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Cannabaceae is a small
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...
s, known as the hemp family. As now circumscribed, the family includes about 170 species grouped in about 11 genera, including ''
Cannabis ''Cannabis'' () is a genus of flowering plants in the family Cannabaceae that is widely accepted as being indigenous to and originating from the continent of Asia. However, the number of species is disputed, with as many as three species be ...
'' (hemp), ''
Humulus ''Humulus'', or hop, is a small genus of flowering plants in the family Cannabaceae. The hop is native to temperate regions of the Northern Hemisphere. Hops are the female flowers (seed cones, strobiles) of the hop species '' H. lupulus''; as a ...
'' (
hops Hops are the flowers (also called seed cones or strobiles) of the hop plant ''Humulus lupulus'', a member of the Cannabaceae family of flowering plants. They are used primarily as a bittering, flavouring, and stability agent in beer, to whic ...
) and ''
Celtis ''Celtis'' is a genus of about 60–70 species of deciduous trees, commonly known as hackberries or nettle trees, in the hemp family Cannabaceae. It has a cosmopolitan distribution. Description ''Celtis'' species are generally medium-sized tree ...
'' (hackberries). ''Celtis'' is by far the largest genus, containing about 100 species.Stevens, P.F. (2001 onwards
"Cannabaceae"
''Angiosperm Phylogeny Website'', retrieved 2014-02-25
Cannabaceae is a member of the
Rosales Rosales (, ) are an order of flowering plants. Peter F. Stevens (2001 onwards). "Rosales". At: Trees At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below) Well-known members of Rosales include: ...
. Members of the family are erect or climbing plants with petalless flowers and dry, one-seeded fruits. Hemp (''Cannabis'') and hop (''Humulus'') are the most economically important species. Other than a shared evolutionary origin, members of the family have few common characteristics; some are trees (e.g. ''Celtis''), others are
herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
s (e.g. ''Cannabis'').


Description

Members of this family can be trees (e.g. ''
Celtis ''Celtis'' is a genus of about 60–70 species of deciduous trees, commonly known as hackberries or nettle trees, in the hemp family Cannabaceae. It has a cosmopolitan distribution. Description ''Celtis'' species are generally medium-sized tree ...
''), erect herbs (e.g. ''
Cannabis ''Cannabis'' () is a genus of flowering plants in the family Cannabaceae that is widely accepted as being indigenous to and originating from the continent of Asia. However, the number of species is disputed, with as many as three species be ...
''), or twining herbs (e.g. ''
Humulus ''Humulus'', or hop, is a small genus of flowering plants in the family Cannabaceae. The hop is native to temperate regions of the Northern Hemisphere. Hops are the female flowers (seed cones, strobiles) of the hop species '' H. lupulus''; as a ...
''). Leaves are often more or less palmately lobed or palmately compound and always bear
stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole (botany), petiole). They are primarily found among dicots and rare among monocots. Stipules are considered part ...
s.
Cystolith Cystolith (Gr. "cavity" and "stone") is a botanical term for outgrowths of the epidermal cell wall, usually of calcium carbonate but sometimes of silicon dioxide also, formed in a cellulose matrix in special cell (biology), cells called lithocys ...
s are always present and some members of this family possess
laticifer A laticifer is a type of elongated secretory cell found in the leaves and/or stems of plants that produce latex and rubber as secondary metabolites. Laticifers may be divided into: *Articulated laticifers, i.e., composed of a series of cells joine ...
s. Cannabaceae are often
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
(distinct male and female plants). The flowers are
actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
(radially symmetrical) and not showy, as these plants are pollinated by the wind. As an adaptation to this kind of pollination, the calyx and corolla are radically reduced to only vestigial remnants found as an adherent perianth coating the seed. A reduced and monophyllous cuplike perigonal bract, properly known as the bracteole, immediately surrounds and protects the seed and is often misnamed as a "calyx". Flowers are grouped to form
cymes In botany, an inflorescence is a group or cluster of flowers arranged on a plant's stem that is composed of a main branch or a system of branches. An inflorescence is categorized on the basis of the arrangement of flowers on a main axis ( ped ...
. In the dioecious plants the male inflorescences are long and look like
panicle In botany, a panicle is a much-branched inflorescence. (softcover ). Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers (and fruit) be pedicellate (having a single stem per flower). The branches of a p ...
s, while the female ones are shorter and bear fewer flowers. The
pistil Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl (botany), whorl of a flower; it consists ...
is made of two connate
carpel Gynoecium (; ; : gynoecia) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more ...
s, the usually superior ovary is unilocular; there is no fixed number of
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s. The fruit can be an
achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple fruit, simple dry fruits, dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and Dehi ...
or a
drupe In botany, a drupe (or stone fruit) is a type of fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pip'' (UK), ''pit'' (US), ''stone'', or ''pyrena'') of hardened endocarp with a seed ...
.


