Lamiaceae

Lamiaceae
	Lamium purpureum L.
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Lamiales
Family:	Lamiaceae; Martinov
Type genus
	Lamium; L.
Genera
	See § Genera

The Lamiaceae (/ˌleɪmiˈeɪsi.iː, -ˌaɪ/ LAY-mee-AY-see-ee, -⁠eye)^[3] or Labiatae are a family of flowering plants commonly known as the mint, deadnettle, or sage family. Many of the plants are aromatic in all parts and include widely used culinary herbs like basil, mint, rosemary, sage, savory, marjoram, oregano, hyssop, thyme, lavender, and perilla, as well as other medicinal herbs such as catnip, salvia, bee balm, wild dagga, and oriental motherwort. Some species are shrubs, trees (such as teak), or, rarely, vines. Many members of the family are widely cultivated, not only for their aromatic qualities, but also their ease of cultivation, since they are readily propagated by stem cuttings.^{[citation needed]} Besides those grown for their edible leaves, some are grown for decorative foliage. Others are grown for seed, such as Salvia hispanica (chia), or for their edible tubers, such as Plectranthus edulis, Plectranthus esculentus, Plectranthus rotundifolius, and Stachys affinis (Chinese artichoke). Many are also grown ornamentally, notably coleus, Plectranthus, and many Salvia species and hybrids.

The family has a cosmopolitan distribution.^[4] The enlarged Lamiaceae contain about 236 genera^[5] and have been stated to contain 6,900^[4] to 7,200^[5] species, but the World Checklist lists 7,534.^[6] The largest genera are Salvia (900), Scutellaria (360), Stachys (300), Plectranthus (300), Hyptis (280), Teucrium (250), Vitex (250), Thymus (220), and Nepeta (200).^[5] Clerodendrum was once a genus of over 400 species,^[5] but by 2010, it had been narrowed to about 150.^[7]

The family has traditionally been considered closely related to the Verbenaceae;^[5] in the 1990s, phylogenetic studies suggested that many genera classified in the Verbenaceae should be classified in the Lamiaceae ^[8]^[9] or to other families in the order Lamiales.^[1]

The alternative family name Labiatae refers to the flowers typically having petals fused into an upper lip and a lower lip (labia in Latin). Although this is still considered an acceptable alternative name, most botanists now use the name Lamiaceae in referring to this family. The flowers are bilaterally symmetrical with five united petals and five united sepals. They are usually bisexual and verticillastrate (a flower cluster that looks like a whorl of flowers, but actually consists of two crowded clusters). The leaves emerge oppositely, each pair at right angles to the previous one (decussate) or whorled. The stems are frequently square in cross section,^[10] but this is not found in all members of the family, and is sometimes found in other plant families.

Genera

The last revision of the entire family was published in 2004.^[5] It described and provided keys to 236 genera. These are marked with an asterisk (*) in the list below. A few genera have been established or resurrected since 2004. These are marked with a plus sign (+). Other genera have been synonymised. These are marked with a minus sign (-). The remaining genera in the list are mostly of historical interest only and are from a source that includes such genera without explanation.^[11] Few of these are recognized in modern treatments of the family.

Kew Gardens provides a list of genera that includes additional information.^[12] A list at the Angiosperm Phylogeny Website is frequently updated.^[13] Plants of the World Online currently accepts 227 genera.^[14]

