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 heavily
morphologically from the leaves of vascular plants. In most cases, they lack vascular tissue, are only a
single cell thick, and have no cuticle, stomata, or internal system of intercellular spaces. (The phyllids of
the moss family Polytrichaceae are notable exceptions.) The phyllids of bryophytes are only present on
the gametophytes, while in contrast the leaves of vascular plants are only present on the sporophytes.
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
See also: Glossary of leaf morphology

Leafstem of dog rose with petiole, stipules and leaflets

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 Poaceae (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

Main article: Phyllotaxis

The arrangement of leaves on the stem is known as phyllotaxis.[21] A large variety of phyllotactic
patterns occur in nature:

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 (Senecio angulatus) are alternately arranged.

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 plant.

Cauline

Attached to 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.

Decussate

An opposite arrangement in which each successive pair is rotated 90° from the previous.

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.

In the simplest mathematical models of phyllotaxis, the apex of the stem is represented as a circle. Each
new node is formed at the apex, and it is rotated by a constant angle from the previous node. This angle
is called the divergence angle. The number of leaves that grow from a node depends on the plant
species. When a single leaf grows from each node, and when the stem is held straight, the leaves form a
helix.
The divergence angle is often represented as a fraction of a full rotation around the stem. A rotation
fraction of 1/2 (a divergence angle of 180°) produces an alternate arrangement, such as in Gasteria or
the fan-aloe Kumara plicatilis. Rotation fractions of 1/3 (divergence angles of 120°) occur in beech and
hazel. Oak and apricot rotate by 2/5, sunflowers, poplar, and pear by 3/8, and in willow and almond the
fraction is 5/13.[22] These arrangements are periodic. The denominator of the rotation fraction
indicates the