Zanthoxylum

Zanthoxylum is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the family Rutaceae that are native to warm temperate and subtropical areas worldwide. It is the type genus of the tribe Zanthoxyleae in the subfamily Rutoideae. Several of the species have yellow heartwood, to which their generic name alludes.[3]

Zanthoxylum
Zanthoxylum kauaense
Scientific classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Sapindales
Family: Rutaceae
Subfamily: Zanthoxyloideae
Genus: Zanthoxylum
L.
Type species
Zanthoxylum americanum
Synonyms[2]
List

Description

Plants in the genus Zanthoxylum are typically dioecious shrubs, trees or woody climbers armed with trichomes. The leaves are arranged alternately and are usually pinnate or trifoliate. The flowers are usually arranged in panicles and usually function as male or female flowers with four sepals and four petals, the sepals remaining attached to the fruit. Male flowers have four stamens opposite the sepals. Female flowers have up to five, more or less free carpels with the styles free or sometimes fused near the tip. The fruit is usually of up to four follicles fused at the base, each containing a single seed almost as large as the follicle.[4][5]

Taxonomy

The genus Zanthoxylum was first formally described in 1753 by Carl Linnaeus in the first volume of Species Plantarum.[6][7] The generic name is derived from Ancient Greek words ξανθός (xanthos), meaning "yellow," and ξύλον (xylon), meaning "wood." It is technically misspelled, as the z should be x, but botanical nomenclature does not allow for spelling corrections. It refers to a yellow dye made from the roots of some species.[8] The first species that Linnaeus described was Zanthoxylum trifoliatum, now regarded as a synonym of Eleutherococcus trifoliatus.[7][9] The once separate genus Fagara is now included in Zanthoxylum.[10]

Species list

The following is a list of species accepted by Plants of the World Online as of August 2020:[2]

