Lamiaceae Martinov

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition

Characters of similar families: Rubiaceae: stipules present, leaf margins entire, ovary inferior. Boraginaceae: leaves alternate, flowers usually actinomorphic. Solanaceae: leaves alternate, fruit many-seeded. Scrophulariaceae, Plantaginaceae, Orobanchaceae: fruits usually many-seeded capsules.

Morphology General Habit

Herbs, shrubs, trees or lianas; herbaceous stems usually quadrangular; often the crushed foliage aromatic or foetid; stipules absent

Morphology Leaves

Leaves simple, more rarely compound, opposite (decussate); lamina margins often toothed or lobed; hairs often present, simple to complex

Morphology Reproductive morphology Inflorescences

Inflorescences usually 1–many flowered cymes of various forms, particularly verticillasters or thyrses; terminal or axillary, usually with bracts

Morphology Reproductive morphology Flowers

Flowers usually zygomorphic, more rarely actinomorphic; calyx 2–5-lobed, rarely unlobed or with more than 5 lobes, campanulate, or funnel-shaped, or tubular, persistent, often accrescent; corolla more or less 5-lobed, usually bilabiate unless actinomorphic; stamens 4, or 2 by abortion and staminodes usually present; ovary superior, unlobed to deeply 4-lobed, 2-carpellate, often appearing 4-locular by intrusions of the ovary wall constituting ‘false septa’; 2 ovules per carpel (1 in each ‘locule’), lateral attachment, placentation axile; style gynobasic or terminal, not persistent (except in Australia)

Morphology Reproductive morphology Fruits

Fruit more or less a schizocarp splitting into typically 4 nutlets, or a drupe (1)–4-seeded, both enclosed in the persistent calyx

Morphology Reproductive morphology Seeds

Seeds endospermic to non-endospermic.

Distribution

A family of 236 genera and ca. 7,300 species; almost cosmopolitan.

Note

Leaves opposite. Inflorescence cymose or thyrsoid; calyx persistent, often accrescent; corolla tubular, usually zygomorphic; stamens (2)4, exserted. Fruit schizocarp with (1)–4 nutlets, sometimes a drupe.

Description Author

Gemma Bramley

Timothy Utteridge & Gemma Bramley (2020). The Kew Tropical Plant Families Identification Handbook, Second Edition. Kew Publishing Royal Botanic Gardens, Kew

Note

Leaves opposite. Inflorescence cymose or thyrsoid; calyx persistent, often accrescent; corolla tubular, usually zygomorphic; stamens (2)4, exserted. Fruit a schizocarp of 4 nutlets or drupe with (1)–4(10) seeds.

Recognition

Characters of similar families: Rubiaceae: stipules, leaf margins entire, inferior ovary. Rutaceae: calyx and corolla free. Solanaceae: leaves alternate, fruit many-seeded. Linderniaceae, Orobanchanceae or Gesneriaceae: fruits usually many-seeded capsules. Apocynaceae: white sap, twisted buds, leaf margins entire.

Morphology General Habit

Herbs, shrubs, trees or lianas; often the crushed foliage aromatic or foetid

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves opposite (decussate on the usually square stem), or whorled; simple or compound; lamina margins entire, crenate or serrate; hairs often present, simple to complex

Morphology Reproductive morphology Inflorescences

Inflorescences solitary or in cymes, often arranged in verticils, heads or thyrses; terminal or axillary

Morphology Reproductive morphology Flowers

Flowers usually zygomorphic, more rarely actinomorphic; calyx 2-, 3-, 4- or 5-lobed, rarely unlobed or with more than 5 lobes, campanulate, or funnel-shaped, or tubular, persistent, often accrescent; corolla more or less 5-lobed, usually bilabiate unless actinomorphic; stamens 2 or 4 (rarely 5, 6 or 7); ovary 2-locular but appearing 4-locular by intrusions of the ovary wall constituting ‘false septa’; style gynobasic or terminal, not persistent (except subfamily Prostantheroideae)

Morphology Reproductive morphology Fruits

Fruit more or less a schizocarp, typically of four nutlets, or a drupe (1–)4(–14)-seeded, both enclosed in the persistent calyx

Morphology Reproductive morphology Seeds

Seeds endospermic to non-endospermic.

Distribution

An almost cosmopolitan family with 236 genera and c. 7,200 species. Large genera in the tropics include the woody Vitex and Clerodendrum and the herbaceous Salvia, Hyptis (both mostly New World) and Plectranthus (Old World).

Ecology

Found in drylands and wet tropics, lowland and montane habitats.

Description Author

Gemma Bramley

Lamiaceae (Labiatae), A.J. Paton, G. Bramley, O. Ryding, R.M. Polhill, Y.B. Harvey, M. Iwarsson, F. Willis, P.B. Phillipson, K. Balkwill, C.W. Lukhoba, D.F. Otieno, & R.M. Harley. Flora of Tropical East Africa. 2009

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, actinomorphic to more often slightly to strongly zygomorphic, often 2-lipped, rarely 1-lipped, rarely resupinate; tube short to elongate, rarely spurred, sometimes with a ring of hairs or appendaged within; lobes (2–)4–5(–16), equal or unequal, one or other lip often concave to galeate

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens epipetalous, attached within corolla tube, usually 4 or 2 by abortion and then staminodes often present, or stamens 5–6, when 4 often didynamous, usually free; filaments usually exserted from corolla tube, sometimes included within lip of corolla, rarely held within tube; anthers dithecous, synthecous or monothecous by abortion, opening by longitudinal slits or rarely by pores

Morphology Reproductive morphology Flowers Disc

Disk at base of ovary often present, nectariferous

Morphology Reproductive morphology Flowers Gynoecium

Gynoecium 2-carpellate, often 4-locular by intrusion of carpel margin forming a “false septum”, or rarely imperfectly 2-locular and free towards apex; ovary entire or lobed, with terminal style, or more often deeply 4-lobed, the locules often separated and with style gynobasic; ovules usually 4, anatropous to hemianatropous, usually basal or sub-basal, erect, rarely orthotropous, apical, pendulous, borne submarginally on the carpel wall (placenta); style usually with 2 equal or unequal stigma-lobes, rarely entire with 1 stigma-lobe vestigial, or stigma capitate or very rarely 4-lobed

Morphology Reproductive morphology Fruits

Fruit drupaceous or dry and indehiscent, 4-seeded or fewer by abortion, or frequently splitting into four 1-seeded mericarps, sometimes fewer by abortion; mericarps (nutlets) often with sculptured, tuberculate, hairy or rarely winged pericarp, mucilage cells often present

Morphology Reproductive morphology Seeds

Seeds albuminous or exalbuminous; embryo straight or bent.

