Flavio França* and Sandy Atkins**
Universidade Estadual de Feira de Santana, Feira de Santana, Bahia, Brasil.*Royal Botanic Gardens, Kew, UK.**
Herbs, shrubs or small trees, sometimes lianas, stems terete or quadrangular with varied indumentum, unarmed, or sometimes with prickles or spines; eglandular or with resinous glands on stems, leaves, bracts, calyx or corolla. Leaves opposite, usually decussate, rarely ternate, sometimes verticillate or fascicled, sessile or petiolate, sometimes much reduced, scale-like, or spinose, simple to pinnatisect, margins entire, dentate, crenate, lobed or incised, stipules absent. Inflorescence racemose, dense or lax, sometimes spicate or a compact head, axillary and/or terminal, simple or compound; bracts usually present, inconspicuous or showy. Flowers hermaphrodite, rarely plants dioecious, zygomorphic or sub-actinomorphic, 4- or 5-merous; calyx tubular or campanulate, 4- or 5- (rarely 2-) lobed or annular or sub-truncate, persistent in fruit; corolla hypocrateriform or infundibular, zygomorphic, 4- or 5-lobed, limb spreading, with lobes equal or not, tube sometimes enlarged in the middle or at throat; stamens 4 (5), inserted on the corolla, alternate with the petals, usually didynamous, or reduced to 2, sometimes with 1 or more staminodes present, included, or only slightly exserted; anthers 2-locular, basifixed or dorsifixed; thecae parallel or divergent, dehiscing longitudinally, connective reduced or conspicuous and with or without glandular appendage; ovary superior, glabrous, rarely pilose at apex, seated on a nectariferous disk, syncarpous, 2-carpellate, (or 1-carpellate by abortion of the abaxial carpel, or 4-carpellate by elaboration as in Duranta); carpels 2-locular by the inrolling of the carpel margin; ovules 1 or 2 per locule, erect, rarely pendulous, fixed at base of locule, or at upper part of locule, anatropous, rarely hemi-anatropous, or sub-anatropous, or orthotropous; style terminal, 2-lobed or simple with stigma capitate, oblique or lateral, rarely weakly 4-lobed. Fruit a dry or fleshy schizocarp separating at maturity into two or four 1-seeded mericarps, or into two 2-locular and 2-seeded mericarps, or a drupaceous fruit with one or two pyrenes with one locule and one seed, or with two or four pyrenes, each with two locules and two seeds.
Notes on delimitation
- This circumscription of the family follows Verbenaceae, in The Families and Genera of Vascular Plants Vol. VII Lamiales (ed. Kubitzki) 2004.
- The case is set out there, from molecular and anatomical evidence, for the removal of all the cymose-inflorescenced genera from the traditional circumscription of the family, and placing them in the Labiatae.
- This leaves a more or less monophyletic group of 34 genera, only 1 of which has its origin outside the New World.
Distribution in the Neotropics
- A family of 34 genera with about 1,200 species, mainly from the New World with a few groups in Europe, Asia, Africa and Madagascar.
- The distribution of the New World genera is varied.
- Verbena L. which has its greatest species diversity in the New World, also has species in Europe, Asia and North Africa.
- The closely-allied Glandularia J.F.Gmel. has a disjunct North-South American distribution.
- Lantana L. has species in Africa, as do Lippia L. and Priva Adans.
- The main pattern for the rest of the genera seems to be temperate and tropical America, as found in, for example, Tamonea Aubl. which occurs from Mexico and the Caribbean to northern South America, Brazil and eastern Bolivia.
- Some genera are confined to the southern part of South America, i.e. Urbania Phil., Acantholippia Griseb., Diostea Miers, Lampaya Philippi ex Murillo, all of which are restricted to Argentina and Chile.
Distinguishing characters (always present)
The key characters for the family are:
- Opposite, simple leaves.
- Racemose inflorescence, usually in the form of a head or spike.
- Hermaphrodite flowers.
- Stamens 2 - 4 (5), included.
- Tubular/campanulate calyx.
- Tubular flower with spreading limb.
Other important characters
- Leaves often aromatic.
Key differences from similar families
Superficially resembling the Labiatae, the Verbenaceae can be distinguished from it by:
- Racemose inflorescence.