Zanthoxylum clava-herculis fruits and foliage
Z. piperitum Fruits and seeds
Z. rhetsa bark in Pakke Tiger Reserve
  • Zanthoxylum nadeaudii Drake (French Polynesia)
  • Zanthoxylum nashii P.Wilson (Cuba to Hispaniola)
  • Zanthoxylum nebuletorum (Herzog) P.G.Waterman (Bolivia)
  • Zanthoxylum nemorale Mart. (eastern Brazil)
  • Zanthoxylum neocaledonicum Baker f. (New Caledonia)
  • Zanthoxylum nigrum Mart. (Brazil)
  • Zanthoxylum nitidum (Roxb.) DC. – shiny-leaf prickly ash (tropical and subtropical Asia, Queensland)
  • Zanthoxylum novoguineense T.G.Hartley (New Guinea)
  • Zanthoxylum oahuense Hillebr. – Oahu prickly ash (Hawaii)
  • Zanthoxylum ovalifolium Wight (Andaman Island, Bangladesh, Myanmar, Queensland)
  • Zanthoxylum ovatifoliolatum (Engl.) Finkelstein (Angola to north-west Namibia)
  • Zanthoxylum oxyphyllum Edgew. (Nepal to China and Myanmar)
  • Zanthoxylum panamense P.Wilson (Mexico to central America)
  • Zanthoxylum pancheri P.S.Green (New Caledonia)
  • Zanthoxylum paniculatum Balf.f. (Rodrigues in Mauritius)
  • Zanthoxylum paracanthum (Mildbr.) Kokwaro (south-east Kenya to central Tanzania)
  • Zanthoxylum paulae (Albuq.) P.G. Waterman (Guianas to northern Brazil)
  • Zanthoxylum pentandrum (Aubl.) R.A.Howard (Trinidad to northern Brazil)
  • Zanthoxylum petenense Lundell (Mexico to Guatemala)
  • Zanthoxylum petiolare A.St.-Hil. & Tul. (Bolivia to Brazil and northern Argentina)
  • Zanthoxylum piasezkii Maxim. (China)
  • Zanthoxylum pilosiusculum (Engl.) P.G.Waterman (Democratic Republic of Congo to Angola)
  • Zanthoxylum pilosulum Rehder & E.H.Wilson (south-central China)
  • Zanthoxylum pimpinelloides (Lam.) DC. (Cuba to Hispaniola)
  • Zanthoxylum pinnatum (J.R.Forst. & G.Forst.) W.R.B.Oliv. (Norfolk Island, Lord Howe Island, south-west Pacific.)
  • Zanthoxylum piperitum (L.) DC. (China to South Korea, Japan)
  • Zanthoxylum phyllopterum (Griseb.) Wright (Cuba)
  • Zanthoxylum piasezkii Maxim.
  • Zanthoxylum pilosiusculum (Engl.) P.G.Waterm.
  • Zanthoxylum pilosulum Rehder & E.H. Wilson
  • Zanthoxylum pimpinelloides (Lam.) DC.
  • Zanthoxylum pinnatum Druce
  • Zanthoxylum piperitum (L.) DC. – Japanese pepper, chopi, Korean pepper (China, Japan, Korea)
  • Zanthoxylum pluviatile T.G.Hartley (New Guinea and Solomon Islands)
  • Zanthoxylum poggei (Engl.) P.G.Waterman (western central tropical Africa)
  • Zanthoxylum psammophilum (Aké Assi) P.G.Waterman (Ivory Coast)
  • Zanthoxylum pseudoxyphyllum Babu (Ivory Coast)
  • Zanthoxylum pteracanthum Rehder & E.H.Wilson (China)
  • Zanthoxylum pucro D.M Porter (Panama to north-west Colombia)
  • Zanthoxylum punctatum Vahl – dotted prickly ash (Caribbean)
  • Zanthoxylum quassiifolium (Donn.Sm.) Standl. & Steyerm. (Mexico to Guatemala.)
  • Zanthoxylum quinduense Tul. (western South America to north-west Venezuela)
  • Zanthoxylum renieri (G.C.C.Gilbert) P.G.Waterman (Democratic Republic of Congo to Uganda)
  • Zanthoxylum retroflexum T.G.Hartley (Sumatra)
  • Zanthoxylum retusum (Albuq.) P.G.Waterman (Brazil)
  • Zanthoxylum rhetsa (Roxb.) DC. – Cape yellowwood, Indian ivy-rue, Indian pepper (tropical Asia, Queensland)
  • Zanthoxylum rhodoxylum (Urb.) P.Wilson (Jamaica)
Leafless Z. simulans showing its knobbed bark
  • Zanthoxylum rhoifolium Lam. (tropical and subtropical America)
  • Zanthoxylum rhombifoliolatum C.C.Huang (China)
  • Zanthoxylum riedelianum Engl. (Brazil to north-east Argentina)
  • Zanthoxylum rigidum Humb. & Bonpl. ex Willd. (western South America to Paraguay)
  • Zanthoxylum rubescens Planch. ex Hook. (western tropical Africa to south-west Kenya and Angola)
  • Zanthoxylum sapindifolium Wall. (Myanmar)
  • Zanthoxylum sarasinii Guillaumin (New Caledonia)
  • Zanthoxylum scandens Blume (southern China to Nansei-shoto and western Malesia)
  • Zanthoxylum schinifolium Philipp Franz von SieboldSiebold & Zucc. – mastic-leaf prickly ash, Sichuan pepper (central and eastern China to temperate eastern Asia)
  • Zanthoxylum schlechteri Guillaumin (New Caledonia)
  • Zanthoxylum schreberi (J.F.Gmel.) Reynel ex C.Nelson (Mexico to Venezuela and Bolivia, Caribbean)
  • Zanthoxylum setulosum P.Wilson (Costa Rica to north-west Venezuela)
  • Zanthoxylum simulans Hance (China, South Korea, Taiwan)
  • Zanthoxylum spinosum (Sw.) Sw. (Florida to Caribbean)
  • Zanthoxylum spondiifolium Wall. (Myanmar)
  • Zanthoxylum sprucei Engl. (Ecuador to Peru and western Brazil)
  • Zanthoxylum stelligerum Turcz. (central and eastern Brazil)
  • Zanthoxylum stenophyllum Hemsl. (central and eastern central China)
  • Zanthoxylum subspicatum H. Perrier (Madagascar)
  • Zanthoxylum syncarpum (Tul.) Tul. ex B.D.Jacks. (Venezuela, eastern Brazil)
  • Zanthoxylum taediosum A.Rich. (Cuba)
  • Zanthoxylum tahitense (Nadeaud) J.Florence & N.Hallé (Society Islands)
  • Zanthoxylum tambopatense Reynel (Peru)
  • Zanthoxylum tetraphyllum (urb. & Ekman) J.Jiménez Alm. (Haiti)
  • Zanthoxylum tetraspermum Wight & Arn. (southern India, Sri Lanka)
  • Zanthoxylum thomense (Engl.) A.Chev. ex P.G.Waterman (Gabon and Gulf of Guinea Island)
  • Zanthoxylum thornycroftii (I.Verd.) P.G.Waterman (Limpopo to Mpumalanga)
  • Zanthoxylum thouventii H.Perrier (Madagascar])
  • Zanthoxylum tidorense Miq. (Maluku Islands)
  • Zanthoxylum timoriense Span. (Lesser Sunda Islands)
  • Zanthoxylum tingoassuiba A.St.-Hil. (eastern and southern Brazil)
  • Zanthoxylum tomentellum Hook.f. (Nepal to China and Myanmar)
  • Zanthoxylum tragodes (L.) DC. – Niaragato (Caribbean)
  • Zanthoxylum trijugum (Dunkley) P.G.Waterman (central Tanzania to Zambia)
  • Zanthoxylum tsihanimposa H.Perrier (Madagascar)
  • Zanthoxylum undulatifolium Hemsl. (China)
  • Zanthoxylum unifoliatum Groppo & Pirani (Brazil)
  • Zanthoxylum usambarense (Engl.) Kokwara (Ethiopia to Rwanda)
  • Zanthoxylum usitatum Pierre ex Laness. (Vietnam)
  • Zanthoxylum venosum Leonard (Hispaniola)
  • Zanthoxylum verrucosum (Cuatrec.) P.G.Waterman (Colombia to Ecuador)
  • Zanthoxylum vinkii T.G.Hartley (New Guinea)
  • Zanthoxylum viride (A.Chev.) T.G.Hartley (west tropical Africa to Congo)
  • Zanthoxylum vitiense A.C.Sm. (Fiji)
  • Zanthoxylum wutaiense I.S.Chen (Taiwan)
  • Zanthoxylum xichouense C.C.Huang (China)
  • Zanthoxylum yakumontanum (Sugim.) Nagam. (Japan)
  • Zanthoxylum yuanjiangensis C.C.Huang (China)
  • Zanthoxylum zanthoxyloides (Lam.) Zepern. & Timler – Senegal prickly ash (western tropical Africa)
Z. piperitum as a bonsai

