Bamboo Leaves

Leaves and sheaths

In most woody bamboos, rhizome bracts, culm sheaths and foliage leaves
represent 3 different well distinguished forms of the monocot leaf. Rhizome
bracts and culm sheaths are mainly protective and supportive, while foliage
leaf blades are responsible for photosynthesis. Culm sheaths and their blades
are very important for species identification, those from the lower mid-culm
being most distinctive. Foliage sheaths and blades are also important, but
are rather more variable. Important parts for identification include ligules,
auricles, oral setae, and blades. The sheathing section itself varies in density,
nature, pattern and colour of hairs, and cilia along the margins. The
persistence of the culm sheaths is also important, and they may be more
persistent in the centre so that they hang for a while before falling
completely.

Leaves as sheathing organs with blades

Each internode throughout the bamboo plant is surrounded by a tubular
sheath, basally attached to the node, terminating in a blade. As in all
grasses, these are all technically leaves, a confusing concept when applied
in the bamboos, where the sheaths and blades are much more differentiated
and specialized. Leaves are by definition flat organs capable of
photosynthesis, yet the culm sheath may be tubular and not even green.
Therefore the term leaf is best avoided away from the foliage, with the
flattened apical section of all leaves being called a blade, whether leafy or
not. Sheaths and their blades are adapted for different functions according
to the location of the internodes they surround, and in woody bamboos they
are generally referred to as either rhizome bracts, culm sheaths or foliage
leaves.
The rhizome bract is purely protective in function, tightly and permanently
wrapped around the rhizome axis. The culm sheath is both protective and
also thickened so that it can support the soft, weak elongating culm
internode. It is usually deciduous, revealing the normally green culm
internode inside, which can then also photosynthesize. The foliage leaves
have a protective and supportive sheath in which more distal blades are
wrapped, and a very much larger photosynthetic blade.
Culm sheaths are the most important part of a bamboo for identifying
different species. The variations in shape, hairs, marginal cilia, and the
characteristics of various apical appendages are most evident when the
sheaths are young, and bamboos can be very difficult to identify properly
without them. They vary substantially along the culm, and it is important to
look at those in which the characters are best developed. Ignore those that
look like leaves from the upper part of the culm, and those that look too
much like rhizome bracts from the very base. Those at the base have few
appendages, while those at the apex are not only more similar to leaf
sheaths, usually being narrower, but they also have atypically stronger
blade development, the terminal culm sheath blades often being
considerably larger than even any foliage leaf blade. To standardise for
descriptions, culm sheaths from about one quarter the height of the culm
above the ground are most commonly examined and cited, as the important
characteristics are expressed most comprehensively at that point along the
culm. In the larger bamboos, that height is fortunately also much more
accessible, and culm sheaths from eye-level can be used. A good bamboo
description normally covers the culm sheath and foliage leaf details in
depth, listing the characteristics for each separately.
The importance of inspecting the least leafy of the culm sheaths, rather
than those with well-developed blades is another reason why the term culm
leaf, widely used in other grasses, should not be used in bamboo

descriptions. It leads to a search for a leafy culm sheath, which will not have
the correct features.
Ligules
At the apex of each culm sheath or leaf sheath, where the blade is attached,
a thin membrane is found extending upwards along the apical margin,
pressed against the underlying culm internode, culm sheath or foliage leaf.
This ligule, literally a little tongue, varies in height, lateral breadth, hairiness,
edge shape in terms of serration, and edge ciliation, and these
characteristics are very important to distinguish species.
On the outside of the leaf sheath apex are one or two small less well
developed flaps of tissue, usually known as the exterior ligule, sometimes
called callus. This character is not always described as it varies less between
closely related species, but the ciliation on their margins can be important.
Very rare on the culm sheath.
Blades
The distal part of culm sheath and leaf sheath beyond the ligule is the blade,
shorter and thicker in culm sheaths, thinner, longer and more flattened in
leaf sheaths. The lower side of a leaf is called the abaxialside as it faces
away from the culm when growing upright and parallel to the culm. The
upper side is called adaxial, as it would face the culm instead. Overall shape
varies substantially in the culm sheath, less in foliage leaf blades. Surfaces
vary in hairiness. Venation is variable in leaf sheath blades, especially
presence of transverse veinlets.
Vein patterns can be different, notable is the presence of transverse veinlets
in temperate species. A large longitudinal midrib runs up the centre of the
blade, and is accompanied by two to several dominant parallel veins on each
side, with minor parallel veins between them. Reinforcing these parallel veins
in temperate bamboos are short, minor, transverse veins crossing between
the finer longitudinal veins. Together the longitudinal and transverse veins
produce a reticulate network of tessellate venation. The standard shape and
dimensions for a bamboo foliage blade are linear-lanceolate with length c. 10
times the width, and a shortly acuminate tip. Relative to this they can be
broad or narrow, more linear or more lanceolate, and the tip can be acute as
in Fargesia
nitida to
long-acuminate
as
inCephalostachyum and Neomicrocalamus. The blade is joined to the leaf
sheath by a narrow cylindrical to flattened neck, the pseudopetiole, which
can bend and twist, to make the blade turn to face the light. It is often waxy
or pubescent on one or both surfaces. I do not use the term petiole, as I
consider the familiar petiole of a dicot leaf is probably homologous to the
entire sheath in a woody bamboo.

