Leaf

A leaf (plural leaves) is the principal

lateral appendage of the vascular plant
stem,[1] usually borne above ground and
specialized for photosynthesis. The
leaves and stem together form the
shoot.[2] Leaves are collectively referred
to as foliage, as in "autumn foliage".[3][4]
In most leaves, the primary
photosynthetic tissue, the palisade
mesophyll, is located on the upper side
of the blade or lamina of the leaf[1] but in
some species, including the mature
foliage of Eucalyptus,[5] palisade
mesophyll is present on both sides and
the leaves are said to be isobilateral.
Most leaves are flattened and have
distinct upper (adaxial) and lower
(abaxial) surfaces that differ in color,
hairiness, the number of stomata (pores
that intake and output gases), the
amount and structure of epicuticular wax
and other features. Leaves are mostly
green in color due to the presence of a
compound called chlorophyll that is
essential for photosynthesis as it
absorbs light energy from the sun. A leaf
with white patches or edges is called a
variegated leaf.
The diversity of leaves

Leaf of Tilia tomentosa (Silver lime tree)

Diagram of a simple leaf.
1 Apex • 2 Midvein (Primary vein) • 3 Secondary
vein. • 4 Lamina. • 5 Leaf margin • 6 Petiole • 7 Bud •
8 Stem

Top and right: staghorn sumac, Rhus typhina

(compound leaf)
Bottom: skunk cabbage, Symplocarpus foetidus
(simple leaf)
(simple leaf)
1. Apex
2. Primary vein
3. Secondary vein
4. Lamina
5. Leaf margin
6. Petiole

Leaves can have many different shapes,

sizes, and textures. The broad, flat leaves
with complex venation of flowering
plants are known as megaphylls and the
species that bear them, the majority, as
broad-leaved or megaphyllous plants. In
the clubmosses, with different
evolutionary origins, the leaves are
simple (with only a single vein) and are
known as microphylls.[6] Some leaves,
such as bulb scales, are not above
ground. In many aquatic species, the
leaves are submerged in water.
Succulent plants often have thick juicy
leaves, but some leaves are without
major photosynthetic function and may
be dead at maturity, as in some
cataphylls and spines. Furthermore,
several kinds of leaf-like structures found
in vascular plants are not totally
homologous with them. Examples
include flattened plant stems called
phylloclades and cladodes, and flattened
leaf stems called phyllodes which differ
from leaves both in their structure and
origin.[4][7] Some structures of non-
vascular plants look and function much
like leaves. Examples include the phyllids
of mosses and liverworts.

General characteristics

Play media
3D rendering of a computed tomography scan of a
leaf

Leaves are the most important organs of

most vascular plants.[8] Green plants are
autotrophic, meaning that they do not
obtain food from other living things but
instead create their own food by
photosynthesis. They capture the energy
in sunlight and use it to make simple
sugars, such as glucose and sucrose,
from carbon dioxide and water. The
sugars are then stored as starch, further
processed by chemical synthesis into
more complex organic molecules such
as proteins or cellulose, the basic
structural material in plant cell walls, or
metabolized by cellular respiration to
provide chemical energy to run cellular
processes. The leaves draw water from
the ground in the transpiration stream
through a vascular conducting system
known as xylem and obtain carbon
dioxide from the atmosphere by diffusion
through openings called stomata in the
outer covering layer of the leaf
(epidermis), while leaves are orientated
to maximize their exposure to sunlight.
Once sugar has been synthesized, it
needs to be transported to areas of
active growth such as the plant shoots
and roots. Vascular plants transport
sucrose in a special tissue called the
phloem. The phloem and xylem are
parallel to each other, but the transport of
materials is usually in opposite
directions. Within the leaf these vascular
systems branch (ramify) to form veins
which supply as much of the leaf as
possible, ensuring that cells carrying out
photosynthesis are close to the
transportation system.[9]
Typically leaves are broad, flat and thin
(dorsiventrally flattened), thereby
maximising the surface area directly
exposed to light and enabling the light to
penetrate the tissues and reach the
chloroplasts, thus promoting
photosynthesis. They are arranged on the
plant so as to expose their surfaces to
light as efficiently as possible without
shading each other, but there are many
exceptions and complications. For
instance, plants adapted to windy
conditions may have pendent leaves,
such as in many willows and eucalypts.
The flat, or laminar, shape also
maximizes thermal contact with the
surrounding air, promoting cooling.
Functionally, in addition to carrying out
photosynthesis, the leaf is the principal
site of transpiration, providing the energy
required to draw the transpiration stream
up from the roots, and guttation.

Many gymnosperms have thin needle-like

or scale-like leaves that can be
advantageous in cold climates with
frequent snow and frost.[10] These are
interpreted as reduced from
megaphyllous leaves of their Devonian
ancestors.[6] Some leaf forms are
adapted to modulate the amount of light
they absorb to avoid or mitigate
excessive heat, ultraviolet damage, or
desiccation, or to sacrifice light-
absorption efficiency in favor of
protection from herbivory. For xerophytes
the major constraint is not light flux or
intensity, but drought.[11] Some window
plants such as Fenestraria species and
some Haworthia species such as
Haworthia tesselata and Haworthia
truncata are examples of xerophytes.[12]
and Bulbine mesembryanthemoides.[13]

Leaves also function to store chemical

energy and water (especially in
succulents) and may become specialized
organs serving other functions, such as
tendrils of peas and other legumes, the
protective spines of cacti and the insect
traps in carnivorous plants such as
Nepenthes and Sarracenia.[14] Leaves are
the fundamental structural units from
which cones are constructed in
gymnosperms (each cone scale is a
modified megaphyll leaf known as a
sporophyll)[6]:408 and from which flowers
are constructed in flowering plants.[6]:445

Vein skeleton of a leaf. Veins contain lignin that

make them harder to degrade for microorganisms.

The internal organization of most kinds

of leaves has evolved to maximize
exposure of the photosynthetic
organelles, the chloroplasts, to light and
to increase the absorption of carbon
dioxide while at the same time
controlling water loss. Their surfaces are
waterproofed by the plant cuticle and gas
exchange between the mesophyll cells
and the atmosphere is controlled by
minute (length and width measured in
tens of µm) openings called stomata
which open or close to regulate the rate
exchange of carbon dioxide, oxygen, and
water vapor into and out of the internal
intercellular space system. Stomatal
opening is controlled by the turgor
pressure in a pair of guard cells that
surround the stomatal aperture. In any
square centimeter of a plant leaf, there
may be from 1,000 to 100,000
stomata.[15]

Near the ground these Eucalyptus saplings have

juvenile dorsiventral foliage from the previous year,
but this season their newly sprouting foliage is
isobilateral, like the mature foliage on the adult
trees above

The shape and structure of leaves vary

considerably from species to species of
plant, depending largely on their
adaptation to climate and available light,
but also to other factors such as grazing
animals (such as deer), available
nutrients, and ecological competition
from other plants. Considerable changes
in leaf type occur within species, too, for
example as a plant matures; as a case in
point Eucalyptus species commonly have
isobilateral, pendent leaves when mature
and dominating their neighbors; however,
such trees tend to have erect or
horizontal dorsiventral leaves as
seedlings, when their growth is limited by
the available light.[16] Other factors
include the need to balance water loss at
high temperature and low humidity
against the need to absorb atmospheric
carbon dioxide. In most plants, leaves
also are the primary organs responsible
for transpiration and guttation (beads of
fluid forming at leaf margins).