*Acanthomintha (A.Gray) A.Gray
Acanthoprasium (Benth.) Spach
*Achyrospermum Blume
Acinos Mill., synonym of Clinopodium
Acrocephalus Benth., synonym of Platostoma
*Acrotome Benth. ex Endl.
*Acrymia Prain
Adelosa Blume, synonym of Clerodendrum
*Aegiphila Jacq.
*Aeollanthus C.Mart. ex Spreng.
*Agastache Clayton ex Gronov.
*Ajuga L.
*Ajugoides Makino
*Alajja Ikonn., synonym of Eriophyton
*Alvesia Welw.
*Amasonia L.f.
*Amethystea L.
*Anisochilus Wall. ex Benth., synonym of Coleus
Apatelantha T.C.Wilson & Henwood
*Anisomeles R.Br.
*Asterohyptis Epling
*Ballota L.
*Basilicum Moench
Becium Lindl., synonym of Ocimum
*Benguellia G.Taylor
Betonica L.
*Blephilia (L.) Raf.
*Bostrychanthera Benth., synonym of Chelonopsis
Bovonia Chiov., synonym of Aeollanthus
*Brachysola Rye
*Brazoria Engelm. & A.Gray
*Bystropogon L'Hér.
Calamintha L., synonym of Clinopodium
*Callicarpa L.
Cantinoa Harley & J.F.B.Pastore
*Capitanopsis S.Moore
Capitanya Schweinf. ex Gürke, synonym of Coleus
*Caryopteris Bunge
*Catoferia (Benth.) Benth.
*Cedronella Moench
Ceratanthus F.Muell. ex G.Taylor, synonym of Platostoma
*Chaiturus Ehrh. ex Willd.
*Chamaesphacos Schrenk ex Fisch. & C.A.Mey.
*Chaunostoma Donn.Sm., synonym of Lepechinia
*Chelonopsis Miq.
*Chloanthes R.Br.
*Cleonia L.
*Clerodendrum L.
*Clinopodium L.
*Colebrookea Sm.
Coleus Lour.
*Collinsonia L.
*Colquhounia Wall.
*Comanthosphace S.Moore
Condea Adans.
*Congea Roxb.
*Conradina A.Gray
Coridothymus Rchb.f., synonym of Thymbra
*Cornutia Plum. ex L.
*Craniotome Rchb.
*Cryphia R.Br., synonym of Prostanthera
*Cuminia Colla
*Cunila D.Royen
Cyanocephalus (Pohl ex Benth.) Harley & J.F.B.Pastore
*Cyanostegia Turcz.
*Cyclotrichium (Boiss.) Manden. & Scheng.
*Cymaria Benth.
Dasymalla Endl.
*Dauphinea Hedge, synonym of Capitanopsis
*Dicerandra Benth.
*Dicrastylis Drumm. ex Harv.
Discretitheca P.D.Cantino
*Dracocephalum L.
*Drepanocaryum Pojark.
*Elsholtzia Willd.
*Endostemon N.E.Br.
Englerastrum Briq., synonym of Coleus
+Eplingiella Harley & J.F.B.Pastore
Equilabium Mwany., A.J.Paton & Culham
*Eremostachys Bunge, synonym of Phlomoides
*Eriope Humb. & Bonpl. ex Benth.
*Eriophyton Benth.
Eriopidion Harley
*Eriothymus (Benth.) Rchb.
Erythrochlamys Gürke, synonym of Ocimum
Euhesperida Brullo & Furnari, synonym of Satureja
*Eurysolen Prain
*Faradaya F.Muell., synonym of Oxera
*Fuerstia T.C.E.Fr.
*Galeopsis L.
*Garrettia H.R.Fletcher
Geniosporum Wall. ex Benth., synonym of Platostoma
*Glechoma L.
*Glechon Spreng.
*Glossocarya Wall. ex Griff.
*Gmelina L.
*Gomphostemma Wall. ex Benth.
*Gontscharovia Boriss.