Note

The traditional division of Verbenaceae and Lamiaceae is far from satisfactory, e.g. Bentham & Hooker in G.P. 2: 1131–1223 (1876); Baker & Stapf in F.T.A. 5: 273–502 (1900). The delimitation of these families was based on whether the taxa were mostly woody with a terminal or subterminal style (Verbenaceae) or mostly herbaceous with a gynobasic style (Lamiaceae). This traditional classification is difficult to implement, there being many intermediates, and also it does not represent phylogenetically natural taxa. A full discussion of the problems with this traditional classification and of the new circumscription of Verbenaceae and Lamiaceae is provided by Harley in Kubitzki, Fam. Gen. Vasc. Pl. 7: 188–190 (2004). The Verbenaceae are now restricted to subfamily Verbenoideae of traditional classifications (e.g. Briquet in E.P. Pf. IV, 3a: 132–182 (1895)) which has an indeterminate racemose inflorescence and a salverform corolla with stamens included; whereas in the Lamiaceae the inflorescence is cymose with determinate, usually opposite cymes and the corollas are tubular and usually bilabiate with the stamens usually exserted from the tube, but can be held within the lobes and rarely within the tube. The cymes in Lamiaceae are arranged usually in opposite pairs along an indeterminate axis, forming a thyrse. In some Lamiaceae the cymes are reduced to single flowers though bracteoles are often present below the flower in these cases, indicating the cymose, rather than racemose, nature of the inflorescence. These gross morphological differences are supported by anatomical and pollen characters: the Verbenaceae have their ovules attached marginally on the carpel margin, and have thickened pollen exine near the apertures; the Lamiaceae have the ovules attached submarginally and have an unthickened exine. Good general accounts of the Verbenaceae can be found in Sanders in Harvard Papers in Botany 5: 303–358 (2001) and Atkins in Kubitzki, Fam. Gen. Vasc. Pl. 7: 449–468 (2004). Following this delimitation the Verbenaceae includes 34 genera and around 1200 species, and is most abundant in temperate South America. Within the Flora of Tropical East Africa area the family is represented by 8 genera and 39 species. The genera are: Chascanum, Duranta, Lantana, Lippia, Phyla, Priva, Stachytarpheta and Verbena. Thus only genera 1–8 and the cultivated Aloysia, Citharexylum and Petrea listed by Verdcourt, Flora of Tropical East Africa, Verbenaceae (1992) remain in Verbenaceae. The classification of the Lamiaceae used here follows the account of Harley et al. in Kubitzki, Fam. Gen. Vasc. Pl. 7: 167–275 (2004). There are seven subfamilies. The genera of subfamily Viticoideae found within the Flora area have already been covered by the FTEA treatment of Verbenaceae (Verdcourt, 1992). Premna and Vitex are the only natively occurring genera of the Viticoideae, with teak ( Tectona) and Gmelina being the best known introduced members of this group. Subfamily Ajugoideae within the Flora area comprises Teucrium (genus 1) and Ajuga (2) covered in this volume, plus Clerodendrum (including Rotheca) and Karomia accounted for in the Flora treatment of Verbenaceae (Verdcourt, 1992). Subfamily Scutellarioideae includes Tinnea (3) and Scutellaria (4) in this volume plus the cultivated Holmskioldia included within the Verbenaceae volume. Subfamily Lamioideae includes genera 5–9, and subfamily Nepetoideae comprises genera 9–30. Subfamily Nepetoideae is by far the largest group of Lamiaceae and can be divided into tribe Mentheae (genera 10–14) and tribe Ocimeae (genera 15–30). Tribe Ocimeae is the most speciose group in the Flora area and an account of the relationships within this group can be found in Paton et al., Molecular Phylogenetics and Evolution 31: 277–299 (2004). Of the remaining subfamilies, subfamily Prostantheroideae is Australian and absent from the Flora area, whereas the Asiatic Symphorematoideae is represented only by the cultivated Congea, included in the FTEA Verbenaceae account. In the FTEA area the family is represented by 416 species in 35 genera, 327 species in 30 genera being covered in this volume and 89 in the Flora treatment of Verbenaceae. The Lamiaceae is one of the most widely used and cultivated plant families, with 33 genera cultivated in the FTEA area. Uses include timber ( Tectona and Gmelina), medicine, flavouring and as ornamanentals. The family is best known for its aromatic herbs, many of which belong in subfamily Nepetoideae. A good review of uses and underlying phytochemistry can be found in Harley et al. (2004) along with references to more detailed sources. The account of cultivated Lamiaceae below is unlikely to be comprehensive as new species and cultivars will be introduced as the horticultural trade in southern and tropical Africa develops further. A comprehensive account of cultivated Lamiaceae can be found in Cullen, J. et al. (eds) (2000) European Garden Flora 6: 149–230.