Leaves can also store food and water,

and are modified accordingly to meet
these functions, for example in the
leaves of succulent plants and in bulb
scales. The concentration of
photosynthetic structures in leaves
requires that they be richer in protein,
minerals, and sugars than, say, woody
stem tissues. Accordingly, leaves are
prominent in the diet of many animals.
A leaf shed in autumn.

Correspondingly, leaves represent heavy

investment on the part of the plants
bearing them, and their retention or
disposition are the subject of elaborate
strategies for dealing with pest
pressures, seasonal conditions, and
protective measures such as the growth
of thorns and the production of
phytoliths, lignins, tannins and poisons.
Deciduous plants in frigid or cold
temperate regions typically shed their
leaves in autumn, whereas in areas with
a severe dry season, some plants may
shed their leaves until the dry season
ends. In either case, the shed leaves may
be expected to contribute their retained
nutrients to the soil where they fall.

In contrast, many other non-seasonal

plants, such as palms and conifers, retain
their leaves for long periods; Welwitschia
retains its two main leaves throughout a
lifetime that may exceed a thousand
years.

The leaf-like organs of bryophytes (e.g.,

mosses and liverworts), known as
phyllids, differ morphologically from the
leaves of vascular plants in that they lack
vascular tissue, are usually only a single
cell thick, and have no cuticle stomata or
internal system of intercellular spaces.
The leaves of bryophytes are only
present on the gametophytes, while in
contrast the leaves of vascular plants are
only present on the sporophytes, and are
associated with buds (immature shoot
systems in the leaf axils). These can
further develop into either vegetative or
reproductive structures.[14]

Simple, vascularized leaves

(microphylls), such as those of the early
Devonian lycopsid Baragwanathia, first
evolved as enations, extensions of the
stem. True leaves or euphylls of larger
size and with more complex venation did
not become widespread in other groups
until the Devonian period, by which time
the carbon dioxide concentration in the
atmosphere had dropped significantly.
This occurred independently in several
separate lineages of vascular plants, in
progymnosperms like Archaeopteris, in
Sphenopsida, ferns and later in the
gymnosperms and angiosperms.
Euphylls are also referred to as
macrophylls or megaphylls (large
leaves).[6]

Morphology
Rosa canina: Petiole, two stipules, rachis, five
leaflets

Citrus leaves with translucent glands[17]

A structurally complete leaf of an

angiosperm consists of a petiole (leaf
stalk), a lamina (leaf blade), stipules
(small structures located to either side of
the base of the petiole) and a sheath. Not
every species produces leaves with all of
these structural components. The
proximal stalk or petiole is called a stipe
in ferns. The lamina is the expanded, flat
component of the leaf which contains
the chloroplasts. The sheath is a
structure, typically at the base that fully
or partially clasps the stem above the
node, where the latter is attached. Leaf
sheathes typically occur in grasses and
Apiaceae (umbellifers). Between the
sheath and the lamina, there may be a
pseudopetiole, a petiole like structure.
Pseudopetioles occur in some
monocotyledons including bananas,
palms and bamboos.[18] Stipules may be
conspicuous (e.g. beans and roses),
soon falling or otherwise not obvious as
in Moraceae or absent altogether as in
the Magnoliaceae. A petiole may be
absent (apetiolate), or the blade may not
be laminar (flattened). The tremendous
variety shown in leaf structure (anatomy)
from species to species is presented in
detail below under morphology. The
petiole mechanically links the leaf to the
plant and provides the route for transfer
of water and sugars to and from the leaf.
The lamina is typically the location of the
majority of photosynthesis. The upper
(adaxial) angle between a leaf and a
stem is known as the axil of the leaf. It is
often the location of a bud. Structures
located there are called "axillary".

External leaf characteristics, such as

shape, margin, hairs, the petiole, and the
presence of stipules and glands, are
frequently important for identifying
plants to family, genus or species levels,
and botanists have developed a rich
terminology for describing leaf
characteristics. Leaves almost always
have determinate growth. They grow to a
specific pattern and shape and then stop.
Other plant parts like stems or roots have
non-determinate growth, and will usually
continue to grow as long as they have the
resources to do so.
The type of leaf is usually characteristic
of a species (monomorphic), although
some species produce more than one
type of leaf (dimorphic or polymorphic).
The longest leaves are those of the
Raffia palm, R. regalis which may be up
to 25 m (82 ft) long and 3 m (9.8 ft)
wide.[19] The terminology associated with
the description of leaf morphology is
presented, in illustrated form, at
Wikibooks.

Prostrate leaves in Crossyne guttata

Where leaves are basal, and lie on the
ground, they are referred to as prostrate.

Basic leaf types …

Whorled leaf pattern of the American tiger lily

Perennial plants whose leaves are shed

annually are said to have deciduous
leaves, while leaves that remain through
winter are evergreens. Leaves attached
to stems by stalks (known as petioles)
are called petiolate, and if attached
directly to the stem with no petiole they
are called sessile.[20]

Ferns have fronds.

Conifer leaves are typically needle- or
awl-shaped or scale-like, they are
usually evergreen, but can sometimes
be deciduous. Usually, they have a
single vein.
Flowering plant (Angiosperm) leaves:
the standard form includes stipules, a
petiole, and a lamina.
Lycophytes have microphylls.
 Sheath leaves are the type found in
most grasses and many other
monocots.
Other specialized leaves include those
of Nepenthes, a pitcher plant.

Dicot leaves have blades with pinnate

vegetation (where major veins diverge
from one large mid-vein and have smaller
connecting networks between them).
Less commonly, dicot leaf blades may
have palmate venation (several large
veins diverging from petiole to leaf
edges). Finally, some exhibit parallel
venation.[20]

Monocot leaves in temperate climates

usually have narrow blades, and usually
parallel venation converging at leaf tips
or edges. Some also have pinnate
venation.[20]

Arrangement on the stem …

Different terms are usually used to

describe the arrangement of leaves on
the stem (phyllotaxis):

The leaves on this plant are arranged in pairs

opposite one another, with successive pairs at right
angles to each other (decussate) along the red
stem. Note the developing buds in the axils of these
leaves.
The leaves on this plant are alternately arranged
(Senecio angulatus).

Alternate
One leaf, branch, or flower part
attaches at each point or node on the
stem, and leaves alternate direction, to
a greater or lesser degree, along the
stem.
Basal
Arising from the base of the stem.
Cauline
Arising from the aerial stem.
Opposite
Two leaves, branches, or flower parts
attach at each point or node on the
stem. Leaf attachments are paired at
each node and decussate if, as typical,
each successive pair is rotated 90°
progressing along the stem.
Whorled, or verticillate
Three or more leaves, branches, or
flower parts attach at each point or
node on the stem. As with opposite
leaves, successive whorls may or may
not be decussate, rotated by half the
angle between the leaves in the whorl
(i.e., successive whorls of three rotated
60°, whorls of four rotated 45°, etc.).
Opposite leaves may appear whorled
near the tip of the stem.
Pseudoverticillate describes an
arrangement only appearing whorled,
but not actually so.
Rosulate
Leaves form a rosette.
Rows
The term, distichous, literally means
two rows. Leaves in this arrangement
may be alternate or opposite in their
attachment. The term, 2-ranked, is
equivalent. The terms, tristichous and
tetrastichous, are sometimes
encountered. For example, the "leaves"
(actually microphylls) of most species
of Selaginella are tetrastichous, but not
decussate.
As a stem grows, leaves tend to appear
arranged around the stem in a way that
optimizes yield of light. In essence,
leaves form a helix pattern centered
around the stem, either clockwise or
counterclockwise, with (depending upon
the species) the same angle of
divergence. There is a regularity in these
angles and they follow the numbers in a
Fibonacci sequence: 1/2, 2/3, 3/5, 5/8,
8/13, 13/21, 21/34, 34/55, 55/89. This
series tends to the golden angle, which is
approximately 360° × 34/89 ≈ 137.52° ≈
137° 30′. In the series, the numerator
indicates the number of complete turns
or "gyres" until a leaf arrives at the initial
position and the denominator indicates
the number of leaves in the arrangement.
This can be demonstrated by the
following:

Alternate leaves have an angle of 180°

(or 1⁄2)
120° (or 1⁄3): three leaves in one circle
144° (or 2⁄5): five leaves in two gyres
135° (or 3⁄8): eight leaves in three
gyres.