Gymneia (Benth.) Harley & J.F.B.Pastore
*Hanceola Kudô
*Haplostachys (A.Gray) Hillebr.
*Haumaniastrum P.A.Duvign. & Plancke
*Hedeoma Pers.
Heliacria Bo Li, C.L.Xiang, T.S.Hoang & Nuraliev
*Hemiandra R.Br.
*Hemigenia R.Br.
*Hemiphora (F.Muell.) F.Muell.
*Hemizygia (Benth.) Briq., synonym of Syncolostemon
*Hesperozygis Epling
*Heterolamium C.Y.Wu
*Hoehnea Epling
*Holmskioldia Retz.
*Holocheila (Kudô) S.Chow
Holostylon Robyns & Lebrun, synonym of Coleus
*Horminum L.
*Hosea Ridl.
*Hoslundia Vahl
*Huxleya Ewart, synonym of Volkameria
*Hymenocrater Fisch. & C.A.Mey., synonym of Nepeta
*Hymenopyramis Wall. ex Griff.
*Hypenia (Mart. ex Benth.) Harley
*Hypogomphia Bunge
*Hyptidendron Harley
*Hyptis Jacq.
*Hyssopus L., synonym of Dracocephalum
Isodictyophorus Briq., synonym of Coleus
*Isodon (Benth.) Schrad. ex Spach
*Isoleucas O.Schwartz
+Kalaharia Baill.
*Karomia Dop
Keiskea Miq., synonym of Collinsonia
Killickia Bräuchler, Heubl & Doroszenko
Kudrjaschevia Pojark.
*Kurzamra Kuntze
*Lachnostachys Hook.
*Lagochilus Bunge ex Benth.
*Lagopsis (Bunge ex Benth.) Bunge
*Lallemantia Fisch. & C.A.Mey., synonym of Dracocephalum
*Lamiophlomis Kudô, synonym of Phlomoides
*Lamium L.
*Lavandula L.
*Leocus A.Chev., synonym of Coleus
*Leonotis (Pers.) R.Br.
*Leonurus L.
*Lepechinia Willd.
Leptohyptis Harley & J.F.B.Pastore
*Leucas R.Br.
Leucophae Webb & Berthel., synonym of Sideritis
*Leucosceptrum Sm.
Limniboza R.E.Fr., synonym of Platostoma
*Lophanthus Adans., synonym of Nepeta
*Loxocalyx Hemsl.
*Lycopus L.
*Macbridea Elliott ex Nutt.
*Madlabium Hedge, synonym of Capitanopsis
*Marmoritis Benth., synonym of Nepeta
*Marrubium L.
*Marsypianthes Mart. ex Benth.
+Martianthus Harley & J.F.B.Pastore
*Matsumurella Makino
*Meehania Britton
*Melissa L.
*Melittis L.
*Mentha L.
*Meriandra Benth., synonym of Salvia
Mesona Blume, synonym of Platostoma
Mesosphaerum P.Browne
*Metastachydium Airy Shaw ex C.Y.Wu & H.W.Li, synonym of Phlomoides
*Microcorys R.Br.
*Micromeria Benth.
*Microtoena Prain
*Minthostachys (Benth.) Spach
*Moluccella L.
*Monarda L.
*Monardella Benth.
*Monochilus Fisch. & C.A.Mey.
*Mosla (Benth.) Buch.-Ham. ex Maxim.
Muniria N.Streiber & B.J.Conn
Neohyptis J.K.Morton, synonym of Coleus
Neorapinia Moldenke, synonym of Vitex
*Nepeta L.
*Newcastelia F.Muell.
Nosema Prain, synonym of Platostoma
*Notochaete Benth., synonym of Phlomoides
*Obtegomeria Doroszenko & P.D.Cantino
*Ocimum L.
Octomeron Robyns, synonym of Platostoma
*Ombrocharis Hand.-Mazz.
*Oncinocalyx F.Muell., synonym of Teucrium
Oocephalus (Benth.) Harley & J.F.B.Pastore
