Verbenaceae J.St.-Hil.

Timothy M. A. Utteridge and Laura V. S. Jennings (2022). Trees of New Guinea. Kew Publishing. Royal Botanic Gardens, Kew

Distribution

A family of about 34 genera and approximately 1200 species, mainly from America, with a few genera in Europe, mainland Asia and Africa. In New Guinea, only four genera with about five species are recorded.

Recognition

They are all introduced and often weedy and usually grow in disturbed areas; only two genera have species which can be considered as trees. The family has opposite leaves usually with serrate margins, stipules are lacking (though there is often a line across the node), and with flowers borne in racemes or heads, a trumpet-shaped corolla with anthers usually included within or just exceeding the corolla.

Morphology General Habit

Herbs, shrubs or small trees, stems terete or quadrangular, unarmed or sometimes with prickles or spines

Morphology General

Sap absent

Morphology Leaves Stipules

Stipules absent

Morphology Leaves

Leaves opposite, usually decussate, simple, margins entire to serrate

Morphology Reproductive morphology Inflorescences

Inflorescences indeterminate, racemose, terminal or axillary, dense or lax, sometimes compressed into a spike or compact head. Flowers bisexual, 4- or 5-merous; calyx tubular or campanulate, usually persistent in fruit; corolla trumpet shaped (or salver-shaped), zygomorphic to occasionally somewhat bilabiate, limb spreading with lobes equal or not, colourful; stamens 4, inserted on the corolla, usually didynamous, sometimes d2 fertile and 2 staminodes, not or only just exceeding the corolla tube; ovary superior, seated on a nectariferous disk, syncarpous, (1–)2(–4) carpellate, style terminal-Fruit drupaceous (in New Guinea) or a dry or fleshy schizocarp, separating at maturity into two or four 1-seeded mericarps or four 2-seeded pyrenes.

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

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

Morphology Leaves

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

Morphology Reproductive morphology Inflorescences

Inflorescence racemose, dense or lax, sometimes spicate or a compact head, axillary and/or terminal simple or compound; bracts usually present, inconspicuous or showy

Morphology Reproductive morphology Flowers

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

Morphology Reproductive morphology Fruits

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.

Note

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 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.

Diagnostic

"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 3. Ovary 1-carpellate, ovules 2 — 43. Ovary 2 or 4-carpellate, ovules 4 or 8 — 17 4. Fruit a dry or fleshy schizocarp of two 1-seeded mericarps — 54. Fruit drupaceous, with one 2-locular, 2-seeded  pyrene — 15 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""—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 —109. 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 — 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 —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 — 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 18. Fertile stamens 2; calyx slightly inflated in fruit — Hierobotana18. 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 — 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 reticulateexocarp... Junellia 21.  Inflorescence a simplespike 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 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 — 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 stigma tiferous, and the other lobe reduced — Priva 25. Calyx enlarged in fruit, completely surrounding the fruit — Parodianthus25. 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 — 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 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 — 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" 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. 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. 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. Leaves often aromatic.

Distribution

Native 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.

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.

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

Morphology General Habit

Herbs, shrubs or trees, or sometimes woody vines; branches often 4-angled, sometimes armed with thorns; leaves opposite or whorled, simple or palmately compound, the margins entire or variously toothed, lobed or incised; stipules absent

Morphology Reproductive morphology Flowers

Flowers usually perfect, in axillary or terminal spikes, racemes, cymes or panicles, or sometimes in heads

Morphology Reproductive morphology Flowers Calyx

Calyx 4–5- toothed or -lobed, persistent and often enlarging in fruit; corolla gamopetalous, regular or irregular, funnel-or salver-shaped, 4–5-lobed, often more or less 2-lipped

Morphology Reproductive morphology Flowers Androecium Stamens

Stamens usually 4 (2 long and 2 short) or rarely only 2, the filaments inserted in the corolla-tube; anthers 2- celled, opening lengthwise; staminodes sometimes present

Morphology Reproductive morphology Flowers Gynoecium Ovary

Ovary superior, usually 2- carpellate but often 4-lobed and usually 2–5-celled (rarely 1-celled), with 1 or 2 ovules in each cavity, basally or laterally attached to the placentas; style terminal and solitary

Morphology Reproductive morphology Fruits

Fruit a drupe with 1 or more stones, or a dry schizocarp separating into 2 or 4 nutlets.

Distribution

A rather large, widely distributed family of about 100 genera and 2,600 or more species.