Morphology General Habit

Annual or perennial herbs, shrubs or climbers, rarely trees, aromatic or not

Morphology Stem

Stems often square in cross-section

Morphology Leaves

Leaves simple or sometimes compound and then digitate or pinnate, opposite, often decussate, sometimes whorled, very rarely alternate, rarely forming a basal rosette, exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence composed of cymes and often arranged in a terminal, lax or congested indeterminate thyrse which may be paniculate, raceme-like with cymes 1-flowered, or spike-like, or rarely congested into a head, usually with bracts and sometimes with bracteoles

Morphology Reproductive morphology Flowers

Flowers actinomorphic to zygomorphic, hypogynous, usually bisexual

Morphology Reproductive morphology Flowers Calyx

Calyx gamosepalous, sometimes 2-lipped, often accrescent; lobes 2–many, often 5, equal or unequal, rarely obsolete, some lobes often fused, or lips entire

Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://creativecommons.org/licenses/by/3.0

Morphology General Habit

Trees, shrubs, subshrubs or perennial or annual herbs, [rarely climbers ], aromatic or not.  Stems often quadrangular, erect to prostrate, sometimes forming stolons or large or slender rhizomes

Morphology General Indumentum

Indumentum usually present, of glandular and non- glandular trichomes, often hair-like, rarely scale-like, usually multicellular-uniseriate, simple, branched, dendroid or stellate, sometimes gland -tipped, large-headed subsessile glands rarely absent

Morphology Leaves

Leaves opposite, often decussate, sometimes whorled, very rarely alternate simple, entire, toothed or lobed, sometimes compound and then digitate, petiolate or sessile, rarely forming a basal rosette, exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence often bracteate, bracts persistent or deciduous, rarely spirally arranged, composed of cymes, bracteolate or not, and often arranged in a terminal, lax or congested indeterminate thyrse which may be paniculate, raceme -like with cymes often 1-flowered, or spike -like, or rarely congested into a capitulum, with or without a distinct involucre of bracteoles, sometimes conspicuous

Morphology Reproductive morphology Flowers

Flowers hypogynous, usually bisexual, or less often unisexual due to gynodioecy or gynomonoecy, very rarely due to dioecy

Morphology Reproductive morphology Flowers Perianth

Perianth biseriate, sepals 4-5[(--7)], connate, actinomorphic to zygomorphic, sometimes 2-lipped, lobes 2--many, often 5, equal or unequal, rarely obsolete, some lobes often fused, or lips entire, calyx -tube (5--)10--15-nerved, straight or curved, throat hairy or glabrous, calyx often accrescent, rarely inflated or fleshy in fruit

Morphology Reproductive morphology Flowers Corolla

Petals (4--)5(--16) connate, actinomorphic to more often slightly to strongly zygomorphic, often 2-lipped, rarely 1-lipped, lobes (2--)4--5[(--7)], equal or unequal, porrect to patent, one or other lip often concave to galeate, corolla -tube short to elongate, [rarely spurred], often with annulus of hairs or appendaged within, rarely corolla resupinate

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens epipetalous, attached within corolla -tube, usually 4 or 2 by abortion and then staminodes often present, or stamens 5[-6], when 4 often didynamous (rarely a fifth, posterior vestigial staminode present), free or rarely monodelphic, filaments short or often elongate, usually exserted from corolla -tube and sometimes long- exserted from corolla; parallel, divergent or ascending and sometimes included within or lying under the posterior corolla -lip, or declinate and then sometimes included within the anterior corolla -lip, anthers usually dithecous, tetrasporangiate or monothecous by abortion, thecae parallel or divergent, occasionally widely separated by an elongate connective, or apically confluent or synthecous, opening by longitudinal slits or rarely by pores

Disc

Disc at base of ovary often present, usually fleshy, entire or irregularly or often 4- lobed, anterior lobe sometimes longer than others, nectariferous

Morphology Reproductive morphology Flowers Gynoecium

"Gynoecium 2-carpellate, often 4-locular by intrusion of carpel wall forming ""false septum"", [or rarely imperfectly 2-locular and free towards apex ], ovary usually 4-ovuled, [2-locular ovaries generally with loculi 2-ovuled and 4-locular ovaries with 1 ovule per loculus], ovary entire or lobed, with terminal style, or more often deeply 4- lobed, the loculi often separated and with style gynobasic; style not  persistent or rarely persistent in young fruit (Prostantheroideae), usually with 2 equal or unequal stigma -lobes, rarely entire with 1 stigma -lobe vestigial, or stigma capitate or very rarely 4- lobed"

Morphology Reproductive morphology Flowers Gynoecium Ovary Ovules

Ovules anatropous to hemianatropous, usually basal or sub - basal, erect, rarely orthotropous, apical, pendulous, borne laterally or submarginally on the placenta, unitegmic, tenuinucellate

Morphology Reproductive morphology Fruits

Fruit drupaceous, often with pyrenes, or dry, indehiscent, or frequently, four 1-seeded mericarps, sometimes fewer by abortion

Morphology Reproductive morphology Flowers Gynoecium Carpels

Mericarps (nutlets) often with sculptured, tuberculate, hairy or rarely winged pericarp, mucilage cells often present

Morphology Reproductive morphology Seeds

Seeds albuminous or exalbuminous, epigeal-Embryo straight or bent, investing or spatulate.