*Origanum L.
*Orthosiphon Benth.
*Otostegia Benth.
+Ovieda L.
*Oxera Labill.
*Panzerina Soják
*Paralamium Dunn
*Paraphlomis Prain
*Paravitex H.R.Fletcher, synonym of Vitex
*Peltodon Pohl, synonym of Hyptis
*Pentapleura Hand.-Mazz.
*Perilla L.
*Perillula Maxim.
*Peronema Jack
-Perovskia Kar., synonym of Salvia
Perrierastrum Guillaumin, synonym of Capitanopsis
Petitia Jacq.
*Petraeovitex Oliv.
*Phlomidoschema (Benth.) Vved.
*Phlomis L.
*Phlomoides Moench
*Phyllostegia Benth.
Physominthe Harley & J.F.B.Pastore
*Physopsis Turcz.
*Physostegia Benth.
*Piloblephis Raf.
Pitardia Batt. ex Pit., synonym of Nepeta
*Pityrodia R.Br.
*Platostoma P.Beauv.
*Plectranthus L.Hér.
*Pogogyne Benth.
*Pogostemon Desf.
*Poliomintha A.Gray
*Prasium L.
*Premna L.
*Prostanthera Labill.
*Prunella L.
*Pseuderemostachys Popov, synonym of Phlomoides
*Pseudocarpidium Millsp.
*Pseudocaryopteris (Briq.) P.D.Cantino
Pseudochamaesphacos Parsa
Pseudodictamnus Fabr.
*Pseudomarrubium Popov, synonym of Phlomoides
Puntia Hedge, syononym of Endostemon
*Pycnanthemum Michx.
*Pycnostachys Hook., synonym of Coleus
Quoya Gaudich.
Rabdosiella Codd, synonym of Coleus
*Renschia Vatke
*Rhabdocaulon (Benth.) Epling
*Rhaphiodon Schauer
*Rhododon Epling
-Rosmarinus L., synonym of Salvia
*Rostrinucula Kudô
*Rotheca Raf.
*Roylea Wall. ex Benth.
*Rubiteucris Kudô
+Rydingia Scheen & V.A.Albert
Sabaudia Buscal. & Muschl., synonym of Lavandula
*Saccocalyx Coss. & Durieu
Salazaria Torr., synonym of Scutellaria
*Salvia L.
*Satureja L.
*Schizonepeta (Benth.) Briq.
*Schnabelia Hand.-Mazz.
*Scutellaria L.
*Sideritis L.
*Siphocranion Kudô
Solenostemon Thonn., synonym of Coleus
*Spartothamnella Briq., synonym of Teucrium
*Sphenodesme Jack
*Stachydeoma Small
*Stachyopsis Popov & Vved., synonym of Eriophyton
*Stachys L.
*Stenogyne Benth.
*Sulaimania Hedge & Rech.f., synonym of Moluccella
*Suzukia Kudô
*Symphorema Roxb.
Symphostemon Hiern, synonym of Coleus
*Synandra Nutt.
*Syncolostemon E.Mey. ex Benth.
*Tectona L.f.
*Teijsmanniodendron Koord.
+Tetraclea A.Gray
*Tetradenia Benth.
*Teucridium Hook.f., synonym of Teucrium
*Teucrium L.
*Thorncroftia N.E.Br.
*Thuspeinanta T.Durand
*Thymbra L.
*Thymus L.
*Tinnea Kotschy ex Hook.f.
*Trichostema L.
*Tripora P.D.Cantino
*Tsoongia Merr., synonym of Vitex
*Vitex L.
*Viticipremna H.J.Lam, synonym of Vitex
+Volkameria L.
*Warnockia M.W.Turner
*Wenchengia C.Y.Wu & S.Chow
*Westringia Sm.
Wiedemannia Fisch. & C.A.Mey., synonym of Lamium
*Wrixonia F.Muell., synonym of Prostanthera
Xenopoma Willd., synonym of Clinopodium
*Zataria Boiss.
*Ziziphora L.