Divisions of the blade …

A leaf with laminar structure and pinnate venation

Two basic forms of leaves can be
described considering the way the blade
(lamina) is divided. A simple leaf has an
undivided blade. However, the leaf may
be dissected to form lobes, but the gaps
between lobes do not reach to the main
vein. A compound leaf has a fully
subdivided blade, each leaflet of the
blade being separated along a main or
secondary vein. The leaflets may have
petiolules and stipels, the equivalents of
the petioles and stipules of leaves.
Because each leaflet can appear to be a
simple leaf, it is important to recognize
where the petiole occurs to identify a
compound leaf. Compound leaves are a
characteristic of some families of higher
plants, such as the Fabaceae. The middle
vein of a compound leaf or a frond, when
it is present, is called a rachis.

Palmately compound
Leaves have the leaflets radiating from
the end of the petiole, like fingers of
the palm of a hand; for example,
Cannabis (hemp) and Aesculus
(buckeyes).
Pinnately compound
Leaves have the leaflets arranged
along the main or mid-vein.

Odd pinnate
With a terminal leaflet; for example,
Fraxinus (ash).
Even pinnate
Lacking a terminal leaflet; for example,
Swietenia (mahogany). A specific type of
even pinnate is bipinnate, where leaves
only consist of two leaflets; for example,
Hymenaea.
Bipinnately compound
Leaves are twice divided: the leaflets
are arranged along a secondary vein
that is one of several branching off the
rachis. Each leaflet is called a pinnule.
The group of pinnules on each
secondary vein forms a pinna; for
example, Albizia (silk tree).
Trifoliate (or trifoliolate)
A pinnate leaf with just three leaflets;
for example, Trifolium (clover),
 Laburnum (laburnum), and some
species of Toxicodendron (for
instance, poison ivy).
Pinnatifid
Pinnately dissected to the central vein,
but with the leaflets not entirely
separate; for example, Polypodium,
some Sorbus (whitebeams). In
pinnately veined leaves the central vein
in known as the midrib.

Characteristics of the petiole …

The overgrown petioles of rhubarb (Rheum

rhabarbarum) are edible.
Leaves which have a petiole (leaf stalk)
are said to be petiolate.

Sessile (epetiolate) leaves have no

petiole and the blade attaches directly to
the stem. Subpetiolate leaves are nearly
petiolate or have an extremely short
petiole and may appear to be sessile.

In clasping or decurrent leaves, the blade

partially surrounds the stem.

When the leaf base completely surrounds

the stem, the leaves are said to be
perfoliate, such as in Eupatorium
perfoliatum.
In peltate leaves, the petiole attaches to
the blade inside the blade margin.

In some Acacia species, such as the koa

tree (Acacia koa), the petioles are
expanded or broadened and function like
leaf blades; these are called phyllodes.
There may or may not be normal pinnate
leaves at the tip of the phyllode.

A stipule, present on the leaves of many

dicotyledons, is an appendage on each
side at the base of the petiole,
resembling a small leaf. Stipules may be
lasting and not be shed (a stipulate leaf,
such as in roses and beans), or be shed
as the leaf expands, leaving a stipule
scar on the twig (an exstipulate leaf). The
situation, arrangement, and structure of
the stipules is called the "stipulation".

Free, lateral
As in Hibiscus.
Adnate
Fused to the petiole base, as in Rosa.
Ochreate
Provided with ochrea, or sheath-
formed stipules, as in Polygonaceae;
e.g., rhubarb.
Encircling the petiole base

Interpetiolar
Between the petioles of two
opposite leaves, as in Rubiaceae.

Intrapetiolar
 Between the petiole and the
subtending stem, as in
Malpighiaceae.

Veins …

Branching veins on underside of taro leaf

The venation within the bract of a lime tree

Micrograph of a leaf skeleton

Veins (sometimes referred to as nerves)

constitute one of the more visible leaf
traits or characteristics. The veins in a
leaf represent the vascular structure of
the organ, extending into the leaf via the
petiole and providing transportation of
water and nutrients between leaf and
stem, and play a crucial role in the
maintenance of leaf water status and
photosynthetic capacity.They also play a
role in the mechanical support of the
leaf.[21][22] Within the lamina of the leaf,
while some vascular plants possess only
a single vein, in most this vasculature
generally divides (ramifies) according to
a variety of patterns (venation) and form
cylindrical bundles, usually lying in the
median plane of the mesophyll, between
the two layers of epidermis.[23] This
pattern is often specific to taxa, and of
which angiosperms possess two main
types, parallel and reticulate (net like). In
general, parallel venation is typical of
monocots, while reticulate is more
typical of eudicots and magnoliids
("dicots"), though there are many
exceptions.[24][23][25]
The vein or veins entering the leaf from
the petiole are called primary or first-
order veins. The veins branching from
these are secondary or second-order
veins. These primary and secondary
veins are considered major veins or lower
order veins, though some authors include
third order.[26] Each subsequent
branching is sequentially numbered, and
these are the higher order veins, each
branching being associated with a
narrower vein diameter.[27] In parallel
veined leaves, the primary veins run
parallel and equidistant to each other for
most of the length of the leaf and then
converge or fuse (anastomose) towards
the apex. Usually, many smaller minor
veins interconnect these primary veins,
but may terminate with very fine vein
endings in the mesophyll. Minor veins are
more typical of angiosperms, which may
have as many as four higher orders.[26] In
contrast, leaves with reticulate venation
there is a single (sometimes more)
primary vein in the centre of the leaf,
referred to as the midrib or costa and is
continuous with the vasculature of the
petiole more proximally. The midrib then
branches to a number of smaller
secondary veins, also known as second
order veins, that extend toward the leaf
margins. These often terminate in a
hydathode, a secretory organ, at the
margin. In turn, smaller veins branch
from the secondary veins, known as
tertiary or third order (or higher order)
veins, forming a dense reticulate pattern.
The areas or islands of mesophyll lying
between the higher order veins, are called
areoles. Some of the smallest veins
(veinlets) may have their endings in the
areoles, a process known as
areolation.[27] These minor veins act as
the sites of exchange between the
mesophyll and the plant's vascular
system.[22] Thus, minor veins collect the
products of photosynthesis
(photosynthate) from the cells where it
takes place, while major veins are
responsible for its transport outside of
the leaf. At the same time water is being
transported in the opposite
direction.[28][24][23]

The number of vein endings is very

variable, as is whether second order
veins end at the margin, or link back to
other veins.[25] There are many elaborate
variations on the patterns that the leaf
veins form, and these have functional
implications. Of these, angiosperms have
the greatest diversity.[26] Within these the
major veins function as the support and
distribution network for leaves and are
correlated with leaf shape. For instance,
the parallel venation found in most
monocots correlates with their elongated
leaf shape and wide leaf base, while
reticulate venation is seen in simple
entire leaves, while digitate leaves
typically have venation in which three or
more primary veins diverge radially from
a single point.[29][22][27][30]

