards the length of the internodes and diameter of the culm. Grazing therefore
tends to produce a depauperate form with unusually small green leaves situated close to the
eround surface, the mature flowering culms of which are minute, often filiform and a few
inches high, with a panicle sometimes not exceeding 1 inch long. Grass-cutting (on a lawn
for instance) obviously has precisely the same effect, the interference with the plant’s develop-
ment being more severe, the ewer and nearer the ground the shoots are cut or bitten off.

Owing to the period of vegetative activity being prolonged in areas which are grazed,
or where the grass is cut, and to the fact that the subterranean stolons during the period of
vegetative activity continue 1c produce fresh young culms, this species is capable of produc-
ing a practically perennial supply of palatable young shoots and small leaves which consti-
tute good fodder, and incidentally it may be noted that, for a similar reason, it is capable
of forming a fairly good green sward, or lawn.

As regards the best treatment for pastures of this grass it is obvious that the growth
on those areas where the plants have been allowed to develop undisturbed through the
rains is too coarse to be of ary value and this ought to be fired if good fodder is desired.

Duthie writes as follows regarding this grass: “ Cattle relish it when young.” * *
“Tn Australia it is called ‘blady grass’ and the young succulent foliage which springs up
after the occurrence of a fire is much relished by stock. I have observed the same effect
resulting from periodicai fires on certain parts of the Himalaya where this grass is plenti-
UN aaa

“ It forms the chief constituent of the turf in the Botanical Gardens at Calcutta.”

79. This species is not an injurious competi-
tor of the Sal tree and in this locality it is only dominant on limited areas which are as a
rule unsuitable for the growth and development of Sal seedlings on account of the soil, in-
sufficient aeration, a deficiency of available moisture at certain critical times of year, frost
or other causes. Its importance as regards the forest therefore consists mainly in the
danger of fire resulting from the masses of inflammable dry leaves existing in the areas
occupied by it during the fire season. This danger is best dealt with, in the usual way, by
firing the grass under suitable control. It should be noted that if areas which have thus
been burnt could be well grazed throughout the rains, the result would be to produce a non-
inflammable sward which, if continually grazed, would require no further firing. In
localities where this grass is plentiful, or could be cheaply introduced, treating fire-lines
and blanks in this way might considerably reduce the cost of fire protection.

80. Like kans this species is liable to take
possession of agricultural lands which have lain fallow and is then exceedingly difficult to

' Fodder Grasses of Northern India by J. F. Duthie, p. 23.
2 Listof the Grasses of North-West2rn India (1883) by J. F. Duthie, p. 14,

[ 100 }

HOLE: SOME INDIAN GRASSES AND THEIR @COLOGY. 101

eradicate, as might be expected, on account of its deep-going’ much branched stolons.
Hackel (p. 95) notes that in the East India Islands it occupies vast areas of fallow lands,
+o the exclusion of almost all other plants. It is there called alang-alang.

In India the succulent white stolons are eaten by pigs and areas which have been well-
worked by pigs in their search for the stolons are not infrequently seen in the forest. It
is possible that in some cases the eradication of the species might be cheaply accomplished
hy the aid of pigs.

Schimper,’ quoting Junghuhn, speaks of this grass as follows in the Malayan Archi-
pelago :—

“When the soil remains uncultivated after clearing the forest, as a rule the social
and dense-growing alang-grass first replaces the vanished forests, then areas
extending for miles, even indeed for whole days’ journeys, are transformed
into a uniform wilderness of dense grass three to five feet high, while on
mountain-slopes the same grass extends far beyond its original zone, and
spreading over everything it ranges up to altitudes of 6,000—7,000 feet, being
almost insensible to differences of temperature. Its silken haired seeds, light
as the tenderest down, are wafted away in millions by the slightest breath of
wind and greatly facilitate its general distribution, while its creeping and
deeply penetrating roots increase the difficulty of eradication when once this
grass, so tenacious of life, has established itself. I have reasons for believing
that while the land was in its original condition the alang-grass was restrict-
ed to sundry sterile, arid, waterless tracts of the hot zone, and was chiefly con-
fined to heavy, hard, easily dried clay soil, with an iron-pan, but that at the
present time, wherever we meet this grass on a fertile light soil and on
mountain-slopes at above 2,000 feet, this state of affairs is first brought about
by the hand of man. . . In Northern Sumatra, especially in the Batta
country that has been devastated by war, grassy wastes have consequently
come into existence which cover everything far and wide with a hideous

uniformity and overrun plain, mountain, and valiey with their whitish-green
mantle.”

D.—Economic USEs.

81. In addition to its value for fodder this
grass under the name of “ lalang ” has recently been favourably reported on as a paper-
making material :

“The ultimate fibre obtained from this grass is very similar in most respects to
Esparto; the yield of bleached fibre being about the same. This is a favourable indication
inasmuch as ‘ Esparto’ is one of the best known and most useful sources of supply to the

1 Plant-Geography, 1903, p. 162.
EO

102 INDIAN FOREST MEMOIRS.

trade * *. The results obtained from the chemical analysis show that the grass is capable
of yielding a good quality of cellulose, suitable in every way for the manufacture of
paper.”*

The value of the leaf for paper manufacture probably depends mainly on the existence
of the well developed groups of sclerenchymatous fibres which form the girders of the
vascular bundles.

The leaves are largely used for thatching.

Triraphis madagascariensis, Hook f. ex Prain in Bengal Plants, II, 1219.
(Plates IV and XXXI to XXXIII.)

References. F. B.I., VII, 305. (Under Neyraudia madagascariensis.)
IP, JOE, IAG),
Distribution. Tropical Himalaya and Sub-Himalayan tract from Punjab to Assam, Burma.

Extends to China, Java, Africa and Madagascar.

A.— DESCRIPTION.

Vernacular Bansi (Dehra Dun).
Name. Erect, slender, attains a height of 10 ft. (Prain gives 15 ft.) and diameter of 0-25 in.
Culm. (F. B. I. gives 0-5 in.).

Usually solid above and fistular below. Sometimes solid throughout.

Glabrous, striate, green in colour, pruinose when young, polished when old. Secondary
culms are often produced from the upper nodes.

These are frequently poorly nourished and may show a diameter of only 0-05 in.

Leaf-sheath. Glabrous.
Lamina. Of uppermost leaf of flowering culm usually well-developed and attaining a length of
2 ft. and width of 0-25 in.

In depauperate culms it may be only 4 in. long with a width of 0-05 in.

Of lower leaves attains a length of 24 ft. and width of 0-5 in. (F. B. I. gives width of
iL) ial.)

Linear, long acuminate, slightly narrowed near the sub-amplexicaul base. Greatest
width attained within the basal eighth, or within 3 inches from base. Usually jointed to,
and deciduous from, the sheath.

Scaberulous on margins aud also on the nerves on both surfaces, especially towards the
apex (F. B. I. gives smooth). A few long hairs on margins and inside at base. :

Ligute. A villous ridge.
Inflorescence. Spikelets solitary, pedicelled, on the fascicled or whorled branches of a terminal panicle
Primary rachis glabrous or scaberulous. Branches and pedicels scaberulous.

1 Agricultural Bulletin of the Straits & F. M. States, Vol. VII, No. 12, December 1908, p. 586.
r 102 J

HOLE: SOME INDIAN GRASSES AND THEIR (ECOLOGY. 103

Panicle attaining a length of 3 ft. but in depauperate specimens only 6 inches long.
Branches of flowering panicle spreading, arcuate, of fruiting panicle, erect, or ascending,
and adpressed to rachis.

0:25 to 0°45 in. long, including the awn, 3—9 flowered, with two empty basal glumes. Spikelets.

(In addition to the empty glumes the lowest flowering glume, III, is sometimes epaleate,
neuter and glabrous, but this has not been found in the local plant.)

Glume I.—Empty, lanceolate, 1 nerved, mucronulate, scaberulous dorsally on
nerve. Membranous, brown in colour.

Glume II.—Empty, usually a little longer than I, oblong-lanceolate, mucronulate,
1 nerved, scaberulous or smooth dorsally on nerve. Membranous,
brown in colour.

Flowering glume (Glume III)—Longer than the empty glumes, broad subulate, 3
nerved, with two lateral keels. Apex slender with two setaceous
points and an interposed scaberulous awn. Awn about 4 length
of glume and recurved in fruiting spikelets, but not twisted.
Dorsally villous with long white hairs along the keels. The
remaining flowering glumes are similar.

Pale.—% to 2 flowering glume, oblong, 2 nerved, with two lateral keels, hyaline,
minutely scaberulous dorsally along the keels, glabrous.

Apex truncate, minutely two dentate.

Lodicules—Minute, two, cuneate or obovate, glabrous.
Anthers—Three, 0°07 in. long, yellow.
Stigmas.—Two, 0-03 in. long, hardly exserted from spikelet.

The rachilla, which is continued beyond the upper flowering glume, is jointed at the
base above the empty glumes and between the flowering glumes. The portion of the rachilla
above the joint and below the next flowering glume (=the callus) is densely bearded with
short white hairs.

The fruiting spikelet breaks up at the joints of the rachilla, from the apex downwards,
the basal empty glumes persisting longest.

B.—TAxonomy.

82. In F. B. I. two forms of the species are

given :—
(1) madagascariensis proper; neuter glume O. Western Himalaya, Burma, Africa,
Madagascar ;
(2) Zollingert; neuter glume present. Eastern Himalaya, Assam, Bengal, Burma,
Java;

foie 9

Habitat,

104 INDIAN FOREST MEMOIRS.

but the author writes : “ I can find no difference whatsoever between the specimens with and
those without the empty glume IIT.” All specimens of the local plant examined have no
neuter glume and fall under (1) above.

The description under (A) above which (with the exception of the characters in paren-
theses) refers exclusively to the local plant (i.e. to the form madagascariensis), growing
under varying conditions of soil-moisture, shows that this is a decidedly less robust plant
than the species as understood by Prain and in the F. B. I. which also includes the form
Zollingeri (=Arundo Reynaudiana Kunth).

The latter also has a different area of distribution and although starved specimens of
both these plants exist which, especially in herbaria, are likely to mask the differences
between them, it seems probable that they will prove to be distinct species. The writer,
however, has at present no knowledge of the E. Himalayan plant in the field and pending
the results of further study it seems advisable to deal with these forms as varieties as
follows :—

(1) madagascariensis proper: Glume IIT similar to IV and fertile.
N. W. tropical Himalaya and Sub-Himalayan traci from Punjab to Kumaon,
Africa, Madagascar.
(2) Reynaudiana: Glumes III and IV heteromorphous, III empty.
E. Himalaya, Assam, Bengal, Burma, China, Java.

Specimens kindly sent me by Dr. Stapf, of the African plant from N. Madagascar
(Revd. R. Baron, 6353) with spikelets 0-31 to 0-34 in. long, appear to be identical with those
of the N. W. Himalayan form.

C.—BIOLOGICAL AND CEcoLocicaL NOTES.

83. This plant is sometimes associated with
Aristida cyanantha on gravel and boulders, but as a rule only becomes dominant where
there is a considerable admixture ot sand, and where there is still insufficient moisture for
a vigorous development of Saccharum Munja.

With it are often found the xeuoplnlions form of Saccharum spontaneum and S. Munja
and also Andropogon monticola.

In many respects this grass closely resembles Aristida cyanantha. Like the latter the-
present species, in its usual local habitat, shows a well-marked resting season during
December—January.

It however also grows vigorously on moist loam under cultivation and then shows a
less sharply defined period of rest.

Plants taken from sand at once established themselves strongly in loam and the species
is therefore apparently not so sensitive as Aristida in respect of soil aeration.