Taxonomy


Classification

Classification systems developed prior to the 1990s, such as those of Cronquist (1981) and Dahlgren (1989), typically recognized the order
Urticales Urticales is an order of flowering plants. Before molecular phylogenetics became an important part of plant taxonomy, Urticales was recognized in many, perhaps even most, systems of plant classification, with some variations in circumscription. Am ...
, which included the families Cannabaceae, Cecropiaceae, Celtidaceae, Moraceae, Ulmaceae and Urticaceae, as then circumscribed. Molecular data from 1990s onwards showed that these families were actually embedded within the order Rosales, so that from the first classification by the
Angiosperm Phylogeny Group The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disc ...
in 1998, they were placed in an expanded Rosales, forming a group which has been called 'urticalean rosids'. Cannabaceae comprises the following genera: * '' Aphananthe'' Planch. (5 spp.) * ''
Cannabis ''Cannabis'' () is a genus of flowering plants in the family Cannabaceae that is widely accepted as being indigenous to and originating from the continent of Asia. However, the number of species is disputed, with as many as three species be ...
'' —Hemp (3 spp.) * ''
Celtis ''Celtis'' is a genus of about 60–70 species of deciduous trees, commonly known as hackberries or nettle trees, in the hemp family Cannabaceae. It has a cosmopolitan distribution. Description ''Celtis'' species are generally medium-sized tree ...
'' L. (73–109 spp.) * '' Chaetachme'' Planch. (1 sp.) * '' Gironniera'' Gaudich. (6 spp.) * ''
Humulus ''Humulus'', or hop, is a small genus of flowering plants in the family Cannabaceae. The hop is native to temperate regions of the Northern Hemisphere. Hops are the female flowers (seed cones, strobiles) of the hop species '' H. lupulus''; as a ...
'' L.—Hop (3 spp.) * '' Lozanella'' Greenm. (2 spp.) * '' Parasponia'' Miq. (5–10 spp.) * ''
Pteroceltis ''Pteroceltis'' is a genus of small trees in the family Cannabaceae and containing the living species ''Pteroceltis tatarinowii''. The genus is now restricted to an endemic range in China and Mongolia. The genus has a fossil record which include ...
'' Maxim. (1 sp.) * ''
Trema Trema may refer to: * a Greek and Latin root meaning ''hole'' * Trema, a term for the two dots (diacritic) ** Tréma, (French), a diaeresis * ''Trema'' (plant), a genus of about 15 species of small evergreen trees * Tréma (record label), a Fr ...
'' Lour. (12–42 spp.)


Phylogeny

Cannabaceae likely originated in East Asia during the Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
. The oldest known pollen typical of members of Cannabaceae is from the Late Cretaceous (
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
~94–90 million years ago) of
Sarawak Sarawak ( , ) is a States and federal territories of Malaysia, state of Malaysia. It is the largest among the 13 states, with an area almost equal to that of Peninsular Malaysia. Sarawak is located in East Malaysia in northwest Borneo, and is ...
, Borneo. Fossils show Cannabaceae were widely distributed in the Northern Hemisphere during the early Cenozoic, though their distribution shifted towards tropical regions in the later Cenozoic due to changing climates. Modern
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
suggest the following relationships:


Uses

Carbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was ...
has revealed that these plants may have been used for ritual/medicinal purposes in Xinjiang, China as early as 494 B.C. ''
Humulus lupulus ''Humulus lupulus'', the common hop or hops, is a species of flowering plant in the hemp family, Cannabaceae. It is a perennial, herbaceous climbing plant which sends up new shoots in early spring and dies back to a cold-hardy rhizome in autumn. ...
'', the common hop, has been the predominant bittering agent of
beer Beer is an alcoholic beverage produced by the brewing and fermentation of starches from cereal grain—most commonly malted barley, although wheat, maize (corn), rice, and oats are also used. The grain is mashed to convert starch in the ...
for hundreds of years. The flowers' resins are responsible for beer's bitterness and their ability to extend shelf life due to some
antimicrobial An antimicrobial is an agent that kills microorganisms (microbicide) or stops their growth (bacteriostatic agent). Antimicrobial medicines can be grouped according to the microorganisms they are used to treat. For example, antibiotics are used aga ...
qualities. The young shoots can be used as a vegetable. Some plants in the genus ''Cannabis'' are cultivated as
hemp Hemp, or industrial hemp, is a plant in the botanical class of ''Cannabis sativa'' cultivars grown specifically for industrial and consumable use. It can be used to make a wide range of products. Along with bamboo, hemp is among the fastest ...
for the production of fiber, as a source of cheap
oil An oil is any nonpolar chemical substance that is composed primarily of hydrocarbons and is hydrophobic (does not mix with water) and lipophilic (mixes with other oils). Oils are usually flammable and surface active. Most oils are unsaturate ...
, for their nutritious seeds, or their edible leaves. Others are cultivated for medical or recreational use as dried flowers,
extracts An extract (essence) is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol, oil or water. Extracts may be sold as tinctures or absolutes or dried and powdered. The aromatic principles of man ...
, or infused food products. Induced
parthenocarpy In botany and horticulture, parthenocarpy is the natural or artificially induced production of fruit without fertilisation of ovules, which makes the fruit seedless. The phenomenon has been observed since ancient times but was first scientificall ...
in pistilate flowers, and
selective breeding Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant m ...
are used to produce either higher or lower yields of
tetrahydrocannabinol Tetrahydrocannabinol (THC) is a cannabinoid found in cannabis. It is the principal psychoactive constituent of ''Cannabis'' and one of at least 113 total cannabinoids identified on the plant. Although the chemical formula for THC (C21H30O2) de ...
(THC), other
cannabinoid Cannabinoids () are several structural classes of compounds found primarily in the ''Cannabis'' plant or as synthetic compounds. The most notable cannabinoid is the phytocannabinoid tetrahydrocannabinol (THC) (delta-9-THC), the primary psychoact ...
s, as well as
terpenes Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n ≥ 2. Terpenes are major biosynthetic building blocks. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predomi ...
with desired flavors or aromas, such as blueberry, strawberry, or even citrus. Many trees in the genus ''Celtis'' are grown for landscaping and ornamental purposes, and the bark of
Pteroceltis ''Pteroceltis'' is a genus of small trees in the family Cannabaceae and containing the living species ''Pteroceltis tatarinowii''. The genus is now restricted to an endemic range in China and Mongolia. The genus has a fossil record which include ...
is used to produce high-end Chinese paper.


References


External links


Cannabaceae of Mongolia in FloraGREIF
{{Authority control Rosid families Extant Turonian first appearances