In evolutionary terms, early emerging

taxa tend to have dichotomous branching
with reticulate systems emerging later.
Veins appeared in the Permian period
(299–252 mya), prior to the appearance
of angiosperms in the Triassic (252–
201 mya), during which vein hierarchy
appeared enabling higher function, larger
leaf size and adaption to a wider variety
of climatic conditions.[26] Although it is
the more complex pattern, branching
veins appear to be plesiomorphic and in
some form were present in ancient seed
plants as long as 250 million years ago.
A pseudo-reticulate venation that is
actually a highly modified penniparallel
one is an autapomorphy of some
Melanthiaceae, which are monocots; e.g.,
Paris quadrifolia (True-lover's Knot). In
leaves with reticulate venation, veins
form a scaffolding matrix imparting
mechanical rigidity to leaves.[31]

Morphology changes within a

single plant
…

Homoblasty
 Characteristic in which a plant has
small changes in leaf size, shape, and
growth habit between juvenile and
adult stages, in contrast to;
Heteroblasty
Characteristic in which a plant has
marked changes in leaf size, shape,
and growth habit between juvenile and
adult stages.

Anatomy

Medium-scale features …

Leaves are normally extensively

vascularized and typically have networks
of vascular bundles containing xylem,
which supplies water for photosynthesis,
and phloem, which transports the sugars
produced by photosynthesis. Many
leaves are covered in trichomes (small
hairs) which have diverse structures and
functions.

Small-scale features …

The major tissue systems present are

 The epidermis, which covers the upper
and lower surfaces
The mesophyll tissue inside the leaf,
which is rich in chloroplasts (also
called chlorenchyma)
The arrangement of veins (the
vascular tissue)

These three tissue systems typically

form a regular organization at the cellular
scale. Specialized cells that differ
markedly from surrounding cells, and
which often synthesize specialized
products such as crystals, are termed
idioblasts.[32]
Major leaf tissues …
Cross-section of a leaf

Epidermal cells
 Spongy mesophyll cells

Epidermis …

SEM image of the leaf epidermis of Nicotiana alata,

showing trichomes (hair-like appendages) and
stomata (eye-shaped slits, visible at full resolution).
The epidermis is the outer layer of cells
covering the leaf. It is covered with a
waxy cuticle which is impermeable to
liquid water and water vapor and forms
the boundary separating the plant's inner
cells from the external world. The cuticle
is in some cases thinner on the lower
epidermis than on the upper epidermis,
and is generally thicker on leaves from
dry climates as compared with those
from wet climates.[33] The epidermis
serves several functions: protection
against water loss by way of
transpiration, regulation of gas exchange
and secretion of metabolic compounds.
Most leaves show dorsoventral anatomy:
The upper (adaxial) and lower (abaxial)
surfaces have somewhat different
construction and may serve different
functions.

The epidermis tissue includes several

differentiated cell types; epidermal cells,
epidermal hair cells (trichomes), cells in
the stomatal complex; guard cells and
subsidiary cells. The epidermal cells are
the most numerous, largest, and least
specialized and form the majority of the
epidermis. They are typically more
elongated in the leaves of monocots than
in those of dicots.

Chloroplasts are generally absent in

epidermal cells, the exception being the
guard cells of the stomata. The stomatal
pores perforate the epidermis and are
surrounded on each side by chloroplast-
containing guard cells, and two to four
subsidiary cells that lack chloroplasts,
forming a specialized cell group known
as the stomatal complex. The opening
and closing of the stomatal aperture is
controlled by the stomatal complex and
regulates the exchange of gases and
water vapor between the outside air and
the interior of the leaf. Stomata therefore
play the important role in allowing
photosynthesis without letting the leaf
dry out. In a typical leaf, the stomata are
more numerous over the abaxial (lower)
epidermis than the adaxial (upper)
epidermis and are more numerous in
plants from cooler climates.

Mesophyll …

Most of the interior of the leaf between

the upper and lower layers of epidermis
is a parenchyma (ground tissue) or
chlorenchyma tissue called the
mesophyll (Greek for "middle leaf"). This
assimilation tissue is the primary
location of photosynthesis in the plant.
The products of photosynthesis are
called "assimilates".

In ferns and most flowering plants, the

mesophyll is divided into two layers:
An upper palisade layer of vertically
elongated cells, one to two cells thick,
directly beneath the adaxial epidermis,
with intercellular air spaces between
them. Its cells contain many more
chloroplasts than the spongy layer.
Cylindrical cells, with the chloroplasts
close to the walls of the cell, can take
optimal advantage of light. The slight
separation of the cells provides
maximum absorption of carbon
dioxide. Sun leaves have a multi-
layered palisade layer, while shade
leaves or older leaves closer to the soil
are single-layered.
Beneath the palisade layer is the
spongy layer. The cells of the spongy
 layer are more branched and not so
tightly packed, so that there are large
intercellular air spaces between them.
The pores or stomata of the epidermis
open into substomatal chambers,
which are connected to the
intercellular air spaces between the
spongy and palisade mesophyll cell, so
that oxygen, carbon dioxide and water
vapor can diffuse into and out of the
leaf and access the mesophyll cells
during respiration, photosynthesis and
transpiration.

Leaves are normally green, due to

chlorophyll in chloroplasts in the
mesophyll cells. Plants that lack
chlorophyll cannot photosynthesize.
Vascular tissue …

The veins of a bramble leaf

The veins are the vascular tissue of the

leaf and are located in the spongy layer
of the mesophyll. The pattern of the
veins is called venation. In angiosperms
the venation is typically parallel in
monocotyledons and forms an
interconnecting network in broad-leaved
plants. They were once thought to be
typical examples of pattern formation
through ramification, but they may
instead exemplify a pattern formed in a
stress tensor field.[34][35][36]

A vein is made up of a vascular bundle.

At the core of each bundle are clusters of
two distinct types of conducting cells:

Xylem
Cells that bring water and minerals
from the roots into the leaf.
Phloem
Cells that usually move sap, with
dissolved sucrose(glucose to sucrose)
produced by photosynthesis in the leaf,
out of the leaf.
The xylem typically lies on the adaxial
side of the vascular bundle and the
phloem typically lies on the abaxial side.
Both are embedded in a dense
parenchyma tissue, called the sheath,
which usually includes some structural
collenchyma tissue.