Recent changes

The circumscription of several genera has changed since 2004. Tsoongia, Paravitex, and Viticipremna have been sunk into synonymy with Vitex.^[15] Huxleya has been sunk into Volkameria.^[7] Kalaharia, Volkameria, Ovieda, and Tetraclea have been segregated from a formerly polyphyletic Clerodendrum.^[7] Rydingia has been separated from Leucas.^[16] The remaining Leucas is paraphyletic over four other genera.^[17]

Subfamilies and tribes

In 2004, the Lamiaceae were divided into seven subfamilies, plus 10 genera not placed in any of the subfamilies.^[5] The unplaced genera are: Tectona, Callicarpa, Hymenopyramis, Petraeovitex, Peronema, Garrettia, Cymaria, Acrymia, Holocheila, and Ombrocharis. The subfamilies are the Symphorematoideae, Viticoideae, Ajugoideae, Prostantheroideae, Nepetoideae, Scutellarioideae, and Lamioideae. The subfamily Viticoideae is probably not monophyletic.^[15] The Prostantheroideae and Nepetoideae are divided into tribes. These are shown in the phylogenetic tree below.

Phylogeny

Most of the genera of Lamiaceae have never been sampled for DNA for molecular phylogenetic studies.^[18] Most of those that have been are included in the following phylogenetic tree. The phylogeny depicted below is based on seven different sources.^[5]^[9]^[7]^[15]^[19]^[20]^[21]

Lamiaceae

Callicarpa

Tectona

Viticoideae (pro parte)

	Gmelina

	Premna

Viticoideae (pro parte)