Isolated plants are occasionally met with growing well in shady Sal forest and it
probably has a stronger capacity for withstanding shade than 4 ristida.

fi anode

HOLE : SOME INDIAN GRASSES AND THEIR CiCOLOGY. 105

84. The young culm in this species shows an Development
: A ; ; 5 ; of Culms,
average of 10 short basal internodes carrying scales, after which each leaf carries a well- season of
d -e] ed le Ser Tl ae P al eS AS l | ‘ 3 2 ° Vegetative
eveloped lamina. ne average mature flowering culm shows these 10 short basal inter- Activity and
nodes, then 10 long internodes and finally the panicle, thus giving 10 green leaves for the ?*"*
culm which suggests 10 months as the usual period of vegetative activity and indicates that
the average culm usually commences vigorous growth in February. This however has not
yet been satisfactorily determined.
Flowering takes place in September—November and the culms are annual.

The remarks given below, referring to additional points under this head, for Aristida
cyanantha also apply to this species.
85. The remarks given below under these Susceptibil-
© ity to Fire

Ss ia 1st his s 1 ! Damage.
heads for Aristida also apply to this species Tae
Fodder.

D.—Economic USEs.

86. Is of some value on dry sandy, or gravelly,
soil as a pioneer of better vegetation.

Aristida cyanantha, Steud ex Trin. (Plates II and XXXIV to XXXVL.)

F. B. L., VII, 225 (excl. ref. Royle Ill. Bot. Himal.). References.
Western Himalaya and Sub-Himalayan tract from Kashmir to Kumaon. Distribution.
Ascends to 5,000 feet. Extends to Afghanistan.

A.—DESCRIPTION.

Suhi, suhni (Dehra Dun). Vernacular
Erect, slender, attains a height of 10 feet and diameter of 9-21 in. Neve:

Solid throughout.

Glabrous, striate, green in colour.

Secondary culms are often produced from the upper nodes which are usually poorly
nourished and show inferior dimensions.

Glabrous. Leaf-sheath.

The lamina disarticulates from the sheath and after the shedding of the lamina the
culms are covered with the straw-coloured persistent sheaths. The latter are also then shed
and the green culms are left naked from base to apex.

Of uppermost leaf of flowering culm usually well-developed. On poorly nourished tamina.
culms 4 inches long but attaining a length of 14 feet.

Of lower leaves attains a length of 2 feet 4 inches and width of 0-25 in.

Linear acuminate, tapering from the base.

Jointed to and deciduous from the sheath.

[ 105 J

106 INDIAN FOREST MEMOIRS.

Scabrid on margins and dorsally on midrib, especially towards the apex (leaves quite
smooth fide F. B. 1).
Ligule. A minute membranous rim, ciliate.
Pi ee eas Spikelets solitary, pedicelled, on the capillary branches of a terminal panicle.
Primary rachis, branches and pedicels scaberulous.
Panicle 1—2 feet long, conical; branches solitary, fascicled, or sub-verticillate, filiform,
wide-spreading, arcuate.

Sptkelets. Narrow 0:-45—0-65 in. long (excluding awn of glume III), one flowered. Flowering
spikelet is dark purple.
Glume I.—Linear lanceolate, 0-2—0-3 in. long.

Awned, awn 0:05—0-15 in.

Membranous, scaberulous dorsally on keel.

One nerved.

Glume II.—Broad subulate, 0:-4—0-52 in. long.

Awned, awn 0:05—0-13 in.

Membranous, smooth dorsally.

One nerved.

Apex entire or fimbriately toothed.

Glume III.—Linear, 0:3—0-42 in. long, terminating in three long awns.

Awns subequal, or the middle slightly longer than the others, scaberu-
lous, 1-6—2:1 in. long (attain 2°5 in. fide F. B. I.), not articulate
to the glume.

In the flowering spikelet the awns are erect and closely adpressed, in
fruiting spikelet they separate and are wide-spreading.

Glume is chartaceous, convolutely folded, glabrous, three nerved.

Pale—Oblong, hyaline to sub-opaque, two nerved at base, 0-05—0-06 in. long.

Glabrous, completely enveloped by glume ITI.

Apex sub-truncate or 2—3 dentate.

Lodicules—Two, cuneate, obovate, or ob-lanceolate, a little shorter than to subequal
the pale, 0-03 to 0:05 in., glabrous.

Anthers—Three, 0:17—0-2 in. long, yellow, more or less suffused with purple.
Filaments short.

Stigmas.—T wo, 0-05—0-1 in. long, purple; styles short.

The rachilla, which is not produced beyond glume III, is jointed above the empty
elumes I and II, and the latter persist after the fall of the grain. The portion of the
rachilla above the joint and below glume III (=the callus) is bearded with short white
hairs.

Field Charae- The spikelets are often attacked by a fungus Ustilago Aristide-cyananthe Bref. and
ae a spikelet which has been attacked develops a remarkable pod-like structure, which is filled
[ 106 }

HOLE: SOME INDIAN GRASSES AND THEIR GCOLOGY. 107

with the black powdery spores, and these fruit-like bodies give the panicles a most
characteristic appearance.

It is interesting to note that this fungus was originally described by Brefeld from a
specimen “collected in Dehra Dun” and sent by D. D. Cunningham from Calcutta. It
has apparently not been reported from other localities.

C.—BIOLOGICAL AND CEcoLocicaL Notes.

87. This plant is gregarious on the soil-less Habitat.
banks of gravel and shingle found in and near the beds of steep water-courses.

While as a rule markedly gregarious and the dominant species in its own station, this
plant is not infrequently associated with Andropogon monticola and occasionally with
Triraphis. In its local habitat the species completes its annual growth within the period
February——November and shows a well-marked resting season during December—January.

In plants which under cultivation are found to grow well on moist loam the resting
period is less sharply defined, the young culms commencing their development and con-
tinuing to grow slowly after the close of the usual period of activity.

Plants taken from sandy, or stony soil, find some difficulty in establishing themselves
on loam, but some individuals adapt themselves to such changed soil conditions more readily
than others. The species therefore appears to be sensitive to some extent as regards good
soil aeration.

Isolated plants are occasionally seen to have obtained a footing in Sal forest and here
it can be seen to possess some capacity for withstanding shade.

§8. This plant is perennial and is usually Developmeni
gregarious. It consists of a much branched rhizome which constantly grows outwards senconlet
centrifugally in all directions. Aeuisicy, and

The culms are either produced from the branches of the rhizome, or they arise from the “°Y""*’
basal nodes of older culms, either above or below the ground surface.

The culms usually arise close together and well-defined clumps are produced.

The culms are annual, flowering taking place in September—November.

In the moist soil of the garden at Dehra Dun, vigorous growth was observed throughout
the winter of 1909-10 which however was exceptionally mild. In the dry localities where
it is usually found in nature there is as a rule a well-marked period of rest during December
and January.

The aerial culms usually die back after flowering, but their lower internodes occa-
sionally persist and produce secondary culms from their nodes.

The young culm usually shows 6 or 7 very short basal internodes carrying scales, above
which all the leaves have a more or less well-developed green lamina. The mature flowering
culm also shows the short 6—7 basal internodes, as a rule aggregating less than an inch in
length, defined by the interncdes of the scales, then 9 long internodes, then a comparatively

fe torn y

Susceptibil-
ity to Fire
Damage.

Utility as
Fodder,

References.

Distribution.

108 INDIAN FOREST MEMOIRS.

short internode, and finaily the terminal segment carrying the panicle. This gives 10 green
leaves for the mature culm which suggests 10 months as the period of activity and indicates
that the average culm usually commences vigorous growth in February. This, however, has
not yet been satisfactorily determined.

The culms are distinctly green, while the green leaf lamina is often deciduous in
October, the sheath persisting for some time longer. It is possible that the green culms to
some extent function as leaves.

89. This species usually shows a well-marked
period of rest, in the dry localities which here form its natural habitat, during the months
December—January. During these months the clumps contain only the old culms which
fiowered the preceding September—November, while next year’s culms have not yet com-
menced their growth and exist as large buds well protected by hard scaly leaves.

Soon after flowering, also, the leaf-sheaths fall away from the culms and during the
veriod of rest the clumps have a very striking appearance, consisting as they do only of
naked green culms terminating in the panicles, with no leaves and no leafy immature culms.
At this period the old culms are not yet thoroughly dry and the clumps contain practically
no inflammable material.

Although the species has a tufted habit and occurs on sandy soil it is, in consequence
of the above facts, not subject to damage by an early fire.

A late fire, when the old culms and soil are thoroughly dry and the young culms have
commenced their growth, may cause severe damage.

90. This species is believed not to be utilised
for fodder.

D.—Economic USEs.

91. The upper portion of the flowering culm
with the attached panicle is cut off and several of these being tied in small bundles the
latter are used as brooms.

This species is of considerable value as a pioneer on gravel and shingle banks which
there resists erosion and gradually improves the soil and prepares the way for better
vegetation.

Andropogon monticola, Schult. (Plates XX XVII to XXXIX.)

F. B. I., VII, 192; H., pp. 557—558 (under 4. monticola and A. Trini); C., II, 985.

P., II, 1205.

Throughout India, especially in hilly tracts, from N. W. Himalayas southwards, as-
cending to 6,000 feet.

Extends to Burma, Ceylon, Afghanistan, Africa.
i 08) if

HOLE: SOME INDIAN GRASSES AND THEIR (ECOLOGY. 109
A.—DESCRIPTION.

Dhaulu, gurla, gorha (Dehra Dun). Gogad, ghora (Central India, Bombay). Vernacular

Geniculately ascending, attains a height of 7 feet (in local plant usually 3—5 feet) Calas
and diameter of 0:2 in.

Slightly compressed.

Solid.

Smooth.

Axillary leafy and flowering branches are usually developed from all the upper nodes
except the one next below the panicle.

These branches developing within the sheaths, push out the latter away from the culm
and often produce a characteristic fan-shaped appearance.

Glabrous. Leaf-sheath

Compressed, keeled, especially of the lower leaves.

Shorter or longer than the proper internode.

Of uppermost leaf of flowering culm usually mucroniform but attains a length of Lamina.
3°3 in.

Of lower leaves attaining a length of 1 foot 5 inches and width of 0:3 in. Verna-
tion conduplicate.

Linear acuminate, tapering from base.

Seabrid on margins. Sometimes also scabrid dorsally on midrib, and scaberulous
above, especially towards the apex. Often ciliate towards the base with tubercle-based
hairs, at least when young.

A minute membranous rim, ciliate. Tienes

Spikelets in clusters of 3 (a central sessile ¢ spikelet with two lateral pedicelled 3 miorescence
spikelets), the clusters being terminal and solitary on the capillary, sub-verticillate, branches
of a terminal panicle.

Panicle rachis and branches smooth or scaberulous.

Panicle 2 inches—6 inches long. Ovate to sub-cylindric, yellowish to purplish in colour.

The branches of the flowering panicle are more or less horizontally spreading, those of
the fruiting panicle are erect and closely adpressed to the rachis.

Each cluster of three spikelets is deciduous as a whole, an ovate scar, shortly fringed
with stiff rufous hairs, then terminating the panicle branches and becoming conspicuous.

The swollen base of the panicle branches apparently functions as a pulvinus in connec-
tion with the movements of the panicle branches.

The flowers of this species are much visited by small pollen-collecting bees.

Laterally compressed, 0-17—0-23 in. long, excluding the awns (Hackel gives 0-16 to Sessile
0-28 in.). apie
Glume I.—Laterally compressed, narrow-oblong, embracing the margins of II at

; the base.
Chartaceous 2—4 nerved.
[ 109 ]

110 INDIAN FOREST MEMOIRS.

Hispidly ciliate dorsally on keel towards the apex or almost glabrous.

