Cannabaceae
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Cannabaceae is a small family 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'' (hemp), '' Humulus'' (
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'' (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. 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''), erect herbs (e.g. '' Cannabis''), or twining herbs (e.g. '' Humulus''). Leaves are often more or less palmately lobed or palmately compound and always bear stipules. Cystoliths are always present and some members of this family possess laticifers. Cannabaceae are often dioecious (distinct male and female plants). The flowers are actinomorphic (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 the dioecious plants the male inflorescences are long and look like panicles, while the female ones are shorter and bear fewer flowers. The pistil is made of two connate carpels, 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 or a drupe.


Taxonomy


Classification

Classification systems developed prior to the 1990s, such as those of Cronquist (1981) and Dahlgren (1989), typically recognized the order Urticales, 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 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'' —Hemp (3 spp.) * '' Celtis'' L. (73–109 spp.) * '' Chaetachme'' Planch. (1 sp.) * '' Gironniera'' Gaudich. (6 spp.) * '' Humulus'' L.—Hop (3 spp.) * '' Lozanella'' Greenm. (2 spp.) * '' Parasponia'' Miq. (5–10 spp.) * '' Pteroceltis'' Maxim. (1 sp.) * '' Trema'' 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 has revealed that these plants may have been used for ritual/medicinal purposes in Xinjiang, China as early as 494 B.C. '' Humulus lupulus'', the common hop, has been the predominant bittering agent of beer for hundreds of years. The flowers' resins are responsible for beer's bitterness and their ability to extend shelf life due to some antimicrobial 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, for their nutritious seeds, or their edible leaves. Others are cultivated for medical or recreational use as dried flowers, extracts, or infused food products. Induced parthenocarpy in pistilate flowers, and selective breeding are used to produce either higher or lower yields of tetrahydrocannabinol (THC), other cannabinoids, as well as terpenes 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 is used to produce high-end Chinese paper.


References


External links


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