Verbenaceae J.St.-Hil.

França, F. & Atkins, S. (2009). Neotropical Verbenaceae.

Morphology

Description

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.

General Description

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.

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

Status

• Native

Distribution

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.

Diagnostic

Other important characters

• Leaves often aromatic.

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/campanulatecalyx.
• Tubular flower with spreading limb.

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.

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 — 2
1. 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 — Petrea
2. Erect shrub or small tree; calyx without "inner" calyx; anther thecae parallel without enlarged connective — Xolocotzia

3. Ovary 1-carpellate, ovules 2 — 4
3. Ovary 2 or 4-carpellate, ovules 4 or 8 — 17

4. Fruit a dry or fleshy schizocarp of two 1-seeded mericarps — 5
4. Fruit drupaceous, with one 2-locular, 2-seeded  pyrene — 15

5. Fertile stamens 4 — 6
5. 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 — 7
6. 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"—Casselia
7. Inflorescence a sessile or short-pedunculate, terminal or axillary spike or fascicle; posterior stamens with parallel thecae and without glandular connective — 8

8. Corolla campanulate, 4-lobed; occurring in Cuba and the Bahama archipelago ... Nashia
8. Corolla hypocrateriform, 5-lobed; occurring in Brazil, Paraguay, Bolivia, Argentina and Chile — 9

9. Fruiting calyx enlarged and completely surrounding fruit —10
9. Fruiting calyx not enlarged, forming a short cup around the fruit — Baillonia

10. Stamens inserted in middle of corolla tube, filamentous; stigma subglobose; tall shrubs with genistoid habit — Diostea
10. 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 — Phyla
11. 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 — 14
13. 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 — Aloysia
14. 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 —Neosparton
15. 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 — Xeroaloysia
16. 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 — 18
17. 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

18. Fertile stamens 2; calyx slightly inflated in fruit — Hierobotana
18. Fertile stamens 4; calyx not inflated in fruit — 19

19. Shrubs, sometimes forming cushions only a few centimetres high; often with deep, thick, woody rootstock; leaves reduced; sometimes with spines — 20
19. 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 — Urbania
20. Calyx without long hygroscopic hairs; fruit oblong with reticulateexocarp... Junellia

21.  Inflorescence a simplespike or panicle of spikes, often branched; style short, not more than 3 times longer than ovary — Verbena
21. Inflorescence a contracted head, unbranched; style long, more than 3 times longer than the ovary —Glandularia

22.  Fruit a dry or fleshy schizocarp separating at maturity into two 2-locular, 2-seeded mericarps — 23

22. 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 ... 28

23. Perennial herbs — 24
23. Trees or shrubs — 25

24. Plant with deep tuberous rootstock; style recurved at apex, stigma lateral, papillose — Pitraea
24. Plant without deep tuberous rootstock; style 2-lobed at apex with one lobe longer and stigma tiferous, and the other lobe reduced — Priva

25. Calyx enlarged in fruit, completely surrounding the fruit — Parodianthus
25. Calyx not enlarged in fruit —26

26. Fruiting calyx caduceus — Rehdera 
26. Fruiting calyx forming a cup — 27

27. Corolla lobes ± regular; 4 fertile stamens and 1 staminode; fruit obovoid, attenuate at base —Verbenoxylum
27. 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 — Tamonea
28. 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

29. Stigma capitate, obscurely 4-lobed; fruit with four 2-locular, 2-seeded pyrenes ... Duranta
29. Stigma subcapitate or 2-lobed; fruit with two 2-seeded pyrenes ... 30

30. Inflorescence a reduced raceme of 1 -- 2 (5) flowers; stigma 2-lobed (hooked) with one lobe stigma tiferous — Rhaphithamnus
30. 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 — Citharexylum
31. 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 laxspike).
• Stachytarpheta (herbs or sub-shrubs, terminal, spicate inflorescence).
• Duranta (erect or subscandent shrub or small tree, often spinose, inflorescence an elongate laxspike.
• Citharexylum (trees and shrubs, sometimes with spines, inflorescence long-spicate, many-flowered, fruit often brightly-coloured, sitting in a cupped calyx.

Literature

Important literature

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.

Verbenaceae, B. Verdcourt. Flora of Tropical East Africa. 1992

Morphology General Habit

Herbs, shrubs, trees or woody climbers (varying from small annual herbs to large forest trees), sometimes thorny

Morphology Leaves

Leaves predominantly opposite, sometimes whorled, rarely alternate, simple or digitately compound, entire, serrate or variously lobed, frequently aromatic; stipules absent Flowers mostly ± irregular and 2-lipped, sometimes almost regular, usually 4–5-merous, usually hermaphrodite in often bracteate panicles, cymes or spikes

Morphology Reproductive morphology Flowers Calyx

Calyx tubular or campanulate, lobed or subentire, sometimes 2-lipped

Morphology Reproductive morphology Flowers Corolla

Corolla usually with narrow tube and spreading limb, minute to quite large and showy; lobes imbricate in bud