Often scaberulous dorsally on nerves and minutely pubescent with.
adpressed hairs dorsally near margin.

Apex sub-truncate or two dentate.

Glume II.—Laterally compressed, broader than I, obtusely keeled, sub-coriaceous,
three nerved.

Margins broad, hyaline, membranous, ciliate or not.

Very variable as regards its hairiness. Sometimes almost glabrous, at
other times hispidly ciliate dorsally on keel with long white or
rufous hairs more or less from base to apex.

Sometimes also pubescent, or minutely villous, dorsally on keel and on
lateral nerves.

Awned, awn 0:1—0-25 in. long.

Apex entire or two lobed.

Glume III.—From 4 to subequal IT, linear, hyaline, ciliate, enerved or indistinctly
1—3 or more nerved.
Apex obtuse.

Glume IV - -The narrow 3 nerved base of an awn.

Basal half or 2rds hyaline, membranous, upper portion chartaceous.
Awn geniculate 0:-40—0-7 in. long including the twisted column.
(Attains 1:3 in. fide F. B. I.) Margins ciliate or not.

Apex entire or two lobed.

Pale.—Sometimes present, very narrow, 0:05 in. long.
Lodicules—Two, cuneate, glabrous.

Anthers.—Three, 0-07 to 0:13 in. long, yellow or purple.
Stigmas.—Two, 0-06—0-08 in. long, laterally exserted at base of spikelet. Yellow.

iets Dorsally compressed, subequal the sessile spikelet.

Pedicel is less than half the sessile spikelet, usually about 4rd the spikelet, and is
densely ciliate on two margins with stiff rufous (or white, fide F. B. I.) hairs, the upper of
which are shorter than to subequal the spikelet.

Glume I—Lanceolate, membranous, 5—7 nerved. Minutely pubescent with ad-
pressed hairs dorsally, especially towards the apex, or almost
glabrous. Sometimes ciliate dorsally on midrib and on marginal
nerves, especially towards the apex. Acute or shortly awned.

Glume I7.—Subequal I, 3 nerved, margins incurved, long ciliate, apex acute or-
mucronate, glabrous dorsally.

Glumes III and IV.—Linear, hyaline, ciliate, enerved or indistinctly nerved.
Pale.—Sometimes present, as in sessile spikelet, but slightly longer, very narrow.
TO 7

HOLE : SOME INDIAN GRASSES AND THEIR CECOLOGY. ea:

B.—Taxonomy.

92. Hackel separates two species mainly by
the following characters :—

Andropogon monticola, Schult—Glume II of sessile spikelet keeled, the keel from
base to 2 or 2 its length densely pectinate ciliate with long rigid rufous hairs, shortly white
hispid in upper $.

Andropogon Trinii, Stewd—Glume II of sessile spikelet keeled below the apex only,
keel white ciliate.

Remaining 2 or # of the glume dorsally, not keeled, glabrous.

In F. B. I., both of the above are included as varieties in one species named A. monti-
cola, Schult, and the author remarks : “ I am unable to classify the varieties of this common
and variable plant in accordance with geographical areas on other considerations. This, if
possible, must be effected by field botanists in India. There is every gradation from the
coarsely hirsute keel of monticola, to the perfectly smooth of some states of Trini.” The
author, however, differentiates and describes an additional variety which he names robustus.
The latter is founded on a plant collected by Strachey and Winterbottom which shows the
basal 2 or 2 of glume II of sessile spikelet more or less pectinately ciliate with white
hairs. All these three described forms appear to vary greatly, as regards their habit and
vigour of growth, in response to the moisture conditions of the habitat and also according
as whether, or not, the plants are habitually grazed, cut for fodder, or periodically burnt.
The colour of the cilia of glume II of the sessile spikelet (pale or white in robustus and
rufous in monticola), accordingly, appears to be the chief difference between monticola and
robustus, rather than any difference in the habit, and these forms appear to have different
and fairly defined areas of distribution (monticola occurring chiefly in Central and Southern
India, while robustus is mainly found in N. India, in the outer N. W. Himalayas and Sub-
Himalayan tract). In the F. B. I. the author includes under robustus a plant (No. 555 col-
lected by King at Goona, Central India), glume IT of the sessile spikelet of which is dorsally

ciliate in the basal 2 with long rufous hairs and which appears to be referable rather to
monticola. In the locality with which the present paper is chiefly concerned the forms of
this plant which occur are robustus (taken in the narrow sense of Strachey and Winter-
bottom’s plant alone) and Trina. Both when studied with herbarium specimens and also
in the field, however, the local plant gradually and imperceptibly passes from typical
robustus to typical Trini. It is believed that a similar passage takes plaee between monti-
cola and Trinii in Central and 8. India. Further research, however, in the field is required
to establish this and also to decide whether monticola can be separated from robustus. For
the present it seems desirable to follow the F. B. 1. and to unite these forms into a single
species and this has been done in (A) above. So far as the local plant is concerned specimens
with the more hairy glume II tend to occur in localities where there is a scarcity of available
ip ree

Habitat.

112 INDIAN FOREST MEMOIRS.

moisture, both on the dry ridges and slopes of the Siwalik Hills and also (rarely) on water-
logged soil, and the writer believes that the characters which have been utilised to define
these varieties vary in response to the factors of the habitat and particularly in response to
‘he available water supply.’ Provided that the development of the plants has not been
interfered with by grazing, grass-cutting, or other agency, those plants with the more hairy
glume IT are usually less robust and less coarse, or rank, than the others, and they are
therefore as a rule most valued for fodder and are distinguished locally by the vernacular
name of dhaulu, whereas the coarser plants with smooth glume are called gurla. As this
grass affords a valuable fodder and is sometimes cultivated, in consequence, it is important
to determine the extent to which its characteristics are constant. If, as suggested above,
they depend on the available moisture, it is obvious that cultivation of this grass on good
agricultural land, with a large quantity of available moisture, would result in producing an
inferior class of rank, coarse fodder.

The local forms of the species are now being cultivated in the Experimental Garden at
Dehra Dun with the object of studying the effect of different soils and varying quantities
of available moisture on their characters.

This will, it is hoped, indicate which forms (if any) of this species are of sufficient im-
portance and sufficiently constant to deserve separate definition and description.

To separately define forms which are connected in the field by numerous intermediates
greatly increases the difficulty of identification and is of little practical utility inasmuch
as such forms are not likely to remain constant under the ordinary conditions of field-culti-
vation.

C.—BIOLOGICAL AND CEcoLocicaL NoTEs.

93. This plant is in this locality almost in-
variably found on dry sandy, or stony, soil, with little capacity for holding water.

It is a frequent associate of Aristida, Triraphis and Saccharum Munja. These plants.
however are all more vigorous and robust than the present species and Andropogon monti-
cola only exceptionally becomes the dominant species under conditions unfavourable for the.
vigorous development of these stronger competitors. Such conditions are provided by the
hot, waterless, and more or less soil-less, ridges and slopes of the Siwaliks, especially on the
south of the range, and on deeper moister soil where the stronger competitors are prevented
from developing by fire, or other factors. At least one area in the present locality has been
seen near Bullawala, where repeated firing has killed out the coarse savannah species Saccha-
rum Munja and has resulted in making Andropogon monticola the dominant grass. This
would undoubtedly occur more frequently were it not that the burnt grasslands are
frequently heavily grazed.

' The interesting fact which has been noted regarding this species that the flowers are much visited by bees naturally suggested that
intercrossing might be responsible for intermediate forms and for confusing the boundaries of distinct types, but field study indicates the
available mcisture supply as the chief factor affecting the characteristics in question.

Lr Ue J

HOLE : SOME INDIAN GRASSES AND THEIR (ECOLOGY. 113

Andropogon monticola is probably the most valuable fodder grass in this locality and
on grazed areas it is severely handicapped by being constantly eaten down, level with the
ground, which not only seriously diminishes the vigour of the plant, but prevents the pro-
duction of flowers and seed. Moreover in such burnt and grazed areas, on sandy soil,
Zizyphus Jujuba frequently appears, woodland then replacing the grassland.

This plant is not infrequently seen growing in the middle of Zizyphus bushes on
grazing grounds and therefore has some capacity for withstanding shade, but in shady
stations its place is usually taken by Apluda varia. Plants of this species are only rarely
found on poorly aerated heavy soil and on loam its place is usually taken by Andropogon
intermedius, or Andropogon contortus. Plants taken from sandy soil and then cultivated
on loam often adapt themselves with considerable difficulty to the new soil conditions and
the species appears to be as a rule sensitive as regards soil-aeration. This is a point of
considerable importance as this valuable grass is sometimes cultivated and a case has been
seen where the attempt to introduce it on heavy soil had ended in failure.

94. A perennial, usually gregarious and
forming centrifugal clumps as in Aristida. The young basal culms examined of this species
show an average of 11 very short basal internodes, carrying closely imbricating scale-like

leaves, with little or no lamina, after which each leaf carries a more or less well-developed
lamina.

The flowering culms examined show, in addition to the short basal internodes,* 11 long
internodes and finally the panicle, thus giving 11 green leaves for the culm, and indicating
11 months as the period of vigorous vegetative activity. The culms are annual, but the

exact period of activity varies considerably with the locality and the quantity of available
moisture.

In the moist loamy soil of the Dehra Dun Experimental Garden vigorous growth was
observed throughout the winter of 1909-10, which however was exceptionally mild. In
moist localities, therefore, growth apparently continues more or less throughout the year.
In the dry localities where this species is usually found locally, however, there is as a rule
a well-marked period of rest during December and January and in such places the usual
period of vigorous activity is 9 months from February to October.

The majority of the culms start growth in February and flowering takes place in
September—October. Culms commencing vigorous growth in February have shown an
average number of 9 long internodes. After the production of the terminal panicle, the
axillary shoots developing from the lower nodes of the culm (=secondary culms) proceed to
flower in their turn and in this way flowering may continue for several weeks or even
months.

1 It must be noted that the number of short basal internodes varies with the position of the culm. Culms which originate at, or
below, the ground surface usually show the greatest number of short basal internodes, while culms which arise from the upper axils
of older culms often show only 1 or 2 such internodes, and sometimes none at all.

f 113%

Development
of Culms,
Season of
Vegetative
Activity and
Flowering.

Susceptibil-
ity to Fire
Damage.

14 INDIAN FOREST MEMOIRS.

The aerial culms usually die back after flowering, but their lower internodes
occasionally persist and tertiary culms are sometimes produced from the lower nodes of the
secondary culms.

95. In this locality the present species is

usually dominant in dry localities and here it usually shows a well-marked period of rest
during December—January.

During these months the clumps contain only the old culms which flowered the preced-
ing September—November, while next year’s culms are visible as large buds below, or
close to, the ground surface.

Although the species has a tufted habit and usually occurs on dry sandy soil, it is not
as a rule severely damaged by an early fire, owing to the fact that the young culms are
still, as a rule, below the ground surface and well-protected by their imbricating scaly
leaves.

The majority of the leaf-bearing branches of the mature culms also spring from the
upper internodes of the culms and an early fire which destroys the dry inflammable leaves
on the mature culms does not, as a rule, develop a dangerous intensity of heat at the base
of the clumps in the neighbourhood of the buds from which the next season’s culms will be
developed.

This species therefore is as a rule less subject to severe damage by an early fire than is
Saccharum Munja for instance.

D.—Economic.