- Included stamens.
- Thickened stigma lobes with conspicuous stigmatic tissue.
- Colporate pollen.
Number of genera
25 Neotropical genera:
- Aloysia Palau
- Baillonia Bocq.
- Bouchea Cham.
- Casselia Nees & Mart.
- Citharexylum L.
- Duranta L.
- Glandularia J.F.Gmel.
- Hierobotana Briq.
- Junellia Moldenke
- Lampaya Philippi ex Murillo
- Lantana L.
- Lippia L.
- Nashia Millsp.
- Phyla Lour.
- Petrea L.
- Pitraea Turcz.
- Priva Adans.
- Recordia Moldenke
- Rehdera Moldenke
- Stachytarpheta Vahl
- Tamonea Aubl.
- Verbena L.
- Verbenoxylum Tronc.
- Xeroaloysia Tronc.
- Xolocotzia Miranda
Useful tips for generic identification
Key to genera of Neotropical Verbenaceae
1. Calyx corolla-like, often syncolorous with corolla, 5-lobed with lobes reticulately-veined and extending beyond the corolla lobes; leaves asperous … 21. Calyx not brightly coloured, lobes not extending beyond corolla lobes; leaves generally smooth; if asperous then not with accompanying characters … 3
2. Lianas, scandent shrubs or trees; calyx with "inner" calyx in the form of a coronet (0.4 -- 3.5 mm high) situated at the inner base of the calyx lobes; anther thecae introrse with enlarged connective … Petrea2. Erect shrub or small tree; calyx without "inner" calyx; anther thecae parallel without enlarged connective … Xolocotzia
5. Fertile stamens 4 … 65. Fertile stamens 2 with 2 staminodes … Stachytarpheta
6. Fruit a fleshy schizocarp with thickened pericarp and mesocarp, fissure line between the 2 mericarps usually visible … 76. Fruit a dry schizocarp with thin dry pericarp and mesocarp, readily splitting at maturity into 2 mericarps … 11
7. Inflorescence a long-pedunculate, axillary, (1)2--6-flowered raceme; posterior stamens with divergent thecae and with a thick, glandular connective forming a dorsal "crest"…Casselia7. Inflorescence a sessile or short-pedunculate, terminal or axillary spike or fascicle; posterior stamens with parallel thecae and without glandular connective … 8
10. Stamens inserted in middle of corolla tube, filamentous; stigma subglobose; tall shrubs with genistoid habit … Diostea10. Stamens inserted in upper part of corolla tube just below the throat, almost sessile; stigma bilobed with anterior lobe thick and papillose, and reduced posterior lobe; low shrubs forming dense thickets …Lampaya
11. Perennial creeping herb, rooting at nodes; indumentum on calyx of uncinate hairs, some species with medifixed hairs on stem, leaves and bracts … Phyla11. Herbs, subshrubs or shrubs; indumentum only of simple hairs … 12
12. Plants of semi-deserts, xerophytic shrubs with small (up to 1.6 mm long), fleshy leaves …Acantholippia 12. Plants, herbs or shrubs, not xerophytic, without reduced leaves … 13
13. Calyx 2 -- 4-lobed or dentate; corolla 4-lobed; style short, not more than 3 times longer than ovary, stigma oblique or lateral; fruit ovoid or subrotund, rounded at apex … 1413. Calyx 5-lobed or dentate; corolla 5-lobed; style long, more than 3 times longer than ovary; stigma sub-bilobed with one lobe much reduced; fruit linear-oblong, beaked at apex … Bouchea
14. Calyx distinctly 4-toothed; teeth acute or subulate; corolla ± actinomorphic, with lobes ± equal; inflorescence a racemose spike, erect or pendant, flowers very shortly pedicellate … Aloysia14. Calyx obscurely 2- or 4-lobed; corolla weakly zygomorphic, more or less 2-lipped; inflorescence a compact head or spike with sessile flowers … Lippia
15. Glabrous shrubs with cylindric stems; leaves reduced or absent; corolla 5-lobed, ± regular …Neosparton15. Shrubs or herbs, not glabrous with stems 4-sided; leaves normal and present; corolla 4-lobed... 16