Diagnostic

Salvia: Recognizable by its unusual stamen structure.  Salvia only expresses two stamens, and the thecae on each stamen are separated by an elongate connective.  This staminal structure is associated with a pollination syndrome in which the pollinator pushes against the posterior (usually sterile) anther theca while accessing a nectar reward at the base of the corolla tube. This causes the anterior (fertile) anther theca to deposit pollen on the pollinator via a lever-like mechanism. At least 500 species in Central and South America. The infra-generic classification proposed, especially by Briquet (1897), largely based on staminal structure, is out-dated, though the Neotropical subgenera Calosphace Benth. and Audibertia (Benth.) Epling appear to be monophyletic. Hyptis: Flowers arranged variously, often in pedunculate or sessile, congested cymes or involucrate capitula in terminal panicles, or in spikes or axillary. Flowers sometimes in lax cymes, rarely fasciculate on long pedicels in the axils of the leaves. About 280 species, mostly tropical and subtropical savannas, sometimes in humid areas, almost entirely New World, from southern U.S.A. to Caribbean and S to Argentina and Peru, a few species extending into Old World. Grouped into seven subfamilies, five of which are native in the Neotropics: Symphorematoideae: Congea Roxb.(planted): Climbers. Inflorescence capitate, subtended by enlarged, showy bracteoles. Viticoideae (transferred from Verbenaceae sensu Briq.): Tropical shrubs and trees. Characteristic fruit - a drupe, likened to an egg in egg-cup. Terminal style. Vitex usually has digitate leaves. Ajugoideae (some genera transferred from Verbenaceae sensu Briq.): Stamens well exserted. Actinomorphic to 1-lipped corolla. Fruit a drupe with a terminal style but often lobes evident. Scutellarioideae (In Neotropics): Calyx usually 2-lipped with lips entire e.g. Scutellaria Riv. ex L. Posterior lip of calyx usually folded to form a scutellum. Lamioideae (wide range of genera e.g. Stachys L. Several genera widely introduced as weeds (Lamium L., Marrubium L.,Leonitis Spach,Leonurus L.): Gynobasic style. Nepetoideae: Often aromatic. Pollen hexacolpate (other subfamilies usually tricolpate e.g. Salvia L., Hyptis Jacq., Minthostachys (Benth.) Spach, Clinopodium L.). Many genera introduced and cultivated. Prunella L. naturalized in montane areas. Key differences from similar families: Separate from Verbenaceae because of cymose inflorescence, usually lipped (rather than salverform) corolla, stamens well-exserted. Differs from Rubiaceae as fused interpetiolar stipules absent, usually zygomorphic flowers, superior (rather than inferior) ovary. Differs from Solanaceae as usually opposite leaves, fruits with 4 nutlets or 1-5 pyrenes (not copious seed). Separated from Apocynaceae by absence of white sap, petals not twisted in bud, no colleters on petiole base and adjoining stem, seed usually without tuft of hairs; androecium and gynoecium never fused. Separated from Boraginaceae as flowers not regular, cymes usually not scorpioid, often not hispid. Separate from Rutaceae (compound leaves Vitex can be confused with Rutaceae) as flowers not regular corolla and calyx lobes fused (not free), leaves usually opposite (not alternate). Leaves opposite. Usually quadrangular stem when young. Stamens exserted. Corolla often zygomorphic- Style soon deciduous. Sessile glands. Distinguishing characters (always present): Stipules absent. Cymose inflorescence. Tubular corolla and persistent synsepalous calyx. Superiorovary.

Distribution

See Neotropical genera The South American Labiate flora, which contains two very large genera, Salvia L. and Hyptis Jacq., but rather few genera in total, falls naturally into three regions: A. Andean; B. Guianan and Brazilian Shields; C Temperate South America. A. The Andean Labiatae are primarily derived from north temperate genera, and the various groups have many links to Central America, including Mexico. The largest group is Salvia subg. Calosphace Raf. with its Andean centre of diversity linked to a much larger one in Mexico. Scutellaria L., Stachys L. and Clinopodium L. also appear to have been derived from a southern migration from Central America, but have radiated to produce endemic groups, in some cases associated with adaptations to humming-bird pollination. Other Andean Nepetoid genera include the monotypicObtegomeria Doroszenko & P.D.Cantino from Colombia, related to Clinopodium, Minthostachys (Benth.) Spach, which appears most closely related to the Macaronesian Bystropogon L'Hér., and Lepechinia Willd., which extends to Mexico with one Hawaiian extension. The small Neotropical genus Catoferia (Benth.) Benth. (Nepetoideae, Ocimeae) also links Central and S America, but its closest relatives would seem to be eastern Asiatic members of the Old World subtribe Ociminae, perhaps the genus Orthosiphon Benth. & M.Ashby. The only other member of the subtribe native to the Americas is Ocimum L., which extends from Mexico to South America. One species extends into lowland areas of the Andes. B. Guianan and Brazilian Shields. This region, including Amazonian and eastern South America, mostly overlying much older geological formations, has a very different flora, dominated by the Nepetoid Ocimeae, subtribe Hyptidinae. This is composed of the large genus Hyptis with about 300 species, and a number of smaller endemic, satellite genera, such as Peltodon Pohl, Rhaphiodon Schauer, Hypenia Mart. ex Benth., Hyptidendron Harley and Eriope Humb. & Bonpl. ex Benth. The primarily Old World genus Ocimum (Nepetoideae: Ocimeae) is represented by several species endemic to the area. It seems probable that Ocimum originally had an Old World tropical origin, perhaps reaching the New World via W Africa in the early Tertiary. A few species of Salvia (Mentheae) reach the mountains of eastern Brazil and extend south into temperate S America. Other endemic genera to be found in this region include Cornutia (Viticoideae) L., Amasonia L.f. and Monochilus (Ajugoideae) Fisch. & C.A.Mey., and more widespread genera such as Vitex L.,  Clerodendrum (Viticoideae) L. and Aegiphila (Ajugoideae) Jacq. The latter, with over 100 species, extends throughout tropical America. C. Temperate South America. A number of endemic genera occur in this region, belonging to Nepetoideae, tribe Mentheae: Kurzamra Kuntze, Rhabdocaulon (Benth.) Epling, Glechon Spreng. and Hoehnea Epling, while Cunila D.Royen ex L. and Hesperozygis Epling are trans-equatorial, being also represented in Mexico, although this needs confirmation from molecular studies. D. Northern America and Mexico. This region has quite a large number of endemic genera, both in the west, especially California, as well as in the East. Most of these belong to Tribe Mentheae (Nepetoideae) and include: Monarda L., Monardella Benth., Pycnanthemum Michx., Rhododon Epling, Stachydeoma Small, Pogogyne Benth., Acanthomintha A.Gray, Neoeplingia Ramamoorthy, P.Hiriart Valencia & F.González Medrano and Blephilia Raf.  Endemic Lamioideae include: Warnockia M.W.Turner, Physostegia Benth., Macbridea Raf., Brazoria Engelm. & Gray and Synandra Nutt. Trichostema (Ajugoideae) Gronov. ex L. is a near endemic, extending into the Caribbean. More widespread, species-rich genera represented in the area, include Scutellaria (Scutellarioideae) L., Stachys (Lamioideae), Salvia, Lycopus L. Clinopodium and Hedeoma (Nepetoideae) Pers., the last a New World genus extending into Andean and temperate South America. Of particular interest is Lycopus (Mentheae), with a circum-boreal distribution, but well represented in North America, while Mentha L. and Dracocephalum L., both primarily Old World genera, each have one species native in North America. One or two members of the species-rich genus Clerodendrum (Ajugoideae) L., primarily of the Old World tropics, occur in southern areas. The primarily tropical Viticoideae is represented by Vitex L., which in the New World is confined to the Caribbean and Central America from Mexico southwards, but there are also two genera endemic to the Caribbean: Petitia Jacq. and Pseudocarpidium Millsp. Mention should also be made here of Callicarpa (Incertae sedis) L. This is a mainly Asiatic genus, with a few New World species, especially C. americana L., in the S United States and Mexico, and others in the Caribbean.