96. Lisboa writes as follows regarding this
species :—

“Tn Mount Aboo it is reckoned as a good fodder grass and the grain used as food by the
natives,” and under the name of Andropogon serrulatus, Trin., he adds : “ Said to be good
fodder, used much in Poona, but reports from other places unfavourable.”

In the local Siwalik Division it is one of the most valuable fodder grasses.

1 List of Bombay Grasses by J.C. Lisboa, 1896, p. 81.

ee rete]

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 115

CoH ACE DRO: JE Tr

(ii) Summary of Chief Characteristics of Species dealt with.

97. Before proceeding to the next chapter in
which a few suggestions are made regarding the practical treatment of grasslands, it is
perhaps advisable to briefly summarise the principal characteristics of the species described
above.

This is done in the following table in which the species are arranged in order accord-
ing to their relative (1) capacity for growing vigorously in a dry soil, (2) capacity for en-
during shade, (3) capacity for enduring bad soil-aeration and (4) capacity for withstanding
the injurious effect of an early fire in December-—February. The species with the greatest
capacity for thriving in a dry soil, and for enduring shade, bad soil-aeration and fire, res-
pectively, has been in each case placed first.. Species joined by a bracket are believed to have
an approximately equal capacity. With regard to the capacity for thriving in a dry soil,
also, the duration of the culm }s of interest; this has therefore been noted and it will be seen
that the species able to grow well in dry soil usually have annual culms :—

Capacity for thriving in » Duration of | Capacity for enduring | Capacity for enduring bad Capacity for resisting injurious
. | . . . -
dry soil. | Culm. Shade. | Soil-Aeration. action of an early Fire.
. | | . eye
Andropogon monticola. Al | ( Saccharum Narenga. ee arundinacea. ( Aristida cyanantha.
|
Aristida cyanantha. Erianthus Ravenne. | (Saccharum spontaneum. { Triraphis madagascariensis.
Saccharum spontaneum. + Annual. Imperata arundinacea. Erianthus Ravenne. (Andropogon monticola.
Triraphis madagascariensis. | Triraphis madagasca- | ( Triraphis madagascariensis, ( Erianthus Ravenne.
| | riensis. |
(Imperata arundinacea. J | | Saccharum Narenga. Saccharum Narenga.
| | (Saccharum Munja.
4 Saccharum Munja. BY (Sachharum Munja. Imperata arundinacea.
| | Saccharum spontaneum. | |
(Saccharum Narenga. ‘\ Biennial. |4 |< Andropogon monticola. (Saccharum spontaneum.
| | Andropogon monticola. |

| | |
Erianthus Ravenne. B) | Aristida cyanantha. Saccharum Munja.

| Aristida cyanantha.

The above table summarises the information obtained from careful field observations
in the lecality defined in Chapter I above, supplemented by the results obtained from the
limited experimental cultures which alone have been possible up to date. It is therefore
to be expected that instrumental field observations and extended and careful experiments
designed in such a way as to give accurate information regarding the effect of
each individual factor on the growth of the different species will to some extent modify the
conclusions summarised above. On the whole, however, it is believed that the conclusions
drawn are fairly accurate and that they will be of utility. It is believed that the observa-

tious which have been made regarding each species embrace the principal classes of habitat
[ 115 }

116 INDIAN FOREST MEMOIRS.

in which the plant is naturally found and that therefore the resuits obtained will to a
considerable extent apply to areas other than that in which the information has been collect-
ed. Alterations in climate, by affecting the period of vegetative activity, will no doubt to
some extent modify the above arrangement as regards fire-damage.

In the present paper, however, what appears to be the normal life bistory of the culms
of each species has been carefully explained and the connection between this development
and the damage done to the plant by fire has been indicated. This information therefore
will, it is believed, enable anyone to form a good idea of the liability of these species to fire
damage in any particular locality, by observing the differences in the life history produced
by local climatic conditions.

As regards ability to thrive in a dry soil, the arrangement given above merely refers
to what is believed to be the average capacity of the species, as it occurs in nature. Thus
the swamp form of Saccharum spontaneum can only thrive, 2.e. attain average dimensions
on a soil containing large quantities of water and the quantity of water actually utilised by
a plant of average vigour of this form is probably not less than that utilised by an average
plant of Saccharum Munja. On the other hand, however, the majority of the individuals
composing the species Saccharum spontaneum, as it exists in nature, do not belong to the
typical swamp form but to the forms commonly found on dry sand and wet agricultural
lands. Taking these into consideration therefore the average plant of Saccharum spon-
taneum is able to thrive on a drier soil than the average individual of Saccharum Munja.

A similar point of view has been taken in arranging the species with regard to other
factors.

Thus as regards soil-aeration in species like Andropogon monticola, Saccharum Munja,
and Aristida, which in nature as a rule are found on well-aerated soil, a comparatively few
individuals can be found growing well on heavy, less aerated soil. Plants of all these
species taken from well-aerated soil and then cultivated in Dehra Dun on loam have all
shown in a single season :—

(1) Some plants which entirely fail to adapt themselves to the new soil and die, or
which adapt themselves with difficulty and show inferior growth.
(2) Some plants which adapt themselves at once to the new soil and show average

dimensions.

We almost certainly have an example here of individual variation and of the power
possessed by individual plants, in a greater or less degree, of adapting themselves to new
conditions, and if the comparatively few individuals of these species which can be found in
nature growing on heavy soil are experimentally cultivated, they would almost certainly
show greater capacity for enduring poor soil aeration (probably equal to that of Saccharum
Narenga or Erianthus) than those taken from sandy, stony places.

At the same time the average plant of these species, at all events in the present locality,
certainly does not occur in dense, poorly aerated soil and the above arrangement has been

r we 7

HOLE : SOME INDIAN GRASSES AND THEIR CCOLOGY. 117

based on what is believed to be the capacity of the average individual of the total aggregate
of individuals which compose the species in nature.

Again as regards fire-damage, Saccharum spontaneum in a swamp or wet clay is
practically free from early fire-damage, but, on the other hand, individuals on dry sand may
be considerably damaged, and on the whole, therefore, this species cannot be placed in the
first class of fire-resistant plants.

| 11% J

115 INDIAN FOREST MEMOIRS.

CHAP? ER tim:

Suggestions regarding the Practical Treatment of Local Grasslands.

98. Before attempting to make suggestions.
regarding the practical treatment of grasslands it is obvious that we must clearly define the
object which we desire to attain, seeing that it is frequently impossible to satisfactorily
attain two distinct objects by one and the same method of treatment. It is therefore
highly advisable at the outset to draw a line between—

(1) Grasslands required for grazing and for the production of fodder,
(2) Grasslands which are not required for fodder-production and which it is desir-
able to afforest,

seeing that the production of grass and production of trees are two distinct and more or
less mutually incompatible objects which cannot as a rule be satisfactorily attained at the
same time, on one and the same area.

Occasionally grasslands are of course required for purposes other than those defined
above, e.g. for the production of munj fibre, of thatching grass, and so on, for each of
which a more or less special treatment is required, but as a general rule the maximum pro-
duction of the best quality of wood and fodder respectively is undoubtedly the most import-
ant object which we desire to attain by a judicious treatment of our forest grasslands.

A point of some importance consists in the fact that grasslands tend to become highly
inflammable in the hot season and therefore greatly increase the danger of fire-damage in
the adjacent woodlands. It is however possible to prevent such damage, (1) by successfully
afforesting our grasslands and thus killing out the grasses or (2) by managing these areas
chiefly with a view to satisfying the demand in respect of fodder production and grazing
needs, in which case the old, inedible and inflammable grass is either removed by burning,
or else is not allowed to accumulate, owing to the young grass being continually cut or

erazed.
The suggestions which it has been possible to make regarding the treatment of the

grasslands in the locality here dealt with and which are based mainly on the detailed study
of the representative species described above, have therefore been arranged with reference

to these main objects as below :—

A.—Treatment with a view to Fodder-Production.

99. The first point of primary importance
consists in this that the robust hygrophilous, or markedly mesophilous, grass species are, as
a rule, too coarse to yield good fodder and only the young shoots and young leaves, or the

Ef wey

HOLE : SOME INDIAN GRASSES AND THEIR GCOLOGY. 1h,

older shoots and leaves of the less vigorous individuals, of such species are consequently
utilised to any considerable extent by cattle.

Typical species of this description are the common coarse savannah grasses Saccharum
Munja, Saccharum Narenga, Erianthus Ravenne and Imperata. The last, being probably
the most valuable for fodder, may be taken as fairly representative of this class.

If this grass is allowed to develop undisturbed, the grasslands composed of it chiefly
consist throughout the cold and hot season of a mass of immature culms each bearing
several large mature leaves which are more or less dead and dry and useless for fodder
while the still younger shoots are minute and inconspicuous. The presence of the old dead
leaves not only makes it difficult for cattle to find the few young green shoots that do exist,
or to obtain a mouthful of fodder free from a considerable admixture of unpalatable
material, but seriously retards the development of the young shoots by interfering with the
passage of moisture into the soil and with the access of light. Until the rains have well set
in and the development of the young culms has well advanced the quantity of green material
available on such grasslands is exceedingly small and in the rains the majority of it is too
coarse to be palatable. On the other hand, if these grasslands are fired when the old leaves
are thoroughly dry, about December, the young shoots usually start vigorous growth in Janu-
ary—February and a good supply of fodder becomes available from February to June.
If the grasslands are well grazed throughout the rains and thereafter, the vigorous develop-
ment of the plant and the production of the coarse unpalatable full-sized leaves oecomes
impossible and the grassland yields a practically perennial supply of slender culms and
small green leaves. If it is impossible to check vigorous growth during the rains, fire
should be annually employed to increase the fodder supply.

In the case of the majority of the coarse savannah grasses in the present locality and
probably also in other parts of India, fire alone is rarely capable of killing out the species
and is directly beneficial in clearing away inedible material, in enabling cattle to gain easy
access to the young palatable shoots and in increasing the production and accelerating the
erowth of the latter at a season when fodder is usually scarce. The only coarse savannah
species dealt with in this paper which is liable to be killed out by repeated early firing is
Saccharum Munja when growing on dry sandy soil. In this locality at least one area exists
near Bullawala where this has taken place and where a far more valuable fodder-grass,
viz. Andropogon monticola, now occupies the ground, owing to the fact that it suffers less
severely from fire damage.

In addition to the direct action of fire on the grass plants, also, we have to consider
another important point, viz. the effect of fire upon the soil. It is obvious that any factor
which decreases the water-content of the soil, ipso facto, renders a locality unfavourable for
the vigorous growth of robust mesophilous and hygrophilous species and tends to produce
conditions suitable for the growth of more xerophilous plants.

fo biG |

120 INDIAN FOREST MEMOIRS.

This is exactly what fire really does and it appears to act in two principal ways :—

(1) By directly destroying the humus which may exist in the surface soil and by
effectually preventing any further addition to the existing supply through the
destruction of all dead leaves and organic débris.

(2) By altering the character of the clayey constituents of the soil in such a way
that loam to a great extent loses its power of retaining water. The texture
of the soil becomes coarser and percolation is increased.

