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Verbena
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| Verbena | |
|---|---|
| Common vervain, Verbena officinalis | |
| Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Asterids |
| Order: | Lamiales |
| Family: | Verbenaceae |
| Genus: | Verbena L.[1] |
| Type species | |
| Verbena officinalis L.
| |
| Species | |
|
See text | |
| Synonyms[2] | |
|
List
| |
Verbena (/vərˈbiːnə/),[3] also known as vervain or (in obsolete spelling) verveine, is a genus in the family Verbenaceae. It contains about 150 species of annual and perennial herbaceous or semi-woody flowering plants. The majority of the species are native to the Americas and Asia; however, Verbena officinalis, the common vervain or common verbena, is the type species and native to Europe.
Naming
[edit]In English, the name Verbena is usually used in the United States and the United Kingdom; elsewhere, the terms verveine or vervain are in use.[4]
Description
[edit]
Verbena is a herbaceous flowering plant, belonging to the Verbenaceae family, and may be annual or perennial depending on the species. The leaves are usually opposite, simple, and in many species hairy, often densely so. The flowers are small, with five petals, and borne in dense spikes. Typically some shade of blue, they may also be white, pink, or purple, especially in cultivars.[5]
The genus can be divided into a diploid North American and a polyploid South American lineage, both with a base chromosome number of seven. The European species is derived from the North American lineage. It seems that verbena as well as the related mock vervains (Glandularia) evolved from the assemblage provisionally treated under the genus name Junellia; both other genera were usually included in the Verbenaceae until the 1990s.[6] Intergeneric chloroplast gene transfer by an undetermined mechanism – though probably not hybridization – has occurred at least twice from vervains to Glandularia, between the ancestors of the present-day South American lineages and once more recently, between V. orcuttiana or V. hastata and G. bipinnatifida. In addition, several species of verbena are of natural hybrid origin; the well-known garden vervain/verbena has an entirely muddy history. The relationships of this close-knit group are therefore hard to resolve with standard methods of computational phylogenetics.[7]
Cultivation
[edit]
Some species, hybrids and cultivars of verbena are used as ornamental plants. They are drought-resistant, tolerating full to partial sun, and enjoy well-drained, average soils. Plants are usually grown from seed. Some species and hybrids are not hardy and are treated as half-hardy annuals in bedding schemes.[8]
They are valued in butterfly gardening in suitable climates, attracting Lepidoptera such as the Hummingbird hawk-moth, Chocolate albatross, or the Pipevine swallowtail, and also hummingbirds, especially V. officinalis, which is also grown as a honey plant.
The hybrid cultivars "Silver Anne"[9] and "Sissinghurst"[10] have gained the Royal Horticultural Society's Award of Garden Merit.
Pests and diseases
[edit]For some verbena pathogens, see List of verbena diseases. Cultivated verbenas are sometimes parasitized by sweet potato whitefly (Bemisia tabaci) and spread this pest to other crops.
Uses
[edit]Although verbena ("vervain") has been used in herbalism and traditional medicine, usually as an herbal tonic, there is no high-quality evidence for its effectiveness.[11] Verbena has been listed as one of the 38 plants used to prepare Bach flower remedies,[12] a kind of alternative medicine promoted for its effect on health. According to Cancer Research UK, "essence therapists believe that using essences can help to increase your mental, emotional and spiritual wellbeing. However, essences are not used to prevent, control, or cure cancer or any other physical condition."[13]
The essential oil of various species, mainly common vervain, is traded as "Spanish verbena oil". Considered inferior[14] to oil of lemon verbena (Aloysia citrodora) in perfumery,[11] it is of some commercial importance for herbalism.
In culture
[edit]Verbena has long been associated with divine and other supernatural forces. It was called "tears of Isis" in ancient Egypt, and later called "Hera's tears". In ancient Greece, it was dedicated to Eos Erigineia. The generic name is the Latin term for a plant sacred to the ancient Romans.[15][16] Pliny the Elder describes verbena presented on Jupiter altars; it is not entirely clear if this referred to a verbena rather than the general term for prime sacrificial herbs.[non-primary source needed]
Nulla tamen Romae nobilitatis plus habet quam hiera botane. aliqui aristereon, nostri verbenacam vocant. haec est quam legatos ferre ad hostes indicavimus; hac Iovis mensa verritur, domus purgantur lustranturque. genera eius duo: foliosa, quam feminam putant, mas rarioribus foliis.[17]
No plant however is so renowned among the Romans as hiera botane ('sacred plant'). Some call it aristereon, and Latin writers verbenaca. This is the plant which I mentioned as carried to the enemy by envoys. With this the table of Jupiter is swept, and homes are cleansed and purified. There are two kinds of it; one has many leaves and is thought to be female, the other, the male, has fewer leaves.[18]
Pliny the Elder notes "the Magi especially make the maddest statements about the plant: that [among other things] a circle must be drawn with iron round the plant".[18] The common names of verbena in many Central and Eastern European languages often associate it with iron. These include for example the Dutch IJzerhard ("iron-hard"), Danish Læge-Jernurt ("medical ironwort"), German Echtes Eisenkraut ("true ironherb"), Slovak Železník lekársky ("medical ironherb"), and Hungarian vasfű ("iron grass").