Doubtful species

The genus Fagara has been sunk into Zanthoxylum, but as of September 2021, no name seemed to have been provided for the former Fagara externa, which was regarded as an unplaced name by Plants of the World Online.[11]

Fossil record

28 fossil seeds of †Zanthoxylum kristinae from the early Miocene, have been found in the Kristina Mine at Hrádek nad Nisou in North Bohemia, the Czech Republic.[12]

Uses

Many Zanthoxylum species make excellent bonsai and in temperate climates they can be grown quite well indoors. Zanthoxylum beecheyanum and Zanthoxylum piperitum are two species commonly grown as bonsai.[13]

Culinary use
See also: Sichuan pepper

The fruit of several species is used to make the spice Sichuan pepper. They are also used as bonsai trees. Historically, the bark was used in traditional medicine.[13]

Spices are made from a number of species in this genus, especially Zanthoxylum piperitum, Z. simulans, Z. bungeanum, Z. schinifolium Z. nitidum, Z. rhetsa, Z. alatum, and Z. acanthopodium. Sichuan pepper is most often made by grinding the husks that surround Z. piperitum berries.[14] In the states of Maharashtra, Karnataka, and Goa in Western India, the berries of Z. rhetsa are sun-dried and added to foods such as legumes and fish. Because the trees bear fruit during the monsoon season, the berries are associated with the concurrent Krishna Janmashtami festival.[15]

Chemistry

Plants in the genus Zanthoxylum contain the lignan sesamin.