The grassland known as Gola Tappar in the present locality which is shown in Plate
XL is believed to be an instructive example of this drying action of fire on the soil. This
grassland is believed to occupy ground which was at one time covered with Sal forest, the
latter having been cleared for cultivation. The soil is a loam, apparently suitable for Sal
and more or less similar to that in the adjoining Sal forest which surrounds the area on all
sides, while Saccharum Narenga, the Sal-locality-indicator, is a common grass in the area.
This Tappar is reported to have been regularly burnt for many years and the result is
remarkable. Instead of a dense growth of the giant Saccharum Narenga which would
naturally have been expected in such a locality we find a good deal of both Saccharum
Narenga and Saccharum Munja, but only of very inferior vigour and dimensions, while
more significant still, we find the xerophilous Andropogon contortus actually able to occupy
considerable areas and to become dominant in a locality where it would, under ordinary
circumstances, have been ousted by the far more vigorous competitors named. Experience
elsewhere in similar soil has shown that fire is not likely to have here caused the poor growth
of the Saccharum sp. noticed, by direct injury to the plants and both the species here flower
freely which would not be the case if the direct fire damage was severe and most of the one-
year culms destroyed. It is therefore concluded that, in this case, the poor growth of the
coarse savannah grasses and the unusual prevalence of the more xerophilous Andropogon
is mainly due to the drying of the soil and that this area affords a good example of the
power possessed by fires of replacing hygrophilous, or strongly mesophilous growth, by a
xerophilous type of vegetation. It has often been stated that fires favour the development
of coarse species and are injurious to the more valuable fodder grasses. From the above
remarks, however, it will be seen that such a generalisation is impossible, that fire under
certain circumstances can kill out a coarse, comparatively valueless, fodder-grass like
Saccharum Munja and may cause it to be replaced by a more valuable fodder-species like
Andropogon monticola and that, as a general rule, by its action in drying the soil, fire is
able to produce conditions favourable for comparatively xerophilous species which are
usually more valuable for fodder than the hygrophilous or strongly mesophilous grasses.
Tn areas where the latter prevail it must I think be accepted that firing is, on the whole,
decidedly beneficial in improving the fodder supply. Further than this, however, it is
impossible to generalise for, when dealing with the xerophilous or less strongly mesophilous
species, it is obvious that some are of more value as fodder than others and that
some are avoided by cattle on account of their peculiar taste, odour, cutting leaves,

[1207]

HOLE : SOME INDIAN GRASSES AND THEIR CECOLOGY. 27

hairs, barbs, and so on. Whether firing is, or is not, a beneficial treatment to
adopt in grasslands of such species will depend entirely on the characteristics of
the particular species dealt with, on the nature of competing species, and on the condi-
tions of the locality. Thus Andropogon contortus Linn., Andropogon foveolatus Del. and
Andropogon annulatus Forsk. are probably all to be reckoned as xerophilous species and all
yield in consequence fairly good fodder. The first, however, is often avoided by cattle on
account of the characteristic masses of its long-awned and barbed fruiting spikelets and for
this reason it is often considered of little value in India. It is, however, hardly correct to
speak of this plant as a coarse species which is a term best reserved for the more robust
hygrophytes and mesophytes and it should be noted that, apart from its peculiar fruiting
spikelets, this species is a good fodder and Duthie writes as follows regarding it : “ Largely
used as fodder both before and after it has flowered, but chiefly when it is young and tender.
* * On Mount Abu the people consider it the best fodder grass they have. * * In
Australia it is looked upon as a splendid grass for a cattle run, as it produces a great
amount of feed.” It is undoubtedly a fact that this species frequently tends to increase in
burnt grasslands, whereas in similar stations which are protected it has to give way, to a
considerable extent, to such species as Andropogon foveolatus and Andropogon annulatus.
An experiment initiated by Mr. R. 8. Pearson some years ago in the Panch Mahals Division
of Bombay and of which details have kindly been supplied to me has shown this to have
occurred in that locality, while in Dehra Dun a comparison of the grasslands known as
Zabarkhet and Gola Tappar indicates that in protected areas Andropogon contortus is to
a great extent replaced by Andropogon intermedius. The fact that Andropogon contortus
tends to spread in burnt areas is probably due to the fruit being fire-resistant, on account
of the very hard enclosing glumes.

Thus it will be clear that, while a generalisation, to the effect that fire is as a rule bene-
ficial in areas where hygrophilous or coarse mesophilous grasses predominate, appears to be
justifiable, we cannot proceed further to say that fire is as a rule beneficial with all species.

As regards local grasslands, therefore, it may be said that where Erianthus, Saccharum
Narenga, Saccharum Munja and Imperata are dominant, early firing is as a rule decidedly
beneficial so far as the fodder supply is concerned inasmuch as this (1) increases the quan-
tity of palatable fodder available from these species themselves, and (2) either by directly
injuring the plants themselves (as in the case of Saccharum Munja on sand), or indirectly
by drying the soil, tends to replace these coarse species by less vigorous and generally more
valuable fodder-grasses, such as Andropogon monticola on sand and Andropogon contortus
on loam.

The value of Saccharum spontaneum for fodder is also, as a rule, probably increased
by early firing which cleans the clumps of débris and old culms and insures the production
of a good supply of vigorous young shoots from the ground. In places where it cannot be
easily fired owing to the wet conditions of its habitat and where vigorous growth during the

1 Fodder Grasses of Northern Jndia by J. ¥. Duthie, p. 32.
Fie 7

122 INDIAN FOREST MEMOIRS.

rains cannot be checked by grazing, the grass should be cut close to the ground in the cold
season, as this will to a great extent have the same effect as fire, although of course it is a
more expensive treatment.

Triraphis and Aristida are believed to be little used for fodder, but the young shoots
would probably be of value during the period February to June and the removal of the old
flowering culms by fire, or cutting, in the cold season would be advisable where possible.

As regards grasslands where valuable fodder-grasses are already the dominant species
it is obviously advisable, as a rule, to let well alone and not to run the risk of favouring the
introduction or extension of less valuable kinds by firing. On sandy soil where Andro-
pogon monticola is already dominant firing may to some extent directly injure that species
and may result in bringing in less desirable xerophilous species of Andropogon, Eragrostis
and others. On loam where species like Andropogon annulatus, Andropogon intermedius,
Andropogon foveolatus, Iseilema, and others are dominant, firing very frequently results
in increasing the proportion of Andropogon contortus and it is clearly a question for local
officers to decide, in the light of local prejudice and demand, whether or not this is desir-
able.

On grasslands where valuable perennial species like Andropogon monticola are already
dominant, it is clear that the best treatment to apply, in order to obtain the maximum of
good fodder with a minimum of damage to the plants, consists in protecting the areas from
fire and grazing and in cutting the grass when flowering is just commencing. Grass so cut
should be made into hay, or silage, and the cattle stall-fed. Heavy grazing not only directly
weakens the plants by continually depriving the growing culms of their green leaves more
or less completely, but considerable damage is done by the treading of cattle in the way of
destroying young shoots and also by treading down the soil around the plants, and thus
checking their outward growth. In sandy soil the rhizome-branches are often found to be
exposed above the ground surface on account of this injurious action and in loam the soil
becomes injuriously hardened and compacted.

B.—Treatment with a view to Afforestation.

100. As regards the question of afforestation,
the local grasslands must first be divided into—
(1) Those in which the vigorous growth of most woody plants is made impossible by
frost, the latter being in this case the factor of dominant importance.
(2) Those where the frost damage is not severe and where woody plants can establish
themselves.

As examples of the first we may take the areas locally known as Zabarkhet in Lachi-
wala Range, Gola Tappar in Tirsal Range (see Plate XL) and Koilpura Tappar in Ganges

Range.
f 122 4]

HOLE : SOME INDIAN GRASSES AND THEIR CECOLOGY. 123

Examples of the second are Kanswala Tappar and parts of Palasi Tappar in Lachi-
wala Range.

Grasslands of the first class appear to owe their characteristics mainly to the fact that
they are surrounded on all sides by dense forest, on account of which free circulation of air
is impossible. These areas are therefore subjected to intense heat by solar radiation during
the day and hot season and to great cold by terrestrial radiation during the night and cold
season. The natural result of these influences is to diminish the water-content of the soil
and to cause great damage from frost, 7.e. to create conditions only suitable for the develop-
ment of xerophilous, or the less mesophilous, and frost-hardy species.

101. On the loamy soil prevailing in Zabar-
khet, which is fire-protected, the grass which should be dominant is the coarse mesophilous
Saccharum Narenga. The latter, however, is here seen to grow vigorously, as a rule, only
near the edges of the grassland where the shade of the surrounding forest keeps the soil
moist, while, in the most exposed portions of the area, the far less mesophilous Andropogon
intermedius tends to become dominant.

In Gola Tappar which is regularly fired, but which lies higher and where there is
possibly more air circulation, the drying action is intensified by fire and here again
Saccharum Narenga is seen to be of inferior vigour and dimensions. Here, however, pro-
bably on account of its fire-hardy fruiting spikelets, Andropogon contortus to a consider-
able extent takes the place of Andropogon intermedius.

Grasslands of this type appear, as a rule, to have been caused by clearings which were
made in the forest for the purposes of cultivation and then abandoned.

In those grasslands where, owing to the small size or shape of the area cleared, or to
the configuration of the ground, a certain amount of shade is derived from the surrounding
forest, or where a few trees were left scattered over the area at the time of clearing, the
effect of such shade in increasing the water content of the soil is very marked. As
already noted above, Saccharum Narenga is vigorous near the edges of Zabarkhet, while a
walk through Gola Tappar only showed a single vigorous plant of the moisture-loving
Erianthus Ravenne which was situated beneath the ample shade of a large tree of Ficus
bengalensis. ‘This prevalence and increased vigour of strongly mesophilous grasses in
these areas near the shade of trees is very marked, has been frequently noticed, and is a
very striking illustration of the value of shade under these conditions in the way of increas-
ing the water content of the soil.

102. As regards the question of frost, it
should first be noted that low temperature is believed to be injurious to woody plants in two
principal ways :—

(a) Indirectly by desiccation. When the soil temperature falls below a certain point

the roots are no longer able to absorb the necessary supplies of water. While
r ee 7

124, INDIAN FOREST MEMOIRS.

the roots are thus inactive, the leaves may be actively transpiring moisture
under the influence of bright sun-light and dry air-currents. The ieaves and
branches of woody plants may thus become completely dried up and killed, in
the same manner as in a case of drought.

(b) Directly by the low temperature of the air in contact with the aerial leaves, twigs
and branches.

The first mode of action can often be recognised in the forest by the fact that those
aerial portions of a plant tend to be injured first which are furthest removed from the water
supply ascending from the roots, 7.e. the base of the leaf and those parts of the lamina
nearest the main nerves are often found to be uninjured and green when the rest of the
leaf is shrivelled and brown, while the terminal twigs and branches tend to die back first.

On the other hand, the second mode of action can often be recognised by the fact that
those parts of the plant are most injured which are in contact with the coldest air, irrespect-
ive of their position at the base or apex of the plant. Thus those leaves or portions of
leaves which are shaded by neighbouring leaves and thus protected from radiation will be
often found to have escaped damage when the rest of the leaves have been killed. In grass-
lands unrestricted radiation tends to produce a layer of cold air just at, or immediately
above, the level of the grass and that radiation is chiefly responsible for the frost damage
done to woody plants in the grasslands now under consideration is often clearly indicated
by the fact that the leaves and twigs situated just at, or immediately above, the level of the
grass tend to be damaged first, although they are not necessarily at the apex of the plant.

103. With reference to the treatment of this
type of grassland, therefore, it is obvious that we may either try to reduce radiation which is
chiefly responsible for the failure of Sal to establish itself in areas where the soil is generally
suited for it, as indicated by the prevalence of the grass Saccharum Narenga, or we must
introduce frost-hardy and xerophilous species which are able to thrive under the conditions
of the habitat.

The injurious action of radiation might be modified—
(a) By clearing lines through the surrounding forest and thus improving the circula-
tion of air.
(b) By irrigating and thus keeping the soil moist which would result in increasing
the quantity of water vapour in the air and thus in decreasing radiation.