Leaf development
According to Agnes Arber's partial-shoot
theory of the leaf, leaves are partial
shoots,[37] being derived from leaf
primordia of the shoot apex. Early in
development they are dorsiventrally
flattened with both dorsal and ventral
surfaces.[14] Compound leaves are closer
to shoots than simple leaves.
Developmental studies have shown that
compound leaves, like shoots, may
branch in three dimensions.[38][39] On the
basis of molecular genetics, Eckardt and
Baum (2010) concluded that "it is now
generally accepted that compound
leaves express both leaf and shoot
properties."[40]

Ecology

Biomechanics …

Plants respond and adapt to

environmental factors, such as light and
mechanical stress from wind. Leaves
need to support their own mass and align
themselves in such a way as to optimize
their exposure to the sun, generally more
or less horizontally. However, horizontal
alignment maximizes exposure to
bending forces and failure from stresses
such as wind, snow, hail, falling debris,
animals, and abrasion from surrounding
foliage and plant structures. Overall
leaves are relatively flimsy with regard to
other plant structures such as stems,
branches and roots.[41]

Both leaf blade and petiole structure

influence the leaf's response to forces
such as wind, allowing a degree of
repositioning to minimize drag and
damage, as opposed to resistance. Leaf
movement like this may also increase
turbulence of the air close to the surface
of the leaf, which thins the boundary
layer of air immediately adjacent to the
surface, increasing the capacity for gas
and heat exchange, as well as
photosynthesis. Strong wind forces may
result in diminished leaf number and
surface area, which while reducing drag,
involves a trade off of also reducing
photosynthesis. Thus, leaf design may
involve compromise between carbon
gain, thermoregulation and water loss on
the one hand, and the cost of sustaining
both static and dynamic loads. In
vascular plants, perpendicular forces are
spread over a larger area and are
relatively flexible in both bending and
torsion, enabling elastic deforming
without damage.[41]

Many leaves rely on hydrostatic support

arranged around a skeleton of vascular
tissue for their strength, which depends
on maintaining leaf water status. Both
the mechanics and architecture of the
leaf reflect the need for transportation
and support. Read and Stokes (2006)
consider two basic models, the
"hydrostatic" and "I-beam leaf" form (see
Fig 1).[41] Hydrostatic leaves such as in
Prostanthera lasianthos are large and
thin, and may involve the need for
multiple leaves rather single large leaves
because of the amount of veins needed
to support the periphery of large leaves.
But large leaf size favors efficiency in
photosynthesis and water conservation,
involving further trade offs. On the other
hand, I-beam leaves such as Banksia
marginata involve specialized structures
to stiffen them. These I-beams are
formed from bundle sheath extensions of
sclerenchyma meeting stiffened sub-
epidermal layers. This shifts the balance
from reliance on hydrostatic pressure to
structural support, an obvious advantage
where water is relatively scarce. [41] Long
narrow leaves bend more easily than
ovate leaf blades of the same area.
Monocots typically have such linear
leaves that maximize surface area while
minimising self-shading. In these a high
proportion of longitudinal main veins
provide additional support.[41]

Interactions with other organisms …

Some insects, like Kallima inachus, mimic leaves

Although not as nutritious as other

organs such as fruit, leaves provide a
food source for many organisms. The
leaf is a vital source of energy production
for the plant, and plants have evolved
protection against animals that consume
leaves, such as tannins, chemicals which
hinder the digestion of proteins and have
an unpleasant taste. Animals that are
specialized to eat leaves are known as
folivores.

Some species have cryptic adaptations

by which they use leaves in avoiding
predators. For example, the caterpillars
of some leaf-roller moths will create a
small home in the leaf by folding it over
themselves. Some sawflies similarly roll
the leaves of their food plants into tubes.
Females of the Attelabidae, so-called
leaf-rolling weevils, lay their eggs into
leaves that they then roll up as means of
protection. Other herbivores and their
predators mimic the appearance of the
leaf. Reptiles such as some chameleons,
and insects such as some katydids, also
mimic the oscillating movements of
leaves in the wind, moving from side to
side or back and forth while evading a
possible threat.

Seasonal leaf loss …

Leaves shifting color in autumn (fall)

Leaves in temperate, boreal, and

seasonally dry zones may be seasonally
deciduous (falling off or dying for the
inclement season). This mechanism to
shed leaves is called abscission. When
the leaf is shed, it leaves a leaf scar on
the twig. In cold autumns, they
sometimes change color, and turn yellow,
bright-orange, or red, as various
accessory pigments (carotenoids and
xanthophylls) are revealed when the tree
responds to cold and reduced sunlight by
curtailing chlorophyll production. Red
anthocyanin pigments are now thought
to be produced in the leaf as it dies,
possibly to mask the yellow hue left
when the chlorophyll is lost—yellow
leaves appear to attract herbivores such
as aphids.[42] Optical masking of
chlorophyll by anthocyanins reduces risk
of photo-oxidative damage to leaf cells
as they senesce, which otherwise may
lower the efficiency of nutrient retrieval
from senescing autumn leaves.[43]

Evolutionary adaptation

Poinsettia bracts are leaves which have evolved red

pigmentation in order to attract insects and birds to
the central flowers, an adaptive function normally
served by petals (which are themselves leaves
highly modified by evolution).

In the course of evolution, leaves have

adapted to different environments in the
following ways:

Waxy micro- and nanostructures on the

surface reduce wetting by rain and
adhesion of contamination (See Lotus
effect).
Divided and compound leaves reduce
wind resistance and promote cooling.
Hairs on the leaf surface trap humidity
in dry climates and create a boundary
layer reducing water loss.
Waxy plant cuticles reduce water loss.
Large surface area provides a large
area for capture of sunlight.
In harmful levels of sunlight,
specialized leaves, opaque or partly
buried, admit light through a
translucent leaf window for
photosynthesis at inner leaf surfaces
(e.g. Fenestraria).
Kranz leaf anatomy in plants who
perform C4 carbon fixation
Succulent leaves store water and
organic acids for use in CAM
photosynthesis.
Aromatic oils, poisons or pheromones
produced by leaf borne glands deter
herbivores (e.g. eucalypts).
Inclusions of crystalline minerals deter
herbivores (e.g. silica phytoliths in
grasses, raphides in Araceae).
Petals attract pollinators.
Spines protect the plants from
herbivores (e.g. cacti).
Stinging hairs to protect against
herbivory, e.g. in Urtica dioica and
Dendrocnide moroides (Urticaceae).
Special leaves on carnivorous plants
are adapted for trapping food, mainly
invertebrate prey, though some species
trap small vertebrates as well (see
carnivorous plants).
 Bulbs store food and water (e.g.
onions).
Tendrils allow the plant to climb (e.g.
peas).
Bracts and pseudanthia (false flowers)
replace normal flower structures when
the true flowers are greatly reduced
(e.g. spurges and spathes in the
Araceae.

Terminology
Leaf morphology terms

Shape …

Leaves showing various morphologies. Clockwise

from upper left: tripartite lobation, elliptic with
serrulate margin, palmate venation, acuminate odd-
 g ,p ,
pinnate (center), pinnatisect, lobed, elliptic with
entire margin

Edge (margin) …
Image Term Latin Description

Forma
Entire Even; with a smooth margin; without toothing
integra

Ciliate Ciliata Fringed with hairs

Crenate Crenata Wavy-toothed; dentate with rounded teeth

Toothed

May be coarsely dentate, having large teeth

Dentate Dentata
or glandular dentate, having teeth which bear
glands

Denticulate Denticulata Finely toothed

Duplicato-
Doubly serrate Each tooth bearing smaller teeth
dentata

Saw-toothed; with asymmetrical teeth pointing

Serrate Serrata
forward

Serrulate Serrulata Finely serrate

With deep, wave-like indentations; coarsely

Sinuate Sinuosa
crenate

Indented, with the indentations not reaching the

Lobate Lobata
center

Undulate Undulata With a wavy edge, shallower than sinuate

Spiny or
Spiculata With stiff, sharp points such as thistles
pungent

Apex (tip) …
Image Term Latin Description

Long-pointed, prolonged into a narrow, tapering point in a

Acuminate _
concave manner

Acute _ Ending in a sharp, but not prolonged point

Cuspidate _ With a sharp, elongated, rigid tip; tipped with a cusp

Emarginate _ Indented, with a shallow notch at the tip

Mucronate _ Abruptly tipped with a small short point

Mucronulate _ Mucronate, but with a noticeably diminutive spine

Obcordate _ Inversely heart-shaped

Obtuse _ Rounded or blunt

Truncate _ Ending abruptly with a flat end

Base …

Acuminate
Coming to a sharp, narrow, prolonged
point.
Acute
Coming to a sharp, but not prolonged
point.
Auriculate
Ear-shaped.
Cordate
Heart-shaped with the notch towards
the stalk.
Cuneate
Wedge-shaped.
Hastate
Shaped like an halberd and with the
basal lobes pointing outward.
Oblique
Slanting.
Reniform
Kidney-shaped but rounder and
broader than long.
Rounded
Curving shape.
Sagittate
 Shaped like an arrowhead and with the
acute basal lobes pointing downward.
Truncate
Ending abruptly with a flat end, that
looks cut off.