Note

General notes: Number of genera: There are 65 genera of Lamiaceae, of which 48 are native and 17 cultivated/introduced: Acanthomintha (A.Gray) A.Gray, Syn. Fl. N. Amer. 2(1): 365 (1878).California, NW. Mexico. 1 Species Aegiphila Jacq., Select. Stirp. Amer. Hist.: t. 16 (1763).Mexico to Tropical America 156 Species. Agastache Clayton ex Gronov., Fl. Virgin., ed. 2: 88 (1762).C. & E. Asia, N. America. 12 Species Amasonia L.f., Suppl. Pl.: 48 (1782).Trop. America. 5 Species Anisomeles  R.Br., Prodr.: 503 (1810). 1 Species introduced in Neotropics, native to W. Indian Ocean, Trop. & Subtrop. Asia to N. Australia Asterohyptis Epling, Bull. Torrey Bot. Club 60: 17 (1933 publ. 1932).Mexico to C. America, Caribbean. 3 Species Ballota L., Sp. Pl.: 582 (1753).  1 Species introduced in Neotropics, native to Macaronesia, Europe, Medit. to W. Asia, Mauritania, Chad, S. Africa Callicarpa L., Sp. Pl.: 111 (1753).America, W. Indian Ocean to W. Pacific. 31 Species Catoferia (Benth.) Benth. in G.Bentham & J.D.Hooker, Gen. Pl. 2: 1173 (1876).Mexico to Colombia. 4 Species Chaunostoma J.D.Sm., Bot. Gaz. 20: 9 (1895).SE. Mexico to C. America. 1 Species Clerodendrum L., Sp. Pl.: 637 (1753).Trop. & Subtrop. 34 Species Clinopodium L., Sp. Pl.: 587 (1753).Temp. & Subtrop. 57 Species Congea Roxb., Pl. Coromandel 3: 90 (1820) 1 Species introduced in Neotropics, native to India to SC. China and W. Malesia Cornutia Plum. ex L., Sp. Pl.: 628 (1753).Mexico to Trop. America. 9 Species Cunila Royen ex L., Syst. Nat. ed. 10, 2: 1359 (1759), nom. cons.C. & E. U.S.A. to C. America, Brazil to Argentina. 20 Species Eriope Humb. & Bonpl. ex Benth., Labiat. Gen. Spec.: 142 (1833). S. Trop. America. 30 Species Eriothymus (Benth.) Rchb., Handb. Nat. Pfl.-Syst.: 189 (1837).SE. Brazil. 1 Species Glechon Spreng., Syst. Veg. 4(2): 227 (1827).Brazil to Argentina. 7 Species Gmelina. Sp. Pl.: 626 (1753).1 Species introduced in Neotropics, native to  Mascarenes, Trop. & Subtrop. Asia to W. Pacific Hedeoma Pers., Syn. Pl. 2: 131 (1806).America. 34 Species Hesperozygis Epling, Repert. Spec. Nov. Regni Veg. Beih. 85: 132 (1936).Mexico, Brazil. 7 Species Hoehnea Epling, Repert. Spec. Nov. Regni Veg. Beih. 115: 8 (1939).Brazil to Argentina. 4 Species Holmskioldia Retz., Observ. Bot. 6: 31 (1791).  1 Species introduced to Neotropics, native to  Indian Subcontinent to Myanmar Hypenia (Mart. ex Benth.) Harley, Bot. J. Linn. Soc. 98: 91 (1988).N. South America to Brazil. 24 Species Hyptidendron Harley, Bot. J. Linn. Soc. 98: 90 (1988).S. Trop. America. 16 Species Hyptis Jacq., Collectanea 1: 101 (1787).Trop. & Subtrop. America, W. Trop. Africa. 289 Species Lamium L., Sp. Pl.: 579 (1753). 2 Species introduced to Neotropics, native to Temp. Eurasia, Macaronesia to Ethiopia Leonotis (Pers.) R.Br., Prodr.: 504 (1810).Trop. & S. Africa, Madagascar. 1 Species introduced to Neotropics Leonurus L., Sp. Pl.: 584 (1753).Temp. Eurasia. 2 Species introduced to Neotropics Lepechinia Willd., Hort. Berol. 1: 20 (1804).SW. U.S.A. to S. America. 37 Species Leucas R.Br.: 504 (1810). 1 Species introduced to Neotropics, native to Africa to NE. Australia Marrubium L., Sp. Pl.: 583 (1753).Macaronesia to Temp. Eurasia. 1 Species introduced to Neotropics Marsypianthes Mart. ex Benth., Labiat. Gen. Spec.: 64 (1833).Mexico to Trop. America. 6 Species Melissa L., Sp. Pl.: 592 (1753).S. Europe to Malesia. 1 Species introduced to Neotropics  Mentha L., Sp. Pl.: 576 (1753).Cosmopolitan. 5 Species Micromeria Benth., Edwards's Bot. Reg. 15: t. 1282 (1829).Temp. & Subtrop. 3 Species Mintostachys (Benth.) Spach, Hist. Nat. Vég. 9: 164 (1840).S. Trop. America. 11 Species Moluccella L., Sp. Pl.: 587 (1753).Medit. to C. Asia. 1 Species (cultivated/introduced) Monochilus Fisch. & C.A.Mey., Index Seminum (LE) 1: 34 (1835).Brazil. 84. 