Vitex

Symphorematoideae

	Congea

	Symphorema

Ajugoideae

Rotheca

Teucrium

Ajuga

	Oxera

	Faradaya

Kalaharia

Clerodendrum

Volkameria

	Ovieda

	Aegiphila

	Tetraclea

	Amasonia

Prostantheroideae

Chloantheae

Chloanthes

Westringieae

	Prostanthera

	Westringia

Nepetoideae

Ocimeae

Lavandula

Siphocranion

Isodon

Hanceola

Hyptis

	Orthosiphon

	Ocimum

	Plectranthus

	Coleus

Elsholtzieae

	Elsholtzia

	Perilla

Mentheae

	Lepechinia

	Salvia

	Rosmarinus

Prunella

	Nepeta

	Dracocephalum

	Agastache

	Origanum

	Thymus

Mentha

Satureja

	Clinopodium

	Bystropogon

	Pycnanthemum

	Monarda

	Dicerandra

	Conradina

Scutellarioideae

	Holmskioldia

	Scutellaria

Lamioideae

Pogostemon

Phlomis

Lamium

Stachys

Sideritis

Haplostachys

	Stenogyne

	Phyllostegia

	Leucas

	Leonotis

References

^ ^a ^b Stevens, P. F. (July 2012). "Lamiales (Lamiaceae Family)". Angiosperm Phylogeny Website. Archived from the original on 21 March 2015. Retrieved 25 March 2015.
^ Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x. hdl:10654/18083.
^ "Pronunciation of lamiaceae". Archived from the original on 2015-09-24. Retrieved 2014-11-25.
^ ^a ^b Heywood, Vernon H.; Brummitt, Richard K.; Seberg, Ole; Culham, Alastair (2007). Flowering Plant Families of the World. Ontario, Canada: Firefly Books. ISBN 978-1-55407-206-4.
^ ^a ^b ^c ^d ^e ^f ^g ^h Raymond M. Harley, Sandy Atkins, Andrey L. Budantsev, Philip D. Cantino, Barry J. Conn, Renée J. Grayer, Madeline M. Harley, Rogier P.J. de Kok, Tatyana V. Krestovskaja, Ramón Morales, Alan J. Paton, and P. Olof Ryding. 2004. "Labiatae" pages 167-275. In: Klaus Kubitzki (editor) and Joachim W. Kadereit (volume editor). The Families and Genera of Vascular Plants volume VII. Springer-Verlag: Berlin; Heidelberg, Germany. ISBN 978-3-540-40593-1
^ "World Checklist of Selected Plant Families". Archived from the original on 2017-07-09. Retrieved 2022-02-21.
^ ^a ^b ^c ^d Yuan, Yao-Wu; Mabberley, David J.; Steane, Dorothy A.; Olmstead, Richard G. (2010). "Further disintegration and redefinition of Clerodendrum (Lamiaceae): Implications for the understanding of the evolution of an intriguing breeding strategy". Taxon. 59 (1): 125–133. doi:10.1002/tax.591013.
^ Cantino, P.D., Harley, R.M. & Wagstaff, S.J. 1992. Genera of Labiatae: status and classification. Pp. 511-522. In: Raymond M. Harley and Tom Reynolds (editors). Advances in Labiate Science. Richmond, Royal Botanic Gardens, Kew.
^ ^a ^b Wagstaff, Steven J.; Hickerson, Laura; Spangler, Russ; Reeves, Patrick A.; Olmstead, Richard G. (1998). "Phylogeny in Labiatae s.l., inferred from cpDNA sequences". Plant Systematics and Evolution. 209 (3–4): 265–274. Bibcode:1998PSyEv.209..265W. doi:10.1007/bf00985232. S2CID 601658.
^ Parnell, J. and Curtis, T. 2012. Webb's An Irish Flora. Cork University Press ISBN 978-185918-4783
^ "List of genera in Lamiaceae". In: "Lamiaceae". In: "List of families". In: "Families and genera in GRIN. (see External links below)
^ List of Genera in Lamiaceae. At: Vascular Plant Families and Genera. At: World Checklist of Selected Plant Families. At: Electronic Plant Information Center. At: Website of Royal Botanic Gardens, Kew. (see External Links below).
^ "APG list of Lamiaceae genera". Archived from the original on 2022-01-31. Retrieved 2022-02-21.
^ Lamiaceae Martinov. Plants of the World Online. Retrieved 14 April 2024.
^ ^a ^b ^c Bramley, Gemma L.C.; Forest, Félix; Rogier (2009). "Troublesome tropical mints: re-examining generic limits of Vitex and relations (Lamiaceae) in South East Asia". Taxon. 58 (2): 500–510. doi:10.1002/tax.582014.
^ Scheen, Anne-Cathrine; Albert, Victor A. (2007). "Nomenclatural and taxonomic changes within the Leucas clade (Lamioideae; Lamiaceae)". Systematics and Geography of Plants. 77 (2): 229–238. JSTOR 20649740.
^ Scheen, Anne-Cathrine; Albert, Victor A. (2009). "Molecular Phylogenetics of the Leucas Group (Lamioideae; Lamiaceae)". Systematic Botany. 34 (1): 173–181. doi:10.1600/036364409787602366. S2CID 85894904.
^ Zhao, Fei; Chen, Ya-Ping; Salmaki, Yasaman; Drew, Bryan T.; Wilson, Trevor C.; Scheen, Anne-Cathrine; Celep, Ferhat; Bräuchler, Christian; Bendiksby, Mika; Wang, Qiang; Min, Dao-Zhang; Peng, Hua; Olmstead, Richard G.; Li, Bo; Xiang, Chun-Lei (8 January 2021). "An updated tribal classification of Lamiaceae based on plastome phylogenomics". BMC Biology. 19 (1): 2. doi:10.1186/s12915-020-00931-z. PMC 7796571. PMID 33419433.
^ Zhong, Jin-Shun; Li, Jie; Li, Lang; Conran, John G.; Hsi-wen, Li (2010). "Phylogeny of Isodon (Schrad. ex Benth.) Spach (Lamiaceae) and Related Genera Inferred from Nuclear Ribosomal ITS, trnL-trnF Region, and rps16 Intron Sequences and Morphology". Systematic Botany. 35 (1): 207–219. doi:10.1600/036364410790862614. S2CID 54808462.
^ Walker, Jay B.; Sytsma, Kenneth J. (2007). "Staminal Evolution in the Genus Salvia (Lamiaceae): Molecular Phylogenetic Evidence for Multiple Origins of the Staminal Lever". Annals of Botany. 100 (2): 375–391. doi:10.1093/aob/mcl176. PMC 2735309. PMID 16926227.
^ Ryding, P. Olof (2010). "Pericarp structure and phylogeny of tribe Mentheae (Lamiaceae)". Plant Systematics and Evolution. 285 (3–4): 165–175. Bibcode:2010PSyEv.285..165R. doi:10.1007/s00606-010-0270-9. S2CID 24076224.