Cannabaceae

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids I
Ordoo: Rosales

Familia: Cannabaceae
Genera: Aphananthe – Cannabis – Celtis – Chaetachme – Gironniera – Humulopsis – Humulus – Lozanella – Parasponia – Pteroceltis – Trema

Name

Cannabaceae Martynov, Tekhno-Bot. Slovar 99. 1820 [3 Aug 1820] , as 'Cannabinae' (1820).
Synonyms

Heterotypic
Celtidaceae Endl., Ench. Bot.: 171 (1841) [15–21 Aug 1841].
Typus: Celtis L.
Humulaceae Berchtold & J.Presl, Přir. Rostlin: 258 (1820) [Jan–Apr 1820].
Typus: Humulus L.
Lupulaceae Schultz Sch., Nat. Syst. Pflanzenr.: 370 (1832) [30 Jan–10 Feb 1832]
Typus: Lupulus Mill.

References
Primary references

Martinov, I.I. 1820. Техно-ботаническiй словарь, на латинскомъ и россiйскомъ языкахъ, составленный […]. [Techno-Botanical Dictionary, in Latin and Russian, compiled by [...]]. i-vi, 682 p. St.-Petersburg: v tipografii Imperatorskoj Rossijskoj Akademii [Nauk] (Russian) DOI: 10.5962/bhl.title.96260 Reference page.

Additional references

Spitaler, R., Gurschler, S., Ellmerer, E., Schubert, B., Sgarbossa, M. & Zidorn, C. 2008. Flavonoids from Celtis australis (Cannabaceae). Biochemical Systematics and Ecology 37(2): 120–121. DOI: 10.1016/j.bse.2008.11.020

Links

Govaerts, R. et al. 2019. Cannabaceae in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2019 Aug. 8. Reference page.
International Plant Names Index. 2019. 10. Published online. Accessed: Aug. 2019.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page.

Vernacular names
العربية: قنبية
azərbaycanca: Kənafkimilər
беларуская: Каноплевыя
български: конопени
català: Cannabàcies
čeština: Konopovité
dansk: Hamp-familien
Deutsch: Hanfgewächse
English: Hemp family
Esperanto: Kanabacoj
español: Cannabáceas
eesti: Kanepilised
فارسی: شاهدانگان
suomi: Hamppukasvit
Nordfriisk: Henepplaanten
français: Cannabinacée
עברית: קנאביים
hrvatski: Konopljovke
hornjoserbsce: Konopowe rostliny
magyar: Kenderfélék
íslenska: Humlaætt
日本語: アサ科
ქართული: კანაფისებრნი
한국어: 삼과
lietuvių: Kanapiniai
latviešu: Kaņepju dzimta
македонски: Конопи
മലയാളം: കന്നാബേസീ
монгол: Олсныхон
Nederlands: Hennepfamilie
norsk: Hampfamilien
polski: Konopiowate
پنجابی: بھنگ ٹبر
português: Canabiáceas
Runa Simi: Kañamu yura rikch'aq ayllu
română: Canabacee
русский: Коноплёвые
slovenčina: konopovité
svenska: Hampväxter
ไทย: วงศ์กัญชา
Türkçe: Kendirgiller
татарча/tatarça: Киндерчәләр
українська: Коноплеві
oʻzbekcha/ўзбекча: Nashadoshlar
Tiếng Việt: Họ Gai dầu
中文: 大麻科

Cannabaceae is a small family of flowering plants, known as the hemp family. As now circumscribed, the family includes about 170 species grouped in about 11 genera, including Cannabis (hemp) , Humulus (hops) and Celtis (hackberries). Celtis is by far the largest genus, containing about 100 species.[1]

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.[3]

Other than a shared evolutionary origin, members of the family have few common characteristics; some are trees (e.g. Celtis), others are herbaceous plants (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).[1]

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 masculine inflorescences are long and look like panicles, while the feminine 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 stamens.

The fruit can be an achene or a drupe.

Evolutionary history

Cannabaceae likely originated in East Asia during the Late Cretaceous. The oldest known pollen typical of members of Cannabaceae is from the Late Cretaceous (Turonian ~94-90 million years ago) of Sarawak, 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.[4]
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".[2]
Genera
Humulus lupulus with nearly mature flowers (hops)

Cannabaceae comprises the following genera:[5][6][7]

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

Modern molecular phylogenetics suggest the following relationships:[2][8][9][5]
Cladogram of Cannabaceae genera
 
 

Moraceae (outgroup)   Cannabaceae   Aphananthe       Gironniera       Lozanella         Cannabis     Humulus         Celtis         Pteroceltis     Chaetachme       Trema (including Parasponia)

 
 
 
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.[10]

Hop (Humulus lupulus) 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 anti-microbial qualities. The young shoots are used as vegetable.

Some plants in the genus Cannabis are cultivated as hemp 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 rice paper.
References

Stevens, P.F. (2001 onwards) "Cannabaceae", Angiosperm Phylogeny Website, retrieved 2014-02-25
Sytsma, Kenneth J.; Morawetz, Jeffery; Pires, J. Chris; Nepokroeff, Molly; Conti, Elena; Zjhra, Michelle; Hall, Jocelyn C. & Chase, Mark W. (2002), "Urticalean rosids: Circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL–F, and ndhF sequences", Am J Bot, 89 (9): 1531–1546, doi:10.3732/ajb.89.9.1531, PMID 21665755, S2CID 207690258
"Cannabaceae | plant family".
Jin, Jian‐Jun; Yang, Mei‐Qing; Fritsch, Peter W.; Velzen, Robin; Li, De‐Zhu; Yi, Ting‐Shuang (July 2020). "Born migrators: Historical biogeography of the cosmopolitan family Cannabaceae". Journal of Systematics and Evolution. 58 (4): 461–473. doi:10.1111/jse.12552. ISSN 1674-4918.
Yang M-Q, van Velzen R, Bakker FT, Sattarian A, Li D-Z, Yi T-S. (2013). "Molecular phylogenetics and character evolution of Cannabaceae". Taxon. 62 (3): 473–485. doi:10.12705/623.9.
Stevens PF. (2017). "Cannabaceae Genera". Angiosperm Phylogeny Website. Retrieved 4 April 2017.
"!!Cannabaceae Martinov". Tropicos. 2017. Retrieved 4 April 2017.
Zavada MS, Kim M. (1996). "Phylogenetic analysis of Ulmaceae". Plant Syst Evol. 200 (1): 13–20. doi:10.1007/BF00984745.
Yesson C, Russell SJ, Parrish T, Dalling JW, Garwood NC. (2004). "Phylogenetic framework for Trema (Celtidaceae)". Plant Syst Evol. 248 (1): 85–109. doi:10.1007/s00606-004-0186-3.
Jiang, Hong-En; Li, Xiao; Zhao, You-Xing; Ferguson, David K.; Hueber, Francis; Bera, Subir; Wang, Yu-Fei; Zhao, Liang-Cheng; Liu, Chang-Jiang & Li, Cheng-Sin (December 2006), "A new insight into Cannabis sativa (Cannabaceae) utilization from 2500-year-old Yanghai Tombs, Xinjiang, China", Journal of Ethnopharmacology, 108 (3): 414–422, doi:10.1016/j.jep.2006.05.034, PMID 16879937