In the early Christian era, folk legend stated that V. officinalis was used to stanch Jesus' wounds after his removal from the cross. It was consequently called "holy herb"[19] or (e.g. in Wales) "Devil's bane".[20]
According to the Wiccan writer Doreen Valiente, Vervain flowers signify the goddess Diana and are often depicted on cimaruta, traditional Italian amulets.[21] In the 1870 The History and Practice of Magic by "Paul Christian" (Jean-Baptiste Pitois), it is employed in the preparation of a mandragora charm.[22] The book also describes its antiseptic capabilities (p. 336), and use as a protection against spells (pp. 339, 414).[23] Romani people use vervain for love and good luck.[24]
While common vervain is not native to North America, it has been introduced there; for example, the Pawnee have adopted it as an entheogen enhancer and in oneiromancy (dream divination), much as Calea zacatechichi is used in Mexico.[citation needed]
An indeterminate vervain is among the plants on the eighth panel of the New World Tapestry (Expedition to Cape Cod).[25]
In the Victorian language of flowers, verbena held the dual meaning of enchantment and sensibility.[26]
Species
[edit]The following species are accepted:[2] (See also Aloysia and Junellia for species formerly placed here.)
- Verbena alata Otto ex Sweet
- Verbena alejandrana (B.L.Turner) Christenh. & Byng
- Verbena × allenii Moldenke
- Verbena amoena Paxton
- Verbena andalgalensis Moldenke
- Verbena angustilobata (Moldenke) Christenh. & Byng
- Verbena araucana Phil.
- Verbena aristigera S.Moore
- Verbena atacamensis Reiche
- Verbena aurantiaca Speg.
- Verbena bajacalifornica Moldenke
- Verbena balansae Briq.
- Verbena barbata Graham
- Verbena berteroi (Meisn.) Schauer
- Verbena × bingenensis Moldenke
- Verbena bipinnatifida Schauer
- Verbena × blanchardii Moldenke
- Verbena bonariensis L. – purpletop vervain, clustertop vervain, tall verbena, pretty verbena, South American vervain
- Verbena brachyrhynchos (G.L.Nesom & Vorobik) Christenh. & Byng
- Verbena bracteata Cav. ex Lag. & Rodr. – prostrate vervain, large-bracted vervain
- Verbena brasiliensis Vell. – Brazilian verbena, Brazilian vervain
- Verbena cabrerae Moldenke
- Verbena californica Moldenke – California vervain, Red Hills vervain
- Verbena calinfera G.L.Nesom
- Verbena canadensis (L.) Britton
- Verbena canescens Kunth – gray vervain
- Verbena caniuensis Moldenke
- Verbena carnea Medik.
- Verbena carolina L.
- Verbena catharinae Moldenke
- Verbena cheitmaniana Moldenke
- Verbena chiricahensis (Umber) Moldenke
- Verbena × clemensiorum Moldenke
- Verbena cloverae Moldenke
- Verbena corymbosa Ruiz & Pav.
- Verbena cuneifolia Ruiz & Pav.
- Verbena dalloniana Quézel
- Verbena × deamii Moldenke
- Verbena delicatula Mart. & Zucc.
- Verbena delticola Small ex Perry
- Verbena demissa Moldenke
- Verbena dissecta Willd. ex Spreng.
- Verbena dusenii Moldenke
- Verbena ehrenbergiana Schauer
- Verbena elegans Kunth
- Verbena × engelmannii Moldenke
- Verbena ephedroides Cham.
- Verbena falcata G.L.Nesom
- Verbena filicaulis Schauer
- Verbena flava Gillies & Hook.
- Verbena glabrata Kunth
- Verbena gooddingii Briq.
- Verbena × goodmanii Moldenke
- Verbena goyazensis Moldenke
- Verbena gracilescens (Cham.) Herter
- Verbena gracilis Desf.
- Verbena grisea B.L.Rob. & Greenm.
- Verbena guaibensis (P.Peralta & V.Thode) Christenh. & Byng
- Verbena guaranitica (Tronc.) Moldenke
- Verbena gynobasis Wedd.
- Verbena halei Small – Texas vervain
- Verbena hassleriana Briq.
- Verbena hastata L. – swamp verbena, blue vervain
- Verbena hatschbachii Moldenke
- Verbena herteri Moldenke
- Verbena hirta Spreng.
- Verbena hispida Ruiz & Pav.
- Verbena humifusa Cham.
- Verbena × illicita Moldenke
- Verbena incompta P.W.Michael
- Verbena intermedia Gillies & Hook.
- Verbena jessicae Nesom & G.S.Hinton
- Verbena jordanensis Moldenke
- Verbena kuntzeana Moldenke
- Verbena laciniata (L.) Briq.
- Verbena landbeckii Phil.