The pseudopetiole in woody bamboos is a distinctive and diagnostic

character. It can facilitate re-orientation of leaf blades to variable light
sources in the forest understorey that change as tree canopies develop
through the year. It would seem likely that it evolved in primitive grasses and
has been retained in bamboos, while most other grasses, which have
evolved out of the forest understorey, have lost this ability to re-orientate
their leaf blades.
Auricles and oral setae
Where the blade and sheath
connect, lateral projecting lobes
may be found on each side.
These
are
called
auricles,
meaning literally small ears.
Their size and shape are very
good characters for separating
bamboo
species,
especially
those on the culm sheaths,
while on leaf sheaths they can
be more variable.
Projecting bristles, borne on the auricles if any auricles are present, may also
be seen. These are the oral setae, meaning literally mouth bristles, a term
adopted from other grasses, where the circular apex of the sheath is
sometimes referred to as the mouth.
Origins of bamboo auricles and oral setae
Woody bamboos differ from other grasses in the manner of connection of the
blade to the sheath in their foliage leaves. In the bamboos, these are joined
only by a narrow, articulating pseudopetiole. How this junction of the foliage
leaf blade relates to the connection of blades to sheaths in other grasses is
best understood by looking at the culm sheaths of a large tropical bamboo,
in which the development into an efficient photosynthetic organ is secondary
to the need to support and protect the massive but vulnerably soft,
elongating culm. In many of these bamboos the blade is often as broad as
the sheath itself, as in other grasses, with three often only partially
separated sections. These are the larger, central blade, and two smaller,
lateral, ear-like lobes, called auricles. It is hypothesised here
thattogether these correspond to the leaf blade of other grasses, as
evidenced by the parallel leaf veins that travel from the sheath into both
blade and auricles. These largely vestigial veins cross the auricles and are

often extended beyond the margin, forming the stiff bristles known as oral
setae, which can be supplemented by marginal ciliation or fimbriation.
In many other woody bamboos the three sections of the blade are more
separated in the culm sheaths as well as in the leaf sheaths, and less
functional. In many species the two lateral sections, or auricles, are lacking
altogether. Their sporadic occurrence or reoccurrence is typical
of vestigial reduced organs. It is
only when they are exaggeratedly
large, as in species such
as Ampelocalamus
scandens or Yushania
brevipaniculata, that their true
identity, as hypothesised here,
becomes apparent. The spreading
oral setae can then readily be seen
as a kind of leaf skeleton, as
produced by the decomposition of
a fallen dicot leaf, except that the
predominantly lateral venation of
these monocotyledonous leaves
gives a fringed effect rather than a
reticulate pattern.
The remnants of the much finer
transverse leaf veins may even be
evident in species with tessellate
venation, in the scabrid nature of
the oral setae. There are even
species in which very elongated
leaf sheath auricles bear a strong
resemblance to the pseudopetiole,
eg Sarocalamus racemosus, Fargesia dracocephala. Nevertheless, the origin
of the woody bamboo auricle as a reduced sections of a lobed leaf blade has
not apparently been suggested before. It would suggest a primitive bamboo
ancestor with a leaf blade having three sections and two divisions, possibly
more like those seen in blades of Monstera deliciosa (Swiss Cheese Plant)
rather than than the three lobes with prominent midribs as seen in palms
or Arisaema triphyllum(Jack in the Pulpit). No such plants in the Poaceae are
of course extant today. The adaptive advantage of loss of the lamina of the
lateral lobes would be the efficiency gained by more separated rotating
blades, as the sections would shade each other whenever a pseudopetiole
rotated a blade to intercept the light.
Comparison with auricles in other grasses

In other grasses the sheath of some species have lateral naked membranous
projections below or beside the ligule, which are also called auricles. Such
appendages are seen in some bamboo species as well, where they are borne
on the shoulders of the sheath apex, away from the blade. The fact that such
auricles are not homologous with bamboo auricles is shown by the existence
of both types of auricle together in Dendrocalamus hookeri. The culm
sheaths of this species often bear an ovate auricle with oral setae attached
to the blade margin, and a second, more acute, naked auricle on the
shoulder of the sheath, and these two distinct forms of auricle overlapping
side by side to give a partially double auricle.
Length vs width vs height vs breadth
As the acute auricles of other grasses usually extend further laterally rather
than in the same orientation as the blade, their length is by convention given
as their lateral extent, while their height is referred to as their width. On the
other hand, as acute auricles in the bamboos are more parallel to the
longitudinal axis of the sheath than the lateral axis, their length is usually
given as the longitudinal dimension, and their width is the lateral dimension.
This would be supported by their origin as a lateral strip of the blade, rather
than a projection of the sheath instead, as in other grasses. However as the
length of anything is by definition its longest dimension, it is best to refer to
the longitudinal dimension as the height, and to the lateral dimension as
lateral breadth. If in doubt as to what is meant, the best solution is a picture.
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