Surface …

The scale-shaped leaves of the Norfolk Island Pine.

The leaf surface is also host to a large
variety of microorganisms; in this context
it is referred to as the phyllosphere.

Hairiness …

Common mullein (Verbascum thapsus) leaves are

covered in dense, stellate trichomes.
Scanning electron microscope image of trichomes
on the lower surface of a Coleus blumei (coleus)
leaf

"Hairs" on plants are properly called

trichomes. Leaves can show several
degrees of hairiness. The meaning of
several of the following terms can
overlap.

Arachnoid, or arachnose
With many fine, entangled hairs giving
a cobwebby appearance.
Barbellate
With finely barbed hairs (barbellae).
Bearded
With long, stiff hairs.
Bristly
With stiff hair-like prickles.
Canescent
Hoary with dense grayish-white
pubescence.
Ciliate
Marginally fringed with short hairs
(cilia).
Ciliolate
Minutely ciliate.
Floccose
With flocks of soft, woolly hairs, which
tend to rub off.
Glabrescent
Losing hairs with age.
Glabrous
No hairs of any kind present.
Glandular
With a gland at the tip of the hair.
Hirsute
With rather rough or stiff hairs.
Hispid
With rigid, bristly hairs.
Hispidulous
Minutely hispid.
Hoary
With a fine, close grayish-white
pubescence.
Lanate, or lanose
With woolly hairs.
Pilose
With soft, clearly separated hairs.
Puberulent, or puberulous
With fine, minute hairs.
Pubescent
With soft, short and erect hairs.
Scabrous, or scabrid
Rough to the touch.
Sericeous
Silky appearance through fine, straight
and appressed (lying close and flat)
hairs.
Silky
With adpressed, soft and straight
pubescence.
Stellate, or stelliform
With star-shaped hairs.
Strigose
With appressed, sharp, straight and
stiff hairs.
Tomentose
Densely pubescent with matted, soft
white woolly hairs.

Cano-tomentose
Between canescent and tomentose.

Felted-tomentose
Woolly and matted with curly hairs.
Tomentulose
Minutely or only slightly tomentose.
Villous
With long and soft hairs, usually
curved.
Woolly
 With long, soft and tortuous or matted
hairs.

Timing …

Hysteranthous
Developing after the flowers [44]
Synanthous
Developing at the same time as the
flowers [45]

Venation …

Classification …

Hickey primary venation types

1. Pinnate
venation,
Ostrya
virginiana
2. Parallel
venation, Iris

3.
Campylodromo
us venation,
Maianthemum
bifolium
4.
Acrodromous
venation
(basal),
Miconia
calvescens

5.
Actinodromous
venation
(suprabasal),
Givotia
moluccana
 6.
Palinactodrom
ous venation,
Platanus
orientalis

A number of different classification

systems of the patterns of leaf veins
(venation or veination) have been
described,[25] starting with Ettingshausen
(1861),[46] together with many different
descriptive terms, and the terminology
has been described as "formidable".[25]
One of the commonest among these is
the Hickey system, originally developed
for "dicotyledons" and using a number of
Ettingshausen's terms derived from
Greek (1973–1979):[47][48][49] (see also:
Simpson Figure 9.12, p. 468)[25]

Hickey system E…

1. Pinnate (feather-veined, reticulate,

pinnate-netted, penniribbed,
penninerved, or penniveined)
The veins arise pinnately (feather like)
from a single primary vein (mid-vein)
and subdivide into secondary veinlets,
known as higher order veins. These, in
turn, form a complicated network. This
type of venation is typical for (but by
no means limited to) "dicotyledons"
(non monocotyledon angiosperms).
E.g., Ostrya.
There are three subtypes of pinnate
venation:

Craspedodromous (Greek:
kraspedon - edge, dromos - running)
The major veins reach to the margin
of the leaf.

Camptodromous
Major veins extend close to the
margin, but bend before they
intersect with the margin.

Hyphodromous
All secondary veins are absent,
rudimentary or concealed
These in turn have a number of further
subtypes such as eucamptodromous,
where secondary veins curve near the
margin without joining adjacent
secondary veins.

Pinnate

Craspe Campt Hypho

dodro odrom dromo
mous ous us

2. Parallelodromous (parallel-veined,
parallel-ribbed, parallel-nerved,
penniparallel, striate)
Two or more primary veins originating
beside each other at the leaf base, and
running parallel to each other to the
apex and then converging there.
Commissural veins (small veins)
connect the major parallel veins.
Typical for most monocotyledons,
such as grasses.

The additional terms marginal (primary

veins reach the margin), and reticulate
(primary veins do not reach the
margin) are also used.

Parallelodromous

3. Campylodromous (campylos -
curve)
Several primary veins or branches
originating at or close to a single point
and running in recurved arches, then
converging at apex. E.g. Maianthemum
.

Campylodromous

4. Acrodromous
Two or more primary or well developed
secondary veins in convergent arches
towards apex, without basal
recurvature as in Campylodromous.
May be basal or suprabasal depending
on origin, and perfect or imperfect
depending on whether they reach to
2/3 of the way to the apex. E.g.,
Miconia (basal type), Endlicheria
(suprabasal type).

Acrodromous

Imper Imper Perfe Perfe

fect fect ct ct
basal supra basal supra
basal basal

5. Actinodromous
Three or more primary veins diverging
radially from a single point. E.g.,
Arcangelisia (basal type), Givotia
(suprabasal type).

Actinodromous
 Imper Imper
fect fect
margi reticu
nal late

6. Palinactodromous
Primary veins with one or more points
of secondary dichotomous branching
beyond the primary divergence, either
closely or more distantly spaced. E.g.,
Platanus.

Venation of a Poinsettia (Euphorbia pulcherrima)

Venation of a Poinsettia (Euphorbia pulcherrima)
leaf.

Palinactodromous

Types 4–6 may similarly be subclassified

as basal (primaries joined at the base of
the blade) or suprabasal (diverging above
the blade base), and perfect or imperfect,
but also flabellate.

At about the same time, Melville (1976)

described a system applicable to all
Angiosperms and using Latin and English
terminology.[50] Melville also had six
divisions, based on the order in which
veins develop.