- Verbena lasiostachys Link – western vervain
- Verbena lilacina Greene
- Verbena lilloana Moldenke
- Verbena lindbergii Moldenke
- Verbena lindmanii Briq.
- Verbena lipozygioides Walp.
- Verbena litoralis Kunth – seashore vervain
- Verbena livermorensis B.L.Turner & G.L.Nesom
- Verbena lobata Vell.
- Verbena macdougalii A.Heller
- Verbena macrosperma Speg.
- Verbena madrensis G.L.Nesom
- Verbena malmii Moldenke
- Verbena malpaisana (T.Van Devender & G.L.Nesom) Christenh. & Byng
- Verbena maritima Small
- Verbena marrubioides Cham.
- Verbena megapotamica Spreng.
- Verbena mendocina Phil.
- Verbena menthifolia Benth. – mint vervain
- Verbena microphylla Kunth
- Verbena moctezumae G.L.Nesom & T.Van Devender
- Verbena × moechina Moldenke
- Verbena montevidensis Spreng. – Uruguayan verbena
- Verbena moranii G.L.Nesom
- Verbena multiglandulosa Moldenke
- Verbena nana Moldenke
- Verbena neomexicana (A.Gray) Briq. – hillside vervain
- Verbena officinalis L. – common vervain, simpler's joy, holy herb, mosquito plant, wild hyssop, herb of the cross (type species)
- Verbena × ostenii Moldenke
- Verbena ovata Cham.
- Verbena paraguariensis Moldenke
- Verbena paranensis Moldenke
- Verbena parodii (Covas & Schnack) Moldenke
- Verbena paulensis Moldenke
- Verbena perennis Wooton – pinleaf vervain
- Verbena × perriana Moldenke
- Verbena × perturbata Moldenke
- Verbena peruviana (L.) Britton
- Verbena phlogiflora Cham.
- Verbena pinetorum Moldenke
- Verbena platensis Spreng.
- Verbena plicata Greene
- Verbena polyantha (Umber) Moldenke
- Verbena porrigens Phil.
- Verbena pumila Rydb.
- Verbena quadrangulata A.Heller
- Verbena racemosa Eggert
- Verbena radicata Moldenke
- Verbena recta Kunth
- Verbena rectiloba Moldenke
- Verbena regnelliana Moldenke
- Verbena reitzii Moldenke
- Verbena ribifolia Walp.
- Verbena rigida Spreng. – tuberous vervain
- Verbena rugosa Mill.
- Verbena × rydbergii Moldenke
- Verbena sagittalis Cham.
- Verbena santiaguensis (Covas & Schnack) Moldenke
- Verbena scabra Vahl – sandpaper vervain
- Verbena scabrella Sessé & Moc.
- Verbena scrobiculata Griseb.
- Verbena selloi Spreng.
- Verbena sessilis (Cham.) Kuntze
- Verbena simplex Lehm. – narrow-leaved vervain
- Verbena sphaerocarpa L.M.Perry
- Verbena stellarioides Cham.
- Verbena stricta Vent. – hoary vervain
- Verbena strigosa Cham.
- Verbena × stuprosa Moldenke
- Verbena subincana (Tronc.) Shinners
- Verbena sulphurea D.Don
- Verbena supina L.
- Verbena tampensis Nash
- Verbena tecticaulis Tronc.
- Verbena tenera Spreng.
- Verbena teucriifolia M.Martens & Galeotti
- Verbena thymoides Cham.
- Verbena tomophylla Briq.
- Verbena townsendii Svenson
- Verbena tumidula L.M.Perry
- Verbena turneri (G.L.Nesom) Christenh. & Byng
- Verbena tweedieana Niven ex Hook.
- Verbena urticifolia L. – white vervain
- Verbena × uruguayensis Moldenke
- Verbena valerianoides Kunth
- Verbena venturii Moldenke
- Verbena verecunda (Umber) Moldenke
- Verbena villifolia Hayek
- Verbena xutha Lehm. – gulf vervain
Gallery
[edit]-
Verbena obsession cascade pink
References
[edit]- ^ "Genus: Verbena L." Germplasm Resources Information Network. United States Department of Agriculture. 2004-01-29. Retrieved 2011-08-29.
- ^ a b "Verbena L." Plants of the World Online. Board of Trustees of the Royal Botanic Gardens, Kew. Retrieved 14 May 2021.
- ^ Western Garden Book. Sunset Books. 1995. pp. 606–07.
- ^ "Amazing Benefits of Verveine Tea: Is This a Miracle Tea?". Tea Reviews. Archived from the original on 2021-06-13. Retrieved 19 November 2022.
- ^ "Year of the Verbena". National Garden Bureau. 2021-10-19. Retrieved 2022-06-22.
- ^ S. M. Botta; S. Martinez & M. E. Mulguta de Romero (1995). "Novedades nomenclaturales en Verbenaceae" [Nomenclatural revisions in Verbenaceae]. Hickenia. 2: 127–128.
- ^ Yao-Wu