16. Corolla 4-lobed, ± regular, anterior lobe emarginate; stigma capitate; infloresence a lax racemiform spike; flowers minutely pedicellat … Xeroaloysia16. Corolla 4(--5)-lobed, 2-lipped; lobes obtuse or emarginate; stigma oblique or lateral; inflorescences pedunculate heads, dense spikes or sessile clusters; flowers sessile … Lantana
17. Fruit a dry schizocarp, maturing into four 1-seeded mericarps … 1817. Fruit a dry or fleshy schizocarp separating at maturity into two 2-locular, 2-seeded mericarps or fruit a fleshy drupe with 1 pyrene with four 1-seeded locules or fruit a fleshy drupe with 2 or 4 pyrenes (or less by abortion), pyrenes 2-locular and 2-seeded … 22
19. Shrubs, sometimes forming cushions only a few centimetres high; often with deep, thick, woody rootstock; leaves reduced; sometimes with spines … 2019. Perennial or annual herbs, subshrubs, not cushion-forming; without deep, woody rootstock, without spines … 21
20. Calyx with long, hygroscopic hairs which completely hide the corolla; fruit globose with smooth exocarp … Urbania20. Calyx without long hygroscopic hairs; fruit oblong with reticulate exocarp... Junellia
21. Inflorescence a simple spike or panicle of spikes, often branched; style short, not more than 3 times longer than ovary … Verbena21. Inflorescence a contracted head, unbranched; style long, more than 3 times longer than the ovary …Glandularia
23. Perennial herbs … 2423. Trees or shrubs … 25
24. Plant with deep tuberous rootstock; style recurved at apex, stigma lateral, papillose … Pitraea24. Plant without deep tuberous rootstock; style 2-lobed at apex with one lobe longer and stigmatiferous, and the other lobe reduced … Priva
27. Corolla lobes ± regular; 4 fertile stamens and 1 staminode; fruit obovoid, attenuate at base …Verbenoxylum27. Corolla ± 2-lipped; 4 fertile stamens and no staminode; fruit ovoid or subglobose, not attenuate at base … Dipyrena
28. Perennial herbs or subshrubs; posterior stamens with connective glandular, no staminode present; fruit a fleshy drupe with 1 pyrene with four 1-seeded locules; fruit often horned or woody … Tamonea28. Shrubs or small trees; posterior stamens with connective not glandular, staminode often present; fruit a fleshy drupe with 2 or 4 pyrenes (or less by abortion), pyrenes 2-locular and 2-seeded; fruit not horned or woody … 29
30. Inflorescence a reduced raceme of 1 -- 2 (5) flowers; stigma 2-lobed (hooked) with one lobe stigmatiferous … Rhaphithamnus30. Inflorescence a spike, many-flowered; stigma capitate, obscurely 2-lobed... 31
31. Stamens ± sessile, included; stigma short, not more than twice as long as ovary, included; fruit drupaceous with usually two 2-locular pyrenes … Citharexylum31. Stamens filamentous, posterior pair exserted; style long, more than 2 times longer than ovary, exserted; fruit dry with usually one 2-locular pyrene... Recordia
Notable genera and distinguishing features
The most common genera in the Neotropics are:
- Lantana (large aromatic shrubs with head-like inflorescences, usually multicoloured, fruit somewhat fleshy).
- Lippia (smaller aromatic shrubs, fruit dry).
- Phyla (procumbent, creeping herb, indumentum of medifixed hairs.
- Priva (small herb, not aromatic, inflorescence a lax spike).
- Stachytarpheta (herbs or sub-shrubs, terminal, spicate inflorescence).
- Duranta (erect or subscandent shrub or small tree, often spinose, inflorescence an elongate lax spike.
- Citharexylum (trees and shrubs, sometimes with spines, inflorescence long-spicate, many-flowered, fruit often brightly-coloured, sitting in a cupped calyx.
Antino, P.D.; R.M. Harley & S.J. Wagstaff. 1992b. Genera of Labiatae: status and classification. In R.M. Harley & T. Reynolds (eds.), Advances in Labiate science, p. 511-522.
Atkins, S. (2004). Verbenaceae in (Kubitzki ed.) The Families and Genera of Flowering Plants, Vol. VII, Lamiales. Berlin: Springer.