Morphology Reproductive morphology Flowers Androecium

Stamens 4, didynamous, rarely 2 or, in a few genera, the same number as the corolla-lobes, inserted in the corolla-tube and alternate with lobes; filaments free; anthers free or connivent, dorsifixed, 2-thecous, the thecae mostly parallel, opening by longitudinal slits, introrse

Morphology Reproductive morphology Flowers Nectaries

Disk usually present, sometimes conspicuous

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, sessile, entire or slightly 4-lobed, 2(–9)-locular, usually soon 4(or more)-locular by development of false septa, with axile or free central (in Avicennia) placentation; ovules 2 in each true locule (2 per carpel), erect or pendulous (in >i>Avicennia); style terminal, simple and entire or shortly 2(rarely 4–5)-lobed

Morphology Reproductive morphology Fruits

Fruit a drupe, a capsule (infrequent) or dividing at maturity into 2 or 4 nutlets

Morphology Reproductive morphology Seeds

Seeds without or with much reduced endosperm or present and fleshy (in >i>Avicennia); testa membranous

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, epipetalous; filaments distinct; anthers dorsifixed, dehiscing longitudinally, introrse

Distribution

A family of c. 36 genera, mainly tropical and subtropical, with only a limited number of representatives (usually herbs) in temperate regions.

Morphology General Habit

Annual or perennial herbs or shrubs, rarely trees or vines, often aromatic, sometimes armed with prickles and/or spines Annual or perennial herbs or shrubs, rarely trees or vines, often aromatic, sometimes armed with prickles and/or spines

Morphology Stem

Stems frequently square in cross section Stems frequently square in cross section

Morphology Leaves

Leaves simple, opposite or sometimes whorled; lamina entire or serrate to lobed; stipules absent Leaves simple, opposite or sometimes whorled; lamina entire or serrate to lobed; stipules absent

Morphology Reproductive morphology Inflorescences

Inflorescence indeterminate, terminal or axillary, lax or condensed, racemose, spicate or capitate Inflorescence indeterminate, terminal or axillary, lax or condensed, racemose, spicate or capitate

Morphology Reproductive morphology Flowers

Flowers zygomorphic, hermaphrodite, hypogynous, usually with an inconspicuous nectariferous disk, each solitary in the axil of a single bract, sessile or pedicellate Flowers zygomorphic, hermaphrodite, hypogynous, usually with an inconspicuous nectariferous disk, each solitary in the axil of a single bract, sessile or pedicellate

Morphology Reproductive morphology Flowers Calyx

Calyx gamosepalous, persistent, campanulate or tubular, 5-lobed or 5-toothed, sometimes expanded or inflated in fruit Calyx gamosepalous, persistent, campanulate or tubular, 5-lobed or 5-toothed, sometimes expanded or inflated in fruit

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, hypocrateriform (with very narrow tube and abruptly spreading limb), funnel-shaped in Verbena, 5-lobed[a1] , often somewhat bilabiate (with a 2-lobed upper lip and 3-lobed lower lip), variously coloured Corolla gamopetalous, hypocrateriform (with very narrow tube and abruptly spreading limb), funnel-shaped in Verbena, 5-lobed, often somewhat bilabiate (with a 2-lobed upper lip and 3-lobed lower lip), variously coloured

Morphology Reproductive morphology Flowers Androecium

Stamens 4, didynamous, epipetalous; filaments distinct; anthers dorsifixed, dehiscing longitudinally, introrse

Morphology Reproductive morphology Flowers Gynoecium

Gynoecium of 1 pistil, 2-carpellate, though sometimes reduced to one; ovary superior, 2-locular but typically appearing 4-locular due to ovary wall intrusions (false septa), usually slightly to moderately 4-lobed[a2] ; ovules 1 in each apparent locule, usually anatropous, erect, placentation axile with ovules attached to the margin of the false septum (directly to carpel margins, not sub terminal); style 1, terminal; stigma 1, capitate or lobed Gynoecium of 1 pistil, 2-carpellate, though sometimes reduced to one; ovary superior, 2-locular but typically appearing 4-locular due to ovary wall intrusions (false septa), usually slightly to moderately 4-lobed; ovules 1 in each apparent locule, usually anatropous, erect, placentation axile with ovules attached to the margin of the false septum (directly to carpel margins, not sub terminal); style 1, terminal; stigma 1, capitate or lobed

Morphology Reproductive morphology Fruits

Fruit a drupe with 2 or 4 pyrenes, or a schizocarp splitting into 2 or 4 nutlets (mericarps), enclosed or subtended by persistent calyx; endosperm absent; embryo straight, oily Fruit a drupe with 2 or 4 pyrenes, or a schizocarp splitting into 2 or 4 nutlets (mericarps), enclosed or subtended by persistent calyx; endosperm absent; embryo straight, oily.