On the other hand, if these measures are impracticable, the most suitable indigenous
frost-hardy and xerophilous species for afforesting these grasslands is almost certainly the
khair, Acacia Catechu. Another species which would probably be of value in establishing
woodland in such places is Zizyphus Jujuba.

Two exotic species which it is believed might be successful are Cinnamomum Camphora
and Eucalyptus citriodora both of which were observed to successfully survive the excep-
tionally severe winter of 1904-05 in frosty grasslands in this locality.

ie P24

HOLE : SOME INDIAN GRASSES AND THEIR (ECOLOGY. 125

Once these species or some of them have been established, a shrub which it is believed
would be of great value if introduced as an underwood is Adhatoda Vasica. This plant is
xerophilous although a strong shade-bearer and on account of its power of increasing the
humus centent of the soil it would probably help to establish soil moisture conditicns suit:
able for sal and thus facilitate the ultimate introduction of the latter, where this is consi-
dered advisable.

104. As regards those grasslands where frost
damage is not severe it should be remembered that where—

Saccharum Munja is dominant the locality is usually suitable for Dry Miscellaneous
Forest,

Saccharum Narenga is dominant the locality is usually suitable for Sal Forest,

Erianthus Ravenne is dominant the locality is usually suitable for Moist Miscella-
neous Forest,

and that careful protection from fire and grazing will, as a rule, result in naturally replac-
ing these types of grassland by forest of the parallel type. Acacia Catechu has been seen
establishing itself naturally in thickets of Saccharum Munja, Mallotus and Cedrela have
similarly been seen ousting Hrianthus and it is believed that Sal can do the same with
Saccharum Narenga, but artificial sowings would naturally hasten this result.

105. The study of the cecology of Sal has only
recently been commenced at Dehra Dun and owing to the want of necessary staff and appli-
ances it has only been possible to carry out a few preliminary and comparatively rough
experiments for the purpose of determining the factors favourable for the establishment of
Sal reproduction. <A full account of these experiments is beyond the scope of the present
paper and this will be published separately, but it is interesting to note that so far as they
go these experiments show—

(a) That in this iccality a number of sal seedlings die back in September—-Decem-
ber owing to the rapid drying of the soil after the close of the rains. This
injurious action is most marked on sand, less so on stiff loam and least in
garden soil which contained a considerable admixture of humus. There had
been no frost when this dying back was noticed and where the seedlings on
these three classes of soil were regularly watered they did not die back.

(b) That Sal is markedly sensitive to the intensity of light. The leaves are coloured
red when young and even when mature they possess considerable power of
movement frequently turning their edges to intense light and the upper sur-
face of the lamina to diffuse illumination.

That, in this locality, seedlings grown in stiffish loam, similar to that on
which most of the local sal forests are found, develop more successfully [as
judyed independently by (i) the percentage of stems which die back and by
(ii) the air-dried weight of the best stems which survive] when provided with

f 125 }

126 INDIAN FOREST MEMOIRS.

overhead shade and diffuse lateral illumination than when grown in the open

with no shade.
(c) That sal is decidedly sensitive as regards soil-aeration and water-logging proves

rapidly fatal.

These results therefore show that sal is only likely to establish itself on well-drained
soil which is not subject to water-logging and that in this locality any factor which reduces
the humus content of the soil, or which tends to increase the loss of moisture from the soil by
evaporation, tends to retard the establishment of sal reproduction.

The burning of grasslands which it is desired to ultimately afforest with sal therefore
is obviously injurious owing to the destruction of all humus and organic débris while on the
other hand the introduction, where possible, of plants like Kugenia operculata, Adhatoda
and Mallotus into these areas will almost certainly accelerate the establishment of sal
owing to their power of increasing the humus content of the soil, their shade being also
to some extent beneficial.

106. It must of course be understood that the
above remarks apply to the locality more particularly considered in this paper and it is
obvious that the conditions prevailing here and in the Sal forests of Central India are very
different from those of some of the Sal tracts of Bengal and Assam where the tree has
extended into, or close to, the hygrophytic Zone of the tropical evergreen forest. In such
places the dominant factor is frequently not lack of sufficient soil moisture, but the presence
of more vigorous competitors, and it is possible that fires, by reducing the water-content of
the soil, may be beneficial to the Sal by rendering the conditions less favourable for the
vigorous development of its injurious hygrophilous competitors. In these localities, also,
insufficient aeration of the soil may in some cases be responsible for the unsatisfactory
growth of Sal seedlings and here again firing would probably be beneficial.

f 126 J

Y 2 G
: See

S , =
SD ROR
S reassign
Badshahibagh

TOPOGRAPHICAL SYMBOLS.
State Bonndary... 0.

Range Boundary ................-....

Reserved Forest 57.00.0000 | eal G

Railway line with Station ..... ... rrrrmmonrreinns

Main Road ............. sane «SA ase 2
Foot-pathe -----.--0------:eeee ee
Village site ee °
Forest Rest House. -------- oF.R.H,
Pablic Works Dept. -

ublic Works Dept. Bungalow iw pie
Travellers Bungalow... -.......... aT.B,
Principal G T. Station K

36 | With height above M. 5. L. 7699

Compiled in the Forest Map Office by A. Descubes Esq. 0.C. Forest

Major J.M. Burn R,E. O.C. Forest Map Office. Published wi

Reg. No. 79D. April. 1911.—Forest—1 010.

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PLATE 1

Scale } Inch == 4 Miles

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TOPOGRAPHICAL SYMBOLS.
State Boundary...
District Bonndary-.......... Eee

Range Boundary

Reserved Forest ..,,

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with height, Peas
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Compiled in the Forest Map Office b fg. Sur ndia..
yy A. Descubes Esqr. O.C. Forest Map Records. General of I
Major J.M. Burn R,E. 0.C. Forest Map Office. Published under the direction of Colonel §.G, Burrard BAR. F.R.S., Of. Servs4or

Reg. No. 78D, April, 1911.—Forest—lv!0. oad

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Heliozincographed at the Survey
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Explanation of Figures.
Prate II.

Fig.1. Aristida cyanantha the dominant species, with scattered plants of Saccharum
Munja, in dry gravelly bed of torrent near Jhajra which flows southwards
into the Tons River. In the background is seen a line of Saccharum Munja
occupying the sandy deposits on the banks of the Tons River.

Photograph taken 18th October 1909.

, 2. Aristida cyanantha the dominant species, with scattered plants of Saccharum
Munja, showing the Tons River in the background with Saccharum Munja
the dominant species on the sandy deposits on its banks.

Photograph taken 21st October 1909.

PLATE If.

2 pe patty ee wenn oe

ry,
a

AD
ot a ye
ia
6

Explanation of Figures.
JeAGaTe ILILIL,

Fig. 1. in the foreground a portion of the bed of the Tons River below Jhajra. In
the background the Siwalik Hills. Between these lies the Asan River.

The sandy deposits on the banks of these large streams constitute an extensive
savannah of Saccharum Munja.
Photograph taken 18th October 1909.

,, 2. The Asan River below Jhajra. On the water-logged area at the water’s edge
Saccharum spontaneum var. nepalense is dominant. The high well-drained
eround in the background, on the contrary, is held by Saccharum Munja.

Photograph taken 21st October 1909.

ye /NIns A,

Explanation of Figures.
Puiate IV.

Fig.1. Triraphis madagascariensis dominant on sandy bank of stream on edge of Sal
forest near Kansrau.
Photograph taken January 1910.
, 2. Triraphis madagascariensis with Aristida cyanantha in dry gravelly torrent-
bed on edge of Sal forest near Kansrau.
Photograph taken January 1910.
On the right in the foreground are seen some bent and broken twigs of
Orthanthera viminea.

Fig. 2.

Explanation of Figures.
PTA EVE

Fig.1. The Song River above Lachiwala with the Himalayas in the background, show-
ing a patch of Acacia Catechu forest established on an island of boulders and
sand.

On the edges of the forest Saccharum Munja and Saccharum spontaneum (tufted
sand-form) can be seen. Under the Acacia is some undergrowth of Adhatoda
Vasica.

Photograph taken January 1910.

,, 2. The Song River below Doiwala showing Dry Miscellaneous Forest on deposits
of boulders and sand. The forest consists of Bombax malabaricum, Odine
Wodier, Acacia Catechu, Bauhinia racemosa, Dalbergia Sissoo, Grewia
oppositifolia, Moringa pterygosperma (now flowering and conspicuous),
Stereospermum suaveolens, Aigle Marmelos and Cassia Fistula.

In the undergrowth are Calotropis procera, Murraya Koenigu, Zizyphus
Jujuba, Randia dumetorum and Adhatoda Vasica.

On the edge of the forest are Saccharum Munja and Saccharum spontaneum.

Photograph taken January 1910.

ALAMITIS) W/

TG OS,

Explanation of Figures.
PuatTeE VI.

Erianthus Ravenne on shady bank of irrigation canal below Lachiwala.

The water-logged soil at the water’s-edge in the foreground is occupied by Cova
Lachryma-Jobi and Saccharum spontaneum var. nepalense.

On the slightly higher ground behind is Erianthus Ravenne. The trees are Sapium
sebiferum, Trewia nudifiora, Mallotus philippinensis, Salix tetrasperma, Bombazx mala-
baricum, and Cedrela Toona.

Photograph taken 31st October 1909.

WAL

FeAGTs

Explanation of Figures.

Puate VII.

Erianthus Ravenne associated with young trees of Mallotus philippinensis, Cedrela

Toona and Bombax malabaricum near Lachiwala.
Photograph taken 31st October 1909

VII.

PEATE

Explanation of Figures.
Pirate VIII.

Erianthus Ravenne dominant in narrow clearing made on account of the railway line
in Moist Miscellaneous Forest near Lachiwala.

Photograph taken 31st October 1909.

The branches of the younger panicles are still spreading but the majority of the
panicles are fruiting with erect branches. On the left is seen a panicle which has dried
prematurely owing to the culm having been attacked by a larva.

Photograph taken 31st October 1909.

VIII

PIP AGTH E

—

Explanation of Figures.
Prate IX.

Fig.1. Erianthus Ravenne in Moist Miscellaneous Forest near Kansrau. The trees
are Stereospermum suaveolens, Celtis australis, Bombax malabaricum, Ehretia
acuminata and Cedrela Toona.

» 2. Hrianthus Ravenne in Moist Miscellaneous Forest near Kansrau. The trees

are Dalbergia Sissoo, Gmelina arborea, Kydia calycina, Cordia Myxa and
Putranjiva Roxburghi.

Both these photographs taken January 1910.

PLATE IX.

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Explanation of Figures.
PLATE X.

Fig.1. Erianthus Ravenne in Moist Miscellaneous Forest near Kansrau. The trees
are Odina Wodier, Butea frondosa, Mallotus philippinensis, Garuga pinnata,
Gmelina arborea, and Terminalia tomentosa.

3» 2. Hrianthus Ravenne in Moist Miscellaneous Forest near Kansrau. The trees
are Mallotus philippinensis and Cordia Myza.
Both these photographs were taken January 1910.

PLATE

X.

Explanation of Figures.
Prate XI.

Saccharum Narenga the dominant grass in a shady blank in Sal Forest near Jhajra.
Photograph taken 18th October 1909.

|

PLATE

Explanation of Figures.
Piate XII.

Saccharum Narenga the dominant grass in a shady blank in Sal Forest near Jhajra.
Photograph taken 18th October 1909.

Pe alten
shy

Le

Explanation of Figures.
Prate XIII.

Figs. i and 2 flowering panicles, Figs. 3 and 4 fruiting panicles, of the loam-form
of Saccharum spontaneum, as usually found in wet pasture lands. Note the spreading
habit of growth. The grass here is cut annually at the close of the rains.