Atkins, S. 2005. The genus Stachytarpheta (Verbenaceae) in Brazil . Kew Bull. 60: 161-272.
Harley, R.M.; Atkins, S.; Budantsev, A.; Cantino, P.D.; B.J. Conn, R. Grayer; M.M. Harley, R. de Kok, T. Krestovskaya, R. Morales, A. Paton, O. Ryding and T. Upson. 2004. Labiatae In K. Kubitzki et al. The families and genera of vascular plants (Lamiales except Acanthaceae). Berlin: Springer.
Jansen-Jacobs, M.J. 1988. Verbenaceae. In A.R. Görts-Van Rijn, Flora of the Guianas. 116p. Koenigstein: Koeltz Scientific Books.
Junell, S. 1934. Zur gynäceummorphologie und systematik der Verbenaceen und Labiaten. Symbolae Botanicae Upsalienses, 4: 1-219.
Lima, A.B. & F.França. 2007. Flora da Bahia: Priva Adans. - Verbenaceae. Sitientibus, ser. Ciências Biológicas, 7: 32-35.
Lopez-Palacios, S. 1977. Flora de Venezuela: Verbenaceae. 654p. Merida: Universidad de los Andes.
Rueda, R.M. 1994. Systematics and evolution of the genus Petrea (Verbenaceae). Ann. Missouri Bot. Gard. 81: 610-652.
Salimena, F. 2000. Revisão taxonômica de Lippia L. sect. Rhodolippia Schauer (Verbenaceae). Tese de Doutorado. Universidade de São Paulo.
Salimena-Pires, F. & A.M. Giulietti. 1998. Flora da Serra do Cipó, Minas Gerais: Verbenaceae. Boletim de Botânica da Universidade de São Paulo, 17: 155-186.
Schauer, J.C. 1851. Verbenaceae In C.F. Martius, A. Eichler & I. Urban, Flora brasiliensis, 9 (10): 169-308; t. 32-50.
Silva, T.R.S. 1999. Redelimitação e revisão taxonômica do gênero Lantana L. (Verbenaceae) no Brasil. Tese de Doutorado. Universidade de São Paulo.
How to cite
França, F. & Atkins, S. (2009). Neotropical Verbenaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. http://www.kew.org/science/tropamerica/neotropikey/families/Verbenaceae.htm.
Click images to enlarge
Flower and foliage of Aloysia sp. © John Wood, Darwin Initiative Project 161/11/011.
Inflorescences of Aloysia sp. © John Wood, Darwin Initiative Project 161/11/011.
Inflorescence of Aloysia spp.© John Wood, Darwin Initiative Project 161/11/011.
Inflorescence of Aloysia sp. © John Wood, Darwin Initiative Project 161/11/011.
Fruit of Citharexylon andinum © John Wood, Darwin Initiative Project 161/11/011.
Fruits of Citharexylon andinum © John Wood, Darwin Initiative Project 161/11/011.
Flower of Citharexylon ilicifolium © John Wood, Darwin Initiative Project 161/11/011.
Fruit of Citharexylon ilicifolium © John Wood, Darwin Initiative Project 161/11/011.
Lantana cabrerae © John Wood, Darwin Initiative Project 161/11/011.
Lantana cabrerae © John Wood, Darwin Initiative Project 161/11/011.
Flowering Branch of Lantana canescens © John Wood, Darwin Initiative Project 161/11/011.
Lantana hypoleuca © John Wood, Darwin Initiative Project 161/11/011.
Flowers of Lantana hypoleuca © John Wood, Darwin Initiative Project 161/11/011.
Inflorescences of Lantana trifolia © John Wood, Darwin Initiative Project 161/11/011.
Flowers of Lippia bolivinana © John Wood, Darwin Initiative Project 161/11/011.
Yellow-flowered Lippia sp. © John Wood, Darwin Initiative Project 161/11/011.
Inflorescence of Verbena cochabambensis © John Wood, Darwin Initiative Project 161/11/011.
Habit of Verbena cochabambensis © John Wood, Darwin Initiative Project 161/11/011.
Inflorescence of Verbena sp. © John Wood, Darwin Initiative Project 161/11/011.
Red-flowered, Verbena sp. © John Wood, Darwin Initiative Project 161/11/011.