To many, the word 

tree evokes images of such ancient, powerful, and

majestic structures as oaks and sequoias, the latter being among the
most massive and longest-living organisms in the world. Although the
majority of Earth’s terrestrial biomass is represented by trees, the
fundamental importance of these seemingly ubiquitous plants for the
very existence and diversity of life on Earth is perhaps not fully
appreciated. The biosphere is dependent on the metabolism, death,
and recycling of plants, especially trees. Their vast trunks
and root systems store carbon dioxide, move water, and produce
oxygen that is released into the atmosphere. The organic matter of the
soil develops primarily from decayed leaves, twigs, branches, roots,
and fallen trees, all of which recycle nitrogen, carbon, oxygen, and
other important nutrients. There are few organisms as important as
trees for maintaining Earth’s ecology.

This article discusses the historical, popular, and botanical

classifications of trees, their evolution, their importance to humans,
and their general structure and patterns of growth. For more
information on the three botanical groups that include
trees, see fern, gymnosperm (including conifers), and angiosperm (the
flowering plants). For general information on plants, see plant.

Classification Of Trees
The ancient Greeks developed a classification about 300 BCE in which
plants were grouped according to their general form—that is, as trees,
shrubs, undershrubs, and vines. This classification was used for
almost 1,000 years. Modern classifications of plants attempt to assign
a plant to a particular taxon and establish relationships with other
plants based on genetics, cytology, ecology, behaviour, and probable
evolutionary lineages, in addition to gross morphology. Popular
classifications, however, remain useful tools for studying the common
stresses that the environment exerts on all plants and the general
patterns of adaptation that are shown no matter how distantly plants
are related.

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Phylogenetic classifications

Trees are represented in each of the major groups of the vascular

plants: pteridophytes (seedless vascular plants that include the tree
ferns), gymnosperms (cycads, ginkgoes, and conifers),
and angiosperms (flowering plants).

Although tree ferns account for only a small percentage of ferns, many

are conspicuous members of a forest, attaining heights of 7 to 10
metres (23 to 33 feet); some are 15, 18, or occasionally 24 metres tall
(49, 59, or 79 feet). These graceful trees, which are natives of humid
montane forests in the tropics and subtropics and of warm temperate
regions of the Southern Hemisphere, have huge lacy leaves; they are
the remnants of a vastly more numerous flora that populated much of
the Earth during the Carboniferous Period (about 360 to 300 million
years ago).

tree ferns
Tree ferns (Alsophila australis), the largest of all ferns.
Walter Chandoha
Cycads compose the Cycadophyta, a division of gymnospermous
plants consisting of 4 families and approximately 140 species. Natives
of warm regions of the Eastern and Western hemispheres, they also
are remnants of a much larger number of species that in past geologic
ages dominated the Earth’s flora.

The ginkgo is the only living representative of the

division Ginkgophyta. It is a relic that has been preserved in
cultivation around Buddhist temples in China and elsewhere since the
mid-18th century; the tree probably no longer exists in a wild state.

Conifers (division Coniferophyta) include trees and shrubs in

7 extant families and 550 species. Familiar representatives
are araucarias, cedars, cypresses, Douglas
firs, firs, hemlocks, junipers, larches, pines, podocarps, redwoods, spr
uces, and yews.

monkey puzzle tree

Monkey puzzle tree (Araucaria araucana).
Ernest Manewal/Shostal Associates
Angiosperms dominate the Earth’s present flora; they contain more
than 250,000 species, among which are the majority of the world’s
trees. Angiosperms are sometimes divided on the basis of a group of
characteristics into two groups: the monocotyledons and the
dicotyledons. The most numerous of the monocotyledonous trees
are palms; others include agaves, aloes, dracaenas, screw pines,
and yuccas. By far the greatest number of tree species are
dicotyledons; they are represented by such familiar groups
as birches, elms, hollies, magnolias, maples, oaks, poplars, ashes,
and willows.

Ecological and evolutionary classification

The tree is not an immutable biological category but rather a human

concept based on visual criteria. Perhaps a general definition would
describe a tree as a perennial woody plant that develops along a single
main trunk to a height of at least 4.5 metres (15 feet) at maturity. This
may be contrasted with a shrub, which might be loosely defined as a
woody plant with multiple stems that is, in most cases, less than 3
metres (about 10 feet) tall. However, a species fitting the description of
either in one area of the world might not necessarily do so in other
regions, since a variety of stresses shape the habit of the mature plant.
Thus, a given woody species may be a tree in one set of habitats within
its range and a shrub elsewhere. For example, the spruce and fir may
thrive in the tree form at the base of a mountain but assume a shrub
form near the mountaintop, the variation due principally to stresses
exerted by such environmental conditions as altitude, temperature,
and oxygen tension.

As seen in the section above, trees are found among many plant
families that also include shrubs and herbs, so that the concept of tree
is not a phylogenetic one. Further, there is no clear consensus as to
whether the tree form is the advanced or primitive condition. Some
paleobotanists suggest that trees are the most primitive members
within these plant families. However, tree forms are found in all the
vascular plants, from the club mosses and ferns to the gymnosperms
and angiosperms. It is furthermore true that, among the flowering
plants, trees are found not only among the most primitive members
(order Magnoliales) but also among the more specialized, or advanced,
members, such as the roses (order Rosales).

Consequently, from both a taxonomic and a phylogenetic perspective,

the tree is an artificial category. On an ecological basis, however, the
tree can be recognized as a natural construct, as it represents an
adaptive strategy by many different taxa to exploit and dominate the
habitat above the ground.

In the early stages of the development of terrestrial life, land plants

were rootless and leafless. Since they had their origins in
aqueous environments, they did not require the specialized conducting
and supporting tissues afforded by roots and stems, nor did they
require localized regions of carbohydrate synthesis, since each cell was
involved in metabolism, water and nutrient absorption, and
respiration. Habitats farther from the water as well as aerial habitats
represented available uninhabited environments.

One key to exploiting terrestrial habitats is increasing complexity of

the plant’s form to allow specialization of function. This requires
physiological and morphological complexity as well as biological
optimization. If all the tissues of massive tree trunks were alive, for
example, the physiological cost of maintaining these structures in the
living state would be enormous and probably unattainable. An elegant
solution came in the form of tremendous structural adaptations: new
tissues and organs permitted localization of the functions of the plant
body.

The evolution of discrete plant body parts with separate functions

allowed plants to move onto the land and undergo an
incredible adaptive radiation. Leaves evolved as specialized
photosynthetic organs. Stems provided mechanical strength as well as
a conductive capacity to transport water and nutrients from the roots
to the leaves. Roots provided anchorage and absorption of sufficient
water and nutrients to support the remainder of the plant.