Photograph taken at Dehra Dun, September 1909.

|
!

Fig. 3. Fig. 4.

Explanation of Figures.
AE Xenye

. The tufted form of Saccharum spontaneum as usually found on sand.
The four plants shown were collected in May 1909 and planted at once in
the Dehra Dun Experimental Garden, where they were photographed in
December 1909.

The two thin clumps on the left had been burnt while the two on the right had
not been burnt.

ho

. A plant of the tufted sand-form of Saccharum spontaneum collected in January

1909 which had been severely damaged and almost killed by a fire in May
1908.

3. Another plant of the tufted sand-form of Saccharum spontaneum.

Fig. 1.

Ut:

Explanation of Figures.
PrAmn Dove

Figs. 1 and 2, plants of the ordinary spreading loam-form of Saccharum spontaneum,
as usually found in wet pastures.
Fig. 3. The Asan River near Jhajra, with the Siwalik Hills in the background.

The water-logged soil in, and close to, the river bed is occupied by Saccharum
spontaneum var. nepalense. The plants show inferior dimensions, as the area
is heavily grazed and the grass here is also cut for thatching purposes.

Photograph taken October 1909.

Explanation of Figures.

Pruate XVI.

Fig. 1. Flowering panicle of Saccharum spontaneum x .

bb)

2. Culm and leaf of same species x 3.

3. Ligule of same x #.

4. Portion of culm of same with axillary shoot, the culm is solid above and
fistular below, (a) the culm node, (b) the leaf insertion ~ 3.

5. Culms of same with an erect habit of growth originating from the nodes of an
older culm. Note the short internodes C, C, at the base of the culms. (a) the
culm node, (b) the leaf insertion x 3.

6. Base of culm of same showing the spreading habit of growth; CC the short
internodes at base of culm x 3.

7. Flowering spikelet of same x 6.

PACA Mav alh

LA

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a NICD ENN DNB LLG AL LAO eae
eR AISI ESAS ~
s —<

Tah,

Explanation of Figures.
Pirate XVII.

Fig. 1. Fruiting panicle of Saccharum spontaneum x 4.
2. Leaf of same x 3.
, 3. Pair of flowering spikelets of same with joint of axis; note the erect callus hairs,
x 3.

, 4. Fruiting sessile spikelet of same with joint of axis and pedicel of pedicelled
spikelet which has fallen off. Note the wide-spreading callus hairs x 3.
5. Flowering spikelet of same x 6.
6. Glume I of spikelet ]
Setedeathi.) ovitvamer lA
Sh iat nied !
Oe ie CLA ‘5
,, 10. Pale x 6.
,, 11. Lodicules
,, 12. Stamens
,, 13. Ovary
,. 14. Ovary stamens and lodicules

PLATE XVII.

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Explanation of Figures.

Prate XVIII.

Fig. 1. Fruiting panicle of Saccharum spontaneum var. nepalense x 3.

2. Leaf of same x 3.

,, 3. Pedicelled spikelet of same x 24.

,, 4. Sessile spikelet of same with pedicel, from which the pedicelled spikelet has

33

fallen off, and joint of axis x 24.

, 9. Basal portion of culm of same with axillary shoot. Note that culm is

fistular x 3.

,, 6. Upper portion of culm of same. Note that culm ig solid x 3.

Figs.

ke
(

and 8. Culm of sand-form of Saccharum spontaneum showing the efiect
of cutting off the leaves on the length of the internodes. The culm was cut
over at X and thus deprived of all its green leaves which were visible on the
date of cutting. When the culm was drawn however the continued growth of
the apical bud of the culm and of the leaf rudiments which had been uninjured
by the cutting had produced a new terminal tuft of green leaves. This cutting
resulted in the removal of the entire lamina and more or less of the sheath
of 3 leaves numbered 1, 2 and 3. In figure 8 the sheaths of these leaves have
been removed to show their insertions on the culm which are also numbered
1,2and3. Internode (a) had practically completed 1ts growth on the date of
cutting and is of normal length. It will be seen however that internode (b)
has not increased in length since the date of cutting, as is indicated by the
cut apices of the leaf sheaths 1, 2 and 3 having remained at the same level X.
Compare the length of the abnormally short internode (b) with that of the
normal internode (a). C is a leaf of which only the apical portion of the
lamina was destroyed by the cutting.

PLATE XVIII.

Explanation of Figures.
Piate XIX.

Flowering plant of Saccharum Munja.
Photograph taken on the banks of the Tons River near Jhajra on 2ist October 1909.

ATI

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f ath ‘Bap 4 0% : ~
pea Si ks

Explanation of Figures.

PLaTE XX.

Glume 1 of sessile spikelet of same

» 2 0» >» >»
» 3 r» 03 %»
ye: % >» 0
Pale e 0 03
Anthers 3 25 .s
Lodicules _,, ss r»
Glume I of pedicelled spikelet of same
ee Ul x 6 r

. Flowering panicle of Saccharum Munja collected October 1909.
. Three fruiting panicles of Saccharum Munja collected November 1909.
. Pair of spikelets of Saccharum Munja with joint of axis
. Pedicelled spikelet of same

. Sessile spikelet of same

i
|
|

VV

PLATE

i
4
i ;
~
j
= a *
hes 4c
Siaee 4a a is .
r ‘ ¥ oh
ue ° =
’ Ba) ; 1
: ahh
' 1 f ; i‘
1 , LA yy ‘
2) ay ery. x “6 : 5; ;
3 eo if : + i
Sel ae.
i F ie i ‘ i : fin 4
Fi Vi
y B's
ioe . b,
- Noon : ne i, "8 ' = 7 ' ae a
of) + p 7 ‘ > mF eee MP lf oF + ‘
Ree a a | AS

ee ee nk ke
" 7% "= i _ at a rf |

a oie f il 1 | | |
, 7 " i

i a ce
‘

pull

oe

Explanation of Figures.

PLATE X XI.

. Culm of Saccharum Munja. (a) leaf insertion, (b) culm node, (c) axillary bud

x 2,

. Leaf of same showing ligule x 3.
. Base of lamina and upper part of sheath of same, showing ligule and concava.

convex midrib x 3.

. Immature one-year culm of same with the basal leaves removed to show the

short basal internodes. There are 13 short internodes of which the upper
internode is still immature x 3.

. Base of mature flowering culm of same. There are 11 short basal internodes

3
Xx 4

. Base of flowering culm of same with 13 short basal internodes and in addition

to these a second series of short internodes indicating that a change has taken
place in the soil level x 3.

. Portion of rhizome of same showing the aerial culms developing from the

terminal bud of the rhizome. The basal leaves have been removed to show
the short internodes x 3.

PLATE AX,

Explanation of Figures.
PuatTE XXII.

. Saccharum arundinaceum growing in a garden at Dehra Dun.
Photograph taken November 1909.

2. Pair of spikelets of same with joint of axis)

3. Sessile spikelet of same with portion of

joint of axis

. Pedicelled spikelet of same

Glume 1 of sessile spikelet of same

99
33

+9

Pale

Lodicules

Stamens

Ovary

Glume I of pedicelled spikelet of same

be)

II

39

be)

be)

33

LB)

99

be)

33

bb)

393

33

33

Sell Se.

XXII.

PLATE

Pd 0 Aan
ee}

No

Explanation of Figures.
Puate XXIII.

Flowering panicle of Saccharum Narenga Wall. Collected October 1909.

Four fruiting panicles of S. Narenga all collected from the same plant on 20th
November 1909. The two on left of staff and the outer one on the right are-
normal. The panicle next the staff on the right was caused to dry prematurely
on the plant by bending the flowering culm. Note the tenacious panicle
branches.

Two abnormal fruiting panicles of S. Narenga caused by the culms being
damaged by insects and fungi. Note the tenacious panicle branches.
Collected November 1909.

A group of three plants of S. Narenga in Dehra Dun Experimental Garden.
The fragile branches of the fruiting panicles have broken up leaving only
the persistent, spike-like, primary rachis.

Photograph taken December 9th, 1909.

XII.

PLATE

NSS
A)

LA hb

ig LY, VU, yy AY | MAA é

ie, tle

Fig. 4.

Fig. 3.

=

ie ; ?

it, a bo ,
aA oe Nias tas

%

Explanation of Figures.
PLATE XXIV.

Figs. 1—4. Culms of Saccharum Narenga which have become fistular owing to an in-
sect- or fungus-attack at the leaf-insertion. In Figs. 1 and 2 the base of the
leaf-sheath has not been removed, in Fig. 3 the base of the leaf-sheath has
been removed from its insertion showing an axillary bud and the culm-node
immediately above it. In Fig. 4 also the base of the leaf-sheath has been
removed from its insertion and a section cut through the centre of the culm
showing the hollows caused by the attack.

Fig.5. Leaf of Saccharum Narenga.

,, 6. Top of leaf-sheath and base of lamina showing the section of the lamina and
the hgule of Saccharum Narenga.

Figs. 7—8. The uppermost leaf of the flowering culm of Saccharum Narenga.

Fig. 9. Portion of culm of Saccharum Narenga showing the smooth culm and the hairy
leaf-sheath and leaf-insertion.

All figures x 3.

PLATE XXIV.

pea
?
=
-
S =
: aS
+
it
j
ie
4

i
fl i
Diet| 1a
i ‘ie :
ian Me

ms "hubs 7

Fi
a
rr \e ,
oon
a :
br
+
:
3 ”
+
he)
}e

et) ) aes
aart
» AEA:

Figs. I—IV.

V.
VI.

13.

14.

15.

Explanation of Figures.

PLATE XXV.

Glumes I, IT, III and IV of pedicelled spikelet of Saccharum Narenga
x 6.
vie f pedicelled spik
uadionles io pedicelled spikelet of same x 6.
. Glumes 1, 2, 3 and 4 of sessile spikelet of same x 6.
ne tot sessile spikelet of same x 6.
. Stamen oe
. Ovary and Stigmas 5 peices:
. Sessile and pedicelled spikelet of same species with joint of axis x 6.
. Pedicelled spikelet of same x 6.
Saccharum fuscum Roxb., copied from Roxburgh’s drawings, Vol. II,

No. 73. Note the distinct pedicels of both spikelets of each pair.

. Portion of rhizome of Saccharum Narenga showing the aerial culms.

developing from the terminal bud of the rhizome and as axillary shoots
from the basal nodes of older culms. The basal leaves have been
removed to show the internodes x 1/1.

One-year culm of Saccharum Narenga with the basal leaves removed to-
show the short basal internodes. There are 13 internodes of which the
upper three, (a)—(a), are still soft and immature, (b), (b) scars of roots.
which have been cut off, (c) an axillary bud, (d) scar from which an
axillary bud has been removed x 1/1.

Another one-year culm of the same species. There are 12 short basal
internodes of which the upper two, (a), (a), are still soft and imma-
ture. (b), (b) root scars, (c), (c) axillary buds x 1/1.

Portion of rhizome of Saccharum Narenga (a), (a), young buds from
which the scale-like outer leaves have been removed. (b) axillary bud
from which the basal 4 scale-leaves have been removed showing the
next two leaves with small lamina.

(c) immature one-year culm with 12 short basal internodes the upper two
of which are still soft and immature.

(d) mature flowering culm with 8 short basal internodes x 1/1.

PLATE XXV.

Ve

if
afer

oA5

es if
yy

net Lisi

+

Se ok dail a

Explanation of Figures.
PLATE XXVI.