2 Species Neoeplingia Ramamoorthy, Hiriart & Medrano, Bol. Soc. Bot. México 43: 61 (1982).Mexico. 79. 1 Species Nepeta L., Sp. Pl.: 570 (1753).Temp. Eurasia Macaronesia to E. Trop. Africa. 1 Species introduced to Neotropics Obtegomeria Doroszenko & P.D.Cantino, Novon 8: 2 (1998).W. South America. 83. 1 Species Ocimum L., Sp. Pl.: 597 (1753).Trop. & Subtrop. 10 Species Origanum L. 1 Species introduced to Neotropics, native to Macaronesia, Europe, Medit. to C. China Peltodon Pohl, Pl. Bras. Icon. Descr. 1: 66 (1827).S. Trop. America. 5 Species Petitia Jacq., Enum. Syst. Pl.: 1 (1760).Caribbean. 81. 3 Species Physostegia Benth. Edwards's Bot. Reg. 15: t. 1289 (1829) 2 Species, N. America to Mexico Plectranthus L'Hér., Stirp. Nov.: 84, verso (1788).Trop. & Subtrop. 4 Species Poliomintha A.Gray, Proc. Amer. Acad. Arts 8: 295 (1873).S. U.S.A. to Mexico. 8 Species Prunella L., Sp. Pl.: 600 (1753).Temp. & Subtrop. Northern Hemisphere. 1 Species probably naturalised in Neotropics Pseudocarpidium Millsp., Publ. Field Columbian Mus., Bot. Ser. 2: 181 (1906).Caribbean. 81. 9 Species Rhaphiodon Schauer, Flora 27: 345 (1844).Brazil. 1 Species Rosmarinus L., Sp. Pl.: 23 (1753).  1 Species introduced to Neotropics, native to Mediterranean Salvia L., Sp. Pl.: 23 (1753).Cosmopolitan. 605 Species Scutellaria L., Sp. Pl.: 598 (1753).Cosmopolitan. 96 Species Stachys L., Sp. Pl.: 580 (1753).Cosmopolitan. 72 Species Tectona L.f., Suppl. Pl.: 151 (1781), nom. cons.  1 Species introduced to Neotropics, native to Tropical Asia Teucrium L., Sp. Pl.: 562 (1753).Cosmopolitan. 10 Species Tinnea Kotschy & Peyr., Pl. Tinn.: 25 (1867). 1 Species introduced to Neotropics, native to Trop. & S. Africa Trichostema L., Sp. Pl.: 598 (1753).N. America. 8 Species Vitex L., Sp. Pl.: 638 (1753).Trop. & Subtrop. 61 Species Warnockia M.W.Turner, Pl. Syst. Evol. 203: 78 (1996).C. U.S.A. to NE. Mexico. 1 Species The traditional division of Verbenaceae and Lamiaceae is far from satisfactory, e.g., Bentham & Hooker in Gen. Pl. 2: 1131-1223 (1876); Baker & Stapf in F.T.A. 5: 273-502 (1900). The delimitation of these families was based on whether the taxa were mostly woody with a terminal or subterminal style (Verbenaceae) or mostly herbaceous with a gynobasic style (Lamiaceae).  This traditional classification is difficult to implement, there being many intermediates, and also it does not represent phylogenetically natural taxa.  A full discussion of the problems with this traditional classification and of the new circumscription of Verbenaceae and Lamiaceae is provided by Harley in Kubitzki, Fam. Gen. Vasc. Pl. 7: 188-190 (2004). The Verbenaceae is now restricted to subfamily Verbenoideae of traditional classifications (e.g. Briquet in Nat. Pflanzenfam. 4, 3a: 132-182 (1895)) which has an indeterminate racemose inflorescence and a salverform corolla with stamens included; whereas in the Lamiaceae the inflorescence is cymose with determinate, usually opposite cymes and the corollas are tubular and usually bilabiate with the stamens usually exserted from the tube, but can be held within the lobes and rarely within the tube.  The cymes in Lamiaceae are arranged usually in opposite pairs along an indeterminate axis, forming a thyrse. In some Lamiaceae the cymes are reduced to single flowers though bracteoles are often present below the flower in these cases, indicating the cymose, rather than racemose, nature of the inflorescence.  These gross morphological differences are supported by anatomical and pollen characters: the Verbenaceae have their ovules attached marginally on the carpel margin, and have thickened pollen exine near the apertures; the Lamiaceae have the ovules attached submarginally and have an unthickened exine.

Flora Zambesiaca. Vol 8, Pt 7. Avicenniaceae, R. Fernandes. Nesogenaceae, M.A. Diniz. Verbenaceae, R. Fernandes. Lamiaceae, R. Fernandes. 2005.