Arbuscular (arbuscularis)
Branching repeatedly by regular
dichotomy to give rise to a three
dimensional bush-like structure
consisting of linear segment (2
subclasses)
Flabellate (flabellatus)
Primary veins straight or only slightly
curved, diverging from the base in a
fan-like manner (4 subclasses)
Palmate (palmatus)
Curved primary veins (3 subclasses)
Pinnate (pinnatus)
 Single primary vein, the midrib, along
which straight or arching secondary
veins are arranged at more or less
regular intervals (6 subclasses)
Collimate (collimatus)
Numerous longitudinally parallel
primary veins arising from a transverse
meristem (5 subclasses)
Conglutinate (conglutinatus)
Derived from fused pinnate leaflets (3
subclasses)

A modified form of the Hickey system

was later incorporated into the
Smithsonian classification (1999) which
proposed seven main types of venation,
based on the architecture of the primary
veins, adding Flabellate as an additional
main type. Further classification was
then made on the basis of secondary
veins, with 12 further types, such as;

Brochidodromous
Closed form in which the secondaries
are joined together in a series of
prominent arches, as in Hildegardia.
Craspedodromous
Open form with secondaries
terminating at the margin, in toothed
leaves, as in Celtis.
Eucamptodromous
Intermediate form with upturned
secondaries that gradually diminish
apically but inside the margin, and
connected by intermediate tertiary
 veins rather than loops between
secondaries, as in Cornus.
Cladodromous
Secondaries freely branching toward
the margin, as in Rhus.

terms which had been used as subtypes

in the original Hickey system.[51]

Secondary venation patterns

Brochi Craspe Eucam Cladod

dodro dodro ptodro romou
mous mous mous s
Brochidodromo Craspedo Eucampt Cladodromous
us dromous odromo Rhus ovata
Hildegardia Celtis us
migeodii occidenta Cornus
lis officinali
s

Further descriptions included the higher

order, or minor veins and the patterns of
areoles (see Leaf Architecture Working
Group, Figures 28–29).[51]
Flabellate venation, Adiantum cunninghamii

Flabellate
Several to many equal fine basal veins
diverging radially at low angles and
branching apically. E.g. Paranomus.

Flabellate

Analyses of vein patterns often fall into

consideration of the vein orders, primary
vein type, secondary vein type (major
veins), and minor vein density. A number
of authors have adopted simplified
versions of these schemes.[52][25] At its
simplest the primary vein types can be
considered in three or four groups
depending on the plant divisions being
considered;

pinnate
palmate
parallel

where palmate refers to multiple primary

veins that radiate from the petiole, as
opposed to branching from the central
main vein in the pinnate form, and
encompasses both of Hickey types 4 and
5, which are preserved as subtypes; e.g.,
palmate-acrodromous (see National Park
Service Leaf Guide).[53]

Palmate venation, Acer truncatum

Palmate, Palmate-netted, palmate-

veined, fan-veined
Several main veins of approximately
equal size diverge from a common
point near the leaf base where the
petiole attaches, and radiate toward
 the edge of the leaf. Palmately veined
leaves are often lobed or divided with
lobes radiating from the common
point. They may vary in the number of
primary veins (3 or more), but always
radiate from a common point.[54] e.g.
most Acer (maples).

Palmate

Other systems E…

Alternatively, Simpson uses:[25]

Uninervous
Central midrib with no lateral veins
(microphyllous), seen in the non-seed
bearing tracheophytes, such as
horsetails
Dichotomous
Veins successively branching into
equally sized veins from a common
point, forming a Y junction, fanning
out. Amongst temperate woody plants,
Ginkgo biloba is the only species
exhibiting dichotomous venation. Also
some pteridophytes (ferns).[54]
Parallel
Primary and secondary veins roughly
parallel to each other, running the
length of the leaf, often connected by
short perpendicular links, rather than
form networks. In some species, the
parallel veins join together at the base
and apex, such as needle-type
evergreens and grasses. Characteristic
of monocotyledons, but exceptions
include Arisaema, and as below, under
netted.[54]
Netted (reticulate, pinnate)
A prominent midvein with secondary
veins branching off along both sides of
it. The name derives from the ultimate
veinlets which form an interconnecting
net like pattern or network. (The
primary and secondary venation may
be referred to as pinnate, while the net
like finer veins are referred to as netted
or reticulate); most non-monocot
angiosperms, exceptions including
Calophyllum. Some monocots have
reticulate venation, including
Colocasia, Dioscorea and Smilax.[54]

Equiset Ginkgo biloba:

um: Dichotomous venation
Reduce
d
microp
hyllous
leaves
(L)
 arising
in
whorl
from
node

However, these simplified systems allow

for further division into multiple
subtypes. Simpson,[25] (and others)[55]
divides parallel and netted (and some
use only these two terms for
Angiosperms)[56] on the basis of the
number of primary veins (costa) as
follows;

Parallel

Penni-parallel (pinnate, pinnate

parallel, unicostate parallel)
Single central prominent midrib,
secondary veins from this arise
perpendicularly to it and run parallel
to each other towards the margin or
tip, but do not join (anastomose).
The term unicostate refers to the
prominence of the single midrib
(costa) running the length of the leaf
from base to apex. e.g. Zingiberales,
such as Bananas etc.

Palmate-parallel (multicostate
parallel)
Several equally prominent primary
veins arising from a single point at
the base and running parallel
towards tip or margin. The term
multicostate refers to having more
 than one prominent main vein. e.g.
"fan" (palmate) palms (Arecaceae)

Multicostate parallel convergent

Mid-veins converge at apex e.g.
Bambusa arundinacea = B.
bambos (Aracaceae), Eichornia

Multicostate parallel divergent

Mid-veins diverge more or less
parallel towards the margin e.g.
Borassus (Poaceae), fan palms
Netted (Reticulate)

Pinnately (veined, netted,

unicostate reticulate)
Single prominent midrib running
from base to apex, secondary veins
arising on both sides along the
length of the primary midrib, running
towards the margin or apex (tip),
with a network of smaller veinlets
forming a reticulum (mesh or
network). e.g. Mangifera, Ficus
religiosa, Psidium guajava, Hibiscus
rosa-sinensis, Salix alba

Palmately (multicostate reticulate)

More than one primary veins arising
from a single point, running from
base to apex. e.g. Liquidambar
styraciflua This may be further
subdivided;

Multicostate convergent
Major veins diverge from origin at
base then converge towards the
 tip. e.g. Zizyphus, Smilax,
Cinnamomum

Multicostate divergent
All major veins diverge towards
the tip. e.g. Gossypium, Cucurbita,
Carica papaya, Ricinus communis

Ternately (ternate-netted)
Three primary veins, as above, e.g.
(see) Ceanothus leucodermis,[57] C.
tomentosus,[58] Encelia farinosa

Simpson venation patterns

Maranta leuconeura Coccothrinax Bambus

var. erythroneura argentea a
var. erythroneura argentea a
bambos:
(Zingiberales): (Arecaceae): Multicos
Penni-parallel Palmate-parallel tate
parallel
converge
nt

Borassus
sp.:
Multicostat
e parallel
divergent

Sali Liquidambar styraciflua: Ziziphus jujuba:

x Palmately netted Multicostate
x Palmately netted Multicostate

alb palmate
a: convergent
Pin
nat
ely
nett
ed

Gossypium
tomentosum
:
Multicostate
palmate
divergent
These complex systems are not used
much in morphological descriptions of
taxa, but have usefulness in plant
identification, [25] although criticized as
being unduly burdened with jargon.[59]

An older, even simpler system, used in

some flora[60] uses only two categories,
open and closed.[61]

Open: Higher order veins have free

endings among the cells and are more
characteristic of non-monocotyledon
angiosperms. They are more likely to
be associated with leaf shapes that are
toothed, lobed or compound. They may
be subdivided as;
 Pinnate (feather-veined) leaves,
with a main central vein or rib
(midrib), from which the remainder
of the vein system arises
Palmate, in which three or more
main ribs rise together at the base
of the leaf, and diverge upward.
Dichotomous, as in ferns, where
the veins fork repeatedly
Closed: Higher order veins are
connected in loops without ending
freely among the cells. These tend to
be in leaves with smooth outlines, and
are characteristic of monocotyledons.
They may be subdivided into
whether the veins run parallel, as
 in grasses, or have other patterns.
Other descriptive terms …