Species identified in Nigeria contains several types of alkaloids including benzophenanthridines (nitidine, dihydronitidine, oxynitidine, fagaronine, dihydroavicine, chelerythrine, dihydrochelerythrine, methoxychelerythrine, norchelerythrine, oxychelerythrine, decarine and fagaridine), furoquinolines (dictamine, 8-methoxydictamine, skimmianine, 3-dimethylallyl-4-methoxy-2-quinolone), carbazoles (3-methoxycarbazole, glycozoline), aporphines (berberine, tembetarine,[16] magnoflorine, M-methyl-corydine), canthinones (6-canthinone), acridones (1-hydroxy-3-methoxy-10-methylacridon-9-one, 1-hydroxy-10-methylacridon-9-one, zanthozolin), and aromatic and aliphatic amides.[17] Hydroxy-alpha sanshool is a bioactive component of plants from the genus Zanthoxylum, including the Sichuan pepper.

Ecology

Zanthoxylum species are used as food plants by the larvae of some Lepidoptera species including the engrailed (Ectropis crepuscularia).

References
  1. "Zanthoxylum L." TROPICOS. Missouri Botanical Garden. Archived from the original on 2010-09-10. Retrieved 2010-02-26.
  2. "Zanthoxylum". Plants of the World Online. Retrieved 16 August 2020.
  3. Thomas, Val; Grant, Rina (2001). Sappi tree spotting: Highlands: Highveld, Drakensberg, Eastern Cape mountains. illustrations: Joan van Gogh; photographs: Jaco Adendorff (3rd ed.). Johannesburg: Jacana. p. 260. ISBN 978-1-77009-561-8.
  4. "Genus Zanthoxylum". Royal Botanic Garden Sydney. Retrieved 17 August 2020.
  5. Hartley, Thomas G.; Wilson, Annette J.G. (ed.) (2013). Flora of Australia (Volume 26). Canberra: Australian Biological Resources Study. p. 74. Retrieved 17 August 2020.CS1 maint: extra text: authors list (link)
  6. "Zanthoxylum". APNI. Retrieved 17 August 2020.
  7. Linnaeus, Carl (1753). Species Plantarum. Berlin: Junk, 1908. p. 270. Retrieved 17 August 2020.
  8. Quattrocchi, Umberto (2000). CRC World Dictionary of Plant Names. IV R-Z. Taylor & Francis US. p. 2868. ISBN 978-0-8493-2678-3.
  9. "Zanthoxylum trifoliatum". Plants of the World Online. Retrieved 17 August 2020.
  10. Beurton, C. (1994). "Gynoecium and perianth in Zanthoxylum s.l. (Rutaceae)". Plant Systematics and Evolution. 189 (3–4): 165–191. doi:10.1007/bf00939724. S2CID 2655415.
  11. "Fagara externa Skottsb". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2021-09-13.
  12. A review of the early Miocene Mastixioid flora of the Kristina Mine at Hrádek nad Nisou in North Bohemia, The Czech Republic, January 2012 by F. Holý, Z. Kvaček and Vasilis Teodoridis - ACTA MUSEI NATIONALIS PRAGAE Series B – Historia Naturalis • vol. 68 • 2012 • no. 3–4 • pp. 53–118
  13. Wilbur, C. Keith, MD. Revolutionary Medicine 1700-1800. The Globe Pequot Press. Page 23. 1980.
  14. Peter, K. V. (2004). Handbook of Herbs and Spices. 2. Woodhead Publishing. pp. 98–99. ISBN 978-1-85573-721-1.
  15. Bharadwaj, Monisha (2006). Indian Spice Kitchen. Hippocrene Books. pp. 82–83. ISBN 978-0-7818-1143-9.
  16. "{title}". Archived from the original on 2017-12-17. Retrieved 2017-05-18.
  17. The Nigerian Zanthoxylum; Chemical and biological values. S. K. Adesina, Afr. J. Trad. CAM, 2005, volume 2, issue 3, pages 282-301 (article Archived 2016-03-03 at the Wayback Machine)
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