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens 4, didynamous (anterior pair longer) or of equal length, or sometimes reduced to 2, epipetalous, free or rarely fused (monodelphic), usually exserted; filaments hairy or glabrous, rarely appendiculate; anthers basi- or dorsifixed, sometimes with a prominent connective, usually with 2-thecae, rarely with one aborted, introrse; thecae dehiscing longitudinally, rarely apically, parallel to divaricate, sometimes confluent at apex or synthecous Stamens epipetalous, attached within corolla-tube, usually 4, or 2 by abortion and then staminodes often present, more rarely stamens 5–8(16) when 4 often didynamous, usually free; filaments usually exserted from corolla tube, sometimes included within lip of corolla; anther usually dithecous or monothecous by abortion, opening by longitudinal slits or rarely by pores

Note

This family contains many herbs and shrubs used as ornamentals and trees for timber.  Species belonging to genera of the subfamilies Viticoideae and Ajugoideae introduced into the Flora Zambesiaca area, which are grown in cultivation but which have not become naturalized are mentioned below, and are included in the key to the genera of these subfamilies. 1 Alan Paton, introduction, key and cultivated species, genera 2, 15, 16 (with Donald Otieno & Kevin Balkwill), 17–24, 27; Gemma Bramley, genera 6 & 28; Ray Harley, genus 10; Yvette Harvey, genera 5, 7 (with Mattias Iwarsson) & 8; Peter Phillipson, genus 25; Roger Polhill, genera 3, 4, 9, 13, 14; Olof Ryding, genera 11, 12, 26; Fiona Willis, genus 1. Only authors other than Alan Paton indicated in genus footnote.

Distribution

A cosmopolitan family of c. 258 genera, with around 7000 species mainly in temperate zones, and particularly diverse in the Mediterranean region.  Around 35 genera and 300 species are found in the Flora Zambesiaca region. A cosmopolitan family of c.258 genera with around 7000 species, mainly in temperate zones and particularly diverse in the Mediterranean region. Around 34 genera and 330 species are found in the Flora Zambesiaca region; 26 other genera are introduced in cultivation.

Morphology General Habit

Annual or perennial herbs, shrubs, or trees, often aromatic Trees, shrubs or herbs, rarely climbers, aromatic or not Annual or perennial herbs, shrubs, or trees, often aromatic

Morphology Stem

Stems frequently square in cross section Stems often square in cross-section Stems frequently square in cross section

Morphology Leaves

Leaves opposite and decussate or sometimes whorled, very rarely alternate, usually simple rarely pinnately or palmately dissected or compound, usually crenate or serrate, sometimes entire or more deeply toothed, petiolate or sessile, stipules absent Leaves simple, sometimes compound but then digitate or pinnate, opposite, often decussate, sometimes whorled, very rarely alternate, entire, toothed or lobed, petiolate or sessile, rarely forming a basal rosette, exstipulate Leaves opposite and decussate or sometimes whorled, very rarely alternate, usually simple rarely pinnately or palmately dissected or compound, usually crenate or serrate, sometimes entire or more deeply toothed, petiolate or sessile, stipules absent

Morphology Reproductive morphology Inflorescences

Inflorescences terminal or axillary, branched or simple, thyrsoid with determinate cymes arranged along an indeterminate axis, often with paired cymes congested into verticils but sometimes appearing racemose by the reduction of cymes to a single flower, or spicate or capitate by reduction of both the internodes of the inflorescence axis and cyme axes; bracts persistent or not, occasionally coloured; bracteoles present or absent Inflorescence composed of cymes and often arranged in a terminal, lax or congested indeterminate thyrse, which may be paniculate, raceme-like with cymes often 1-flowered, or spike-like, rarely congested into a head; often with bract and sometimes with bracteoles Inflorescences terminal or axillary, branched or simple, thyrsoid with determinate cymes arranged along an indeterminate axis, often with paired cymes congested into verticils but sometimes appearing racemose by the reduction of cymes to a single flower, or spicate or capitate by reduction of both the internodes of the inflorescence axis and cyme axes; bracts persistent or not, occasionally coloured; bracteoles present or absent

Morphology Reproductive morphology Flowers

Flowers zygomorphic occasionally actinomorphic, usually hermaphrodite, hypogynous, usually showy, often with a fleshy nectariferous disk, sometimes resupinate, usually many subtended by a bract, occasionally 1 in bract axils (in reduced cymes), usually pedicellate, rarely sessile Flowers actinomorphic to zygomorphic, hypogynous, usually bisexual-Calyx gamosepalous, sometimes 2-lipped, often enlarging in fruit; lobes 2–many, often 5, equal or unequal, rarely obsolete, some lobes often fused, or lips entire Flowers zygomorphic occasionally actinomorphic, usually hermaphrodite, hypogynous, usually showy, often with a fleshy nectariferous disk, sometimes resupinate, usually many subtended by a bract, occasionally 1 in bract axils (in reduced cymes), usually pedicellate, rarely sessile

Morphology Reproductive morphology Flowers Calyx

Calyx gamosepalous, actinomorphic to bilabiate, persistent, tubular to broadly campanulate or spreading, sometimes bearded at throat; lobes usually 5, sometimes 2 or 3, and often enlarged in fruit, sometimes closing throat Calyx gamosepalous, actinomorphic to bilabiate, persistent, tubular to broadly campanulate or spreading, sometimes bearded at throat; lobes usually 5, sometimes 2 or 3, and often enlarged in fruit, sometimes closing throat

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, tubular, 5-lobed, typically bilabiate with posterior lip 2–4-lobed and anterior lip 1–3-lobed, variously coloured; tube cylindrical, parallel sided, amplified or constricted distally, or saccate below, straight or curved, sometimes annulate within Corolla gamopetalous, tubular, 5-lobed, typically bilabiate with posterior lip 2–4-lobed and anterior lip 1–3-lobed, variously coloured; tube cylindrical, parallel sided, amplified or constricted distally, or saccate below, straight or curved, sometimes annulate within Corolla gamopetalous, actinomorphic to (more often) slightly to strongly zygomorphic, often 2-lipped, rarely 1-lipped; tube short to elongate, rarely spurred, often with a ring of hairs or appendaged within; lobes (2)4–5(16), equal or unequal, one or other lip often concave to galeate

Morphology Reproductive morphology Flowers Androecium

Stamens 4, didynamous (anterior pair longer) or of equal length, or sometimes reduced to 2, epipetalous, free or rarely fused (monodelphic), usually exserted; filaments hairy or glabrous, rarely appendiculate; anthers basi- or dorsifixed, sometimes with a prominent connective, usually with 2-thecae, rarely with one aborted, introrse; thecae dehiscing longitudinally, rarely apically, parallel to divaricate, sometimes confluent at apex or synthecous