There are also many other descriptive

terms, often with very specialized usage
and confined to specific taxonomic
groups.[62] The conspicuousness of veins
depends on a number of features. These
include the width of the veins, their
prominence in relation to the lamina
surface and the degree of opacity of the
surface, which may hide finer veins. In
this regard, veins are called obscure and
the order of veins that are obscured and
whether upper, lower or both surfaces,
further specified.[63][54]
Terms that describe vein prominence
include bullate, channelled, flat, guttered,
impressed, prominent and recessed
(Fig. 6.1 Hawthorne & Lawrence
2013).[59][64] Veins may show different
types of prominence in different areas of
the leaf. For instance Pimenta racemosa
has a channelled midrib on the upper
surfae, but this is prominent on the lower
surface.[59]

Describing vein prominence:

Bullate
Surface of leaf raised in a series of
domes between the veins on the upper
surface, and therefore also with
marked depressions. e.g. Rytigynia
pauciflora,[65] Vitis vinifera
Channelled (canalicululate)
Veins sunken below the surface,
resulting in a rounded channel.
Sometimes confused with "guttered"
because the channels may function as
gutters for rain to run off and allow
drying, as in many
Melastomataceae.[66] e.g. (see)
Pimenta racemosa (Myrtaceae),[67]
Clidemia hirta (Melastomataceae).
Guttered
Veins partly prominent, the crest above
the leaf lamina surface, but with
channels running along each side, like
gutters
Impressed
Vein forming raised line or ridge which
lies below the plane of the surface
which bears it, as if pressed into it, and
are often exposed on the lower
surface. Tissue near the veins often
appears to pucker, giving them a
sunken or embossed appearance
Obscure
Veins not visible, or not at all clear; if
unspecified, then not visible with the
naked eye. e.g. Berberis gagnepainii. In
this Berberis, the veins are only
obscure on the undersurface.[68]
Prominent
Vein raised above surrounding surface
so to be easily felt when stroked with
finger. e.g. (see) Pimenta racemosa,[67]
Spathiphyllum cannifolium[69]
Recessed
Vein is sunk below the surface, more
prominent than surrounding tissues
but more sunken in channel than with
impressed veins. e.g. Viburnum
plicatum.

Types of vein prominence

Vitis vinifera Clidemia hirta

Bullate Channeled
 Cornus mas
Impressed

Berberis gagnepainii Spathiphyllu

Obscure (under surface) m
cannifolium
Prominent

Viburnum
plicatum
Recessed
Describing other features:

Plinervy (plinerved)
More than one main vein (nerve) at the
base. Lateral secondary veins
branching from a point above the base
of the leaf. Usually expressed as a
suffix, as in 3-plinerved or triplinerved
leaf. In a 3-plinerved (triplinerved) leaf
three main veins branch above the
base of the lamina (two secondary
veins and the main vein) and run
essentially parallel subsequently, as in
Ceanothus and in Celtis. Similarly, a
quintuplinerve (five-veined) leaf has
four secondary veins and a main vein.
A pattern with 3-7 veins is especially
 conspicuous in Melastomataceae. The
term has also been used in Vaccinieae.
The term has been used as
synonymous with acrodromous,
palmate-acrodromous or suprabasal
acrodromous, and is thought to be too
broadly defined.[70][70]
Scalariform
Veins arranged like the rungs of a
ladder, particularly higher order veins
Submarginal
Veins running close to leaf margin
Trinerved
2 major basal nerves besides the
midrib
Diagrams of venation patterns …
Image Term Description

Arcuate Secondary arching toward the apex

Dichotomous Veins splitting in two

Longitudinal All veins aligned mostly with the midvein

Parallel All veins parallel and not intersecting

Pinnate Secondary veins borne from midrib

Reticulate All veins branching repeatedly, net veined

Veins coming from the center of the leaf and radiating toward the
Rotate
edges

Tertiary veins running perpendicular to axis of main vein, connecting

Transverse
secondary veins

Size …

The terms megaphyll, macrophyll,

mesophyll, notophyll, microphyll,
nanophyll and leptophyll are used to
describe leaf sizes (in descending order),
in a classification devised in 1934 by
Christen C. Raunkiær and since modified
by others.[71]

See also
Glossary of leaf morphology
Glossary of plant morphology:Leaves
Crown (botany)
Evolutionary history of leaves
Evolutionary development of leaves
Leaf Area Index
Leaf protein concentrate
Leaf sensor – a device that measures
the moisture level in plant leaves
Leaf shape
 Vernation – sprouting of leaves, also
the arrangement of leaves in the bud

References
1. Esau 2006.
2. Cutter 1969.
3. Haupt 1953.
4. Mauseth 2009.
5. James et al 1999.
6. Stewart & Rothwell 1993.
7. Cooney-Sovetts & Sattler 1987.
8. Tsukaya 2013.
9. Feugier 2006.
10. Purcell 2016.
11. Willert et al 1992.
12. Bayer 1982.
13. Marloth 1913–1932.
14. Simpson 2011, p. 356.
15. Krogh 2010.
16. James & Bell 2000.
17. Heywood et al 2007.
18. Simpson 2011, pp. 356–357.
19. Hallé 1977.
20. Botany Illustrated: Introduction to
Plants Major Groups Flowering Plant
Families. Thomson Science. 1984.
p. 21.
21. Rolland-Lagan et al 2009.
22. Walls 2011.
23. Dickison 2000.
24. Rudall 2007.
25. Simpson 2011, Leaf venation
pp. 465–468
26. Sack & Scoffoni 2013.
27. Roth-Nebelsick et al 2001.
28. Ueno et al 2006.
29. Runions et al 2005.
30. Massey & Murphy 1996, Surface-
Venation-Texure
31. Bagchi et al 2016.
32. Cote 2009.
33. Clements 1905.
34. Couder et al 2002.
35. Corson et al 2009.
36. Laguna et al 2008.
37. Arber 1950.
38. Rutishauser & Sattler 1997.
39. Lacroix et al 2003.
40. Eckardt & Baum 2010.
41. Read & Stokes 2006.
42. Doring et al 2009.
43. Feild et al 2001.
44. Kew Glossary: Hysteranthous
45. Kew Glossary: Synanthous
46. Ettingshausen 1861.
47. Hickey 1973.
48. Hickey & Wolfe 1975.
49. Hickey 1979.
50. Melville 1976.
51. Leaf Architecture Working Group
1999.
52. Judd et al 2007.
53. Florissant Leaf Key 2016.
54. Kling et al 2005, Leaf Venation
55. Berg 2007.
56. Angiosperm Morphology 2017,
Venation
57. Simpson 2017, Ceanothus
leucodermis
58. Simpson 2017, Ceanothus
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59. Hawthorne & Lawrence 2013, Leaf
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60. Cullen et al 2011.
61. Beach 1914, Venation
62. Neotropikey 2017.
63. Oxford herbaria glossary 2017.
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prominence
65. Verdcourt & Bridson 1991.
66. Hemsley & Poole 2004, Leaf
morphology and drying p. 254
67. Hughes 2017, Pimenta racemosa
68. Cullen et al 2011, Berberis
gagnepainii vol. II p. 398
69. Kwantlen 2015, Spathiphyllum
cannifolium
70. Pedraza-Peñalosa 2013.
71. Whitten et al 1997.
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