Fig.1. Flowering panicle of Erianthus Ravenna, collected October 1909.
,, 2. Two fruiting panicles of Erianthus Ravenne, collected November 1909.
,, 8. Group of the same species growing in Dehra Dun Experimental Garden.
Photograph taken October 1909.
,, 4. Pair of spikelets of same with joint of axis
1,1, Glume 1 of sessile spikelet of same

Dh OX fe ae i a ss

3}, 83, atte es ws a

4,4, » of - sf . all x 44.
P Pale
S Stamens

O Ovary showing stigmas
L_ Lodicules

XXVI.

PLATE

> in he or ; . :
| a7 VRS ara ; =
= at: . By - a . re = ‘ 7
. Uy rei) Patra Wien) (2) Sera ee Ryn aia a Pena ke | a
7 cc 0 ed | Aled HH YALA Di a wil 1h bak ; ” a
. oe Peak | ei Lr pate (SS), a
Bee ose i put) poiewol) Vp Tiel energy UE of: Se
Betray tReet ed bp q7y fs f, Bite Vi Vs nit D> : h PH) q : _
a ‘es Peo; : + : ,
| oliall hig | 4 oo
oe Raa Tt, | 7

2 ‘ it h " ee ce Re oa

Bee TNAh Ee 10 OC fone as

od

Explanation of Figures.
Puate XXVIII.

Fig. 1. Culm of Hrianthus Ravenna, (a), \eaf-insertion, (b), culm-node, (c), axillary
bud, (d), base of hairy leaf-sheath.
, 2. Lower leaf of same.
,, 3. Uppermost leaf of flowering culm of same.
Figs. 4—5. Base of lamina and top of leaf-sheath of same showing the concave midrib
and ligule.
All figures x #.

PLATE XXVII.

Ue Rae EA ES Eat ERATOR LALO REPO

ounces toa
Sse Sale

~ak

eee FEST ES ATR — - eae
—— ca ila a a me en ao

Se REE

OL LP ee I ez

= See

nS ee

Explanation of Figures.
Piate XXVIII.

. Flowering panicle of savannah form of Imperata arundinacea.

. Fruiting panicle of same.

». Depauperate culm of Imperata arundinacea collected from an area where the
grass is annually cut at the close of the rains and which is then frequently
grazed over, (a), (a), the hearded leaf-insertions, (b), (b), the long villi at base
of lamina.

. Ligule of plant shown in Fig. 3.

Figs. 1—4 all’ x2.

. Pair of spikelets of savannah form of Imperata arundinacea x 24.

. Spikelet of same showing callus hairs x 3.

. Callus hairs of same x 3.

, Glume ILof same

’ Il 32>
TE
oY i be
, Pale of same
Stamens ,,

Ovary:

PLATE XXVIII.

-

by + ! u } ee \ .
SG] A AN eT

Explanation of Figures.

Piatt XXIX.
Fig.1. Leaf of savannah form of I[mperata arundinacea.
, 2. Base of lamina and apex of sheath of same showing ligule.
,, 3. (a) Stolon of same covered with membranous sheaths. (b) young aerial culm

developing from terminal bud of stolon (a). (c), (c), stolon branches devel-
oped from the nodes of stclon (a).

,, 4. Shows the same but with the sheaths removed to show the internodes. Lettering
as in Fig. 3.

, 9. Base of mature flowering-culm of same showing 14 short internodes at the
base (a)—(a). 58, 8, stolons.

,, 6. Base of mature flowering-culm of same showing 11 short internodes at the base
(a)—(a); 58, 8, stolons.

», ¢ Culms of same showing transition from glabrous to densely bearded leaf-
insertions.

All figures x 3.

PLATE XXIX.

aaa diced eel

Explanation of Figures.

PLATE XXX.

. Three plants of Imperata arundinacea growing in Dehra Dun Experimental

Garden. Note the young leafy culms in the foreground springing from the
wide-spreading subterranean stolons.
Photograph taken October 1909.

. Another group of the same in Dehra Dun Experimental Garden. The culm

on the right which originated from a subterranean stolon in May 1909 has
produced a flowering panicle.
Photograph taken December 1909.

. Panicles of the same. The three central panicles were collected from a grass

lawn where the plants had been dwarfed by grass-cutting. The outermost
panicle on the right and the two outermost panicles on the left belong to tke
swamp-form and were collected in a marsh. The remaining three panicles
belong to the ordinary savannah form of the species.

Fig.

F F
yy
.
.
t
fr
MpaieN\
|
4
“Seen: Yi
.
raf
ray ;
in
7
“
ii
_ i
i =
z i
.
_ "
i
= . *
‘
i ¥ . {
~
.
‘
, t
7)
i
.
E _
i ‘
eS i
1
= =
¥ ‘
LE ee : . rs a H .

Explanation of Figures.
PLATE XXXII.

Fig.1. Flowering panicle of Triraphis madagascariensis.
Photograph taken September 1909.

, 2. Fruiting panicles of T'riraphis madagascariensis.
Photograph taken November 1909.

PLATE KXXl

Explanation of Figures.
PiaTE XXXII.

Fig. 1. Triraphis madagascariensis growing in Dehra Dun Experimental Garden.
Photograph taken September 1909.
, 2. Spikelets of same x 3.

,, 38. One spikelet of same x 6 approx. showing the rachilla jointed and bearded he-
tween the flowering glumes.

, 4. One flowering glume of same embracing the pale
I, I, Glume_ Toff same |

IU. ane, B 1B Dee

III, III, Flowering glume of same all ce
P’ Pale of same
L__Lodicules
S Anthers
O Ovary

a9

PLATE XXXII.

=. - =a

Explanation of Figures.
Prats XXXIIT.

Fig.1. Culm of Triraphis madagascariensis.
,, 2. Ditto showing leaves.
, 3. Base of lamina and apex of leaf-sheath of same showing ligule.
, 4. Base of mature flowering culm of same showing the short internodes and
numerous large buds at base.
, 9. The same with the roots cut away to show the short basal internodes more

clearly. In this case there are 13 short basal internodes.
All figures x 3.

PLATE XXXII].

Explanation of Figures.
Piare XXXIV.

Aristida cyanantha in gravelly stream bed near Jhajra.
Photograph taken 21st October 1909.

/

Explanation of Figures.
EAE XOXOXO A

Fig. 1. Flowering panicle of Aristida cyanantha x 3.
2. Flowering spikelet of same x 1 /1. Note the erect adpressed awns.
» 93. Fruiting spikelet of same x 1z. Note the 3 spreading awns.
Figs.4—5. Spikelets of same attacked by Ustilago aristide-cyananthe. Note the re-
markable pod-like structures x 3

”

PLATE XXXV,

SAS “

So

4
+
Hk
to :
= Ga i ‘ aq
a i
.
5)
} 4
a, . :
i
i
H
: E
_ L*¥ '
i
- i f pi
- a » Lis ¥ . 7
:
i
; '
cies } AL a .
j “35
= : ; |
| eu : -
, i
2 + - .
u ' i ! ;
[! is 7 |
. ~ ¥ u .
{
: i bea t =, ae
; = 5 5 i rd :
Obs =F i 7 |
4 4 if I
, eel
reel
7 es
; _
bt: }

Sou

Explanation of Figures.
PLATE XXXVI.

Fig.1. Culm of Aristida cyanantha (a) leaf-sheath from which the lamina has been
shed, (b) leaf-insertion, (c) culm-node, (d) axillary shoot with scaly leaves at
base x 3. .
2. Lamina of same showing ligule at (a) x 3.

., 3. (a) Mature flowering culm of same which has developed as an axillary shoot
from the basal node of an older culm (b). (c) is another axillary shoot spring-
ing from (b) which at present is merely a large bud. There are 9 short inter-
nodes at the base of (b) and 6 short internodes at the base of (a) x 3.

4. Spikelet of same x 1/1
I, I, Glume I of same
1 awed Ese Mere 1
TREE hs ee DA a

P Pale : all x 3.

L _ Lodicules a

S Anther

QO Ovary )

PLATE XXXVI.

,
‘
4 i
?
5 Ps
ty i
} ' Ky
a 3 i
\ ‘
( ”
i
)
* R -
‘ j
t
1
{
x ? "
Fe . i
4 ; i
! c
’
no
,
) : FAA
raed ee Oba
i Ay
i (eo
ei ‘ ; i < F ing Uae Yoetan
7 I i 4 ih A
: t
iM
a i f Me
i } ! ~ }
n a :
iz , a I r 1
f - ' i
b i i x
i i
i ‘ TP i : i
aoe } Fart ie ke) : :
Rat D : i : 2 y vs 1 Bain al Oy, oe

cans

i
s
i=
.
3
{
‘
5
‘
i
.
*
N; 7
od = =. s
= 4 A
i
>
1
bal J
1
2 oe

Explanation of Figures.
PLATE XXXVI.

Fig. 1. Andropogon monticola var. robustus growing in Dehra Dun Experimental
Garden.

Photograph taken October 1909. This is a robust plant which shows little to
no hair on the basal 2rds of Glume II of sessile spikelet and which
approaches var. 7'rinw very closely.

» 2. Another less robust plant of the same variety also growing in Dehra Dun
Experimental Garden.

Photograph taken September 1909. This plant showed the middle third of
Glume IT of sessile spikelet pectinately white-ciliate and was taken
from a dry sandy soil in a locality which is annually fired.

PI
-A
ly
E
X
XX
XV
“all

ee)
fave

jt
t

Explanation of Figures.
PiaTE XXXVITI.

Fig. 1. Flowering panicle of Andropogon monticola. Note the spreading branches
x 2.
,, 2. Fruiting panicle of same. Note the erect branches x 3.
,, 3. Panicle of same after the fall of the spikelets showing the bearded tips of the
branches x 3
Figs.4 & 5. A single cluster of spikelets of same showing the central ¢ and two lateral
é spikelets x 24.
Fig.6. Base of lamina and top of sheath of same showing the ligule x 1/1.
,, ¢. Base of culm of same showing 13 short basal internodes, from which numerous
buds are developing which will produce next year’s culms x #.
,, 8. Culm of same showing the leaves and flowering axillary branches x 3.

PLATE XXXVIII.

.
|

=

ety

Ai)

ms Wi
Tel

Fig. 1.

ea:

acd
i, Te
Tu, Tat,
in, IBY,
eae
2, 2,
aa)
4, 4,

S

O

iL

Explanation of Figures.
Pratt XXXIX.

Culm of Andropogon monticola var. robustus showing the fan-like arrangement
of the branches.
Sessile ¢ spikelet of same
Pedicelled g spikelet of same
Glume fof sessile spikelet of same
c. II 5 2»
oy SET 4 ‘ 53
en elaVy 5) 23 2»
3 1 pedicelled _,, ss
2» 2 %» %» >»
%» 3 2» 2) %»
> a 2» ¥9 %»
Anthers
Ovary
Lodicules

all x 44.

Aa Fs

Al

ZZ)
|

PEIN WO

q
>

~

Ki

Explanation of Figures.
PrLatE XL.

Gola Tappar, a grassland, which is believed to have originated from a clearing being
made in Sal Forest for purposes of cultivation. Note the dense wall of Sal Forest surround-
ing the grassland; the Himalayas are seen in the distance.

This grassland is annually fired, but is closed to grazing. The Sal-locality-indicating
erass Saccharum Narenga is common in this area, but note the inferior dimensions attained
as compared with those shown in Plates XI and XII. In parts of this area, also, the
xerophilous Andropogon contortus is dominant. These characteristics indicate a drying
up of the soil, which has been probably caused partly by the surrounding forest interfering
with the free circulation of air currents and thus causing intense insolation, but which has
undoubtedly been accentuated by the annual fires.

Photograph taken January 1910.

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