Morphology Reproductive morphology Flowers Gynoecium

Gynoecium 2-carpellate, fused to form a pistil; ovary superior, 2-locular but appearing 4-locular due to ovary wall intrusions (false septa), slightly to deeply 4-lobed; ovules 1 in each apparent locule, anatropous, erect, placentation axile (often appearing basal) with ovules laterally attached on the face of the false septum, subterminal (just short of inrolled carpel margins); style 1, gynobasic (arising from central depression of ovary lobes) to terminal, usually bifid at apex; stigmas minute at stylar branch tips Gynoecium 2-carpellate, fused to form a pistil; ovary superior, 2-locular but appearing 4-locular due to ovary wall intrusions (false septa), slightly to deeply 4-lobed; ovules 1 in each apparent locule, anatropous, erect, placentation axile (often appearing basal) with ovules laterally attached on the face of the false septum, subterminal (just short of inrolled carpel margins); style 1, gynobasic (arising from central depression of ovary lobes) to terminal, usually bifid at apex; stigmas minute at stylar branch tips Gynoecium 2-carpellate, often 4-locular by intrusion of carpel wall forming a false septum, or rarely imperfectly 2-locular and free towards apex; ovary entire or lobed with terminal style, more often deeply 4-lobed, the locules often separated and with style gynobasic; ovules usually 4, anatropous to hemianatropous, usually basal or sub-basal, erect, rarely orthotropous, apical, pendulous, borne submarginally on placenta; style usually with 2 equal or unequal stigma-lobes, rarely entire with 1 stigma- lobe vestigial, or stigma capitate or very rarely 4-lobed

Morphology Reproductive morphology Fruits

Fruit a schizocarp splitting into 4 dry nutlets or drupaceous, undivided or 4-lobed with usually 4 pyrenes, sometimes fewer by abortion, subtended by or enclosed within persistent calyx; endosperm present, scant or absent; embryo generally straight, rarely bent Fruit a schizocarp splitting into 4 dry nutlets or drupaceous, undivided or 4-lobed with usually 4 pyrenes, sometimes fewer by abortion, subtended by or enclosed within persistent calyx; endosperm present, scant or absent; embryo generally straight, rarely bent. Fruit a drupe, often with pyrenes, or dry and indehiscent, or separating into two 2-seeded or frequently four 1-seeded mericarps, sometimes fewer by abortion; mericarps (nutlets) often with sculptured, tuberculate, hairy or rarely winged pericarp, mucilage cells often present

Disc

Disk at base of ovary often present, nectariferous

Morphology Reproductive morphology Seeds

Seeds albuminous or exalbuminous; embryo straight or bent.

Labiatae, J.K. Morton. Flora of West Tropical Africa 2. 1963

Morphology General Habit

Herbaceous or rarely woody, often odoriferous

Morphology Stem

Stems usually quadrangular

Morphology Leaves

Leaves opposite or whorled, simple; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, zygomorphic, rarely almost actinomorphic, axillary, whorled, racemose or paniculate

Morphology Reproductive morphology Flowers Calyx

Calyx persistent, of 5 variously united sepals, often 2-lipped

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, hypogynous, tubular; lobes 4-5, imbricate, often forming 2 lips or rarely 1 lip

Morphology Reproductive morphology Flowers Androecium

Stamens inserted in the corolla-tube, 4 or 2; anthers 2-celled, cells often divergent, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovules 4 in each ovary, erect Ovary superior, of 2 deeply lobed carpels, the style (gynobasic) arising from the inner base of the lobes; stigmas mostly bifid

Morphology Reproductive morphology Fruits

Fruit of 4 achene-like nutlets, free or cohering in pairs

Morphology Reproductive morphology Seeds

Seeds with usually straight embryo without endosperm, or the latter very scanty

George R. Proctor (2012). Flora of the Cayman Isands (Second Edition). Royal Botanic Gardens, Kew

Morphology General Habit

Herbs or shrubs, rarely trees, often aromatic and usually with 4-angled stems; leaves opposite or whorled, simple, and entire, crenate, serrate or lobed, often glandular; stipules absent. A ring-shaped or unilateral disc or gland present

Morphology Reproductive morphology Flowers

Flowers mostly perfect, rarely unisexual, solitary in the leaf-axils or more often in compact cymes, these axillary or terminal, often condensed into whorls spaced on the axes of racemes or panicles, or crowded into a spike or head; bracts small or large; bracteoles often present

Morphology Reproductive morphology Flowers Calyx

Calyx tubular or bell-shaped, regular or 2-lipped, basically 5- lobed but the upper 3 teeth or lobes often united

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, tubular at the base, the limb basically 5-lobed but usually 2-lipped, the 2 upper lobes usually to form an entire or notched hood, or rarely absent; the 3 lower lobes partly or wholly united to form the lower lip

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens usually 4 in 2 pairs inserted at different levels of the corolla-tube, or sometimes only 2; anthers 2-locular or sometimes 1-locular by abortion, opening lengthwise inwardly

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, of 2 bilobed carpels, becoming more or less 4-lobed and 4-locular, each cavity with 1 basal, erect ovule; style slender, central and usually gynobasic (i.e.) arising basally between the lobes of the ovary); stigma 2-lobed or entire

Morphology Reproductive morphology Fruits

Fruit of 4 free or paired, dry, 1-seeded nutlets, usually enclosed by the persistent calyx; seeds with little or no endosperm, the embryo straight and with flat cotyledons.

Distribution

A large, cosmopolitan family of about 200 genera and 3,200 species.

Use

The family contains many herbs and shrubs used as ornamentals and trees for timber. It is best known for its aromatic herbs, many of which belong in subfamily Nepetoideae. A good review of uses and underlying phytochemistry can be found in Harley et al. (in Kadereit, Fam. Gen. Vasc. Pl. 7: 167–275, 2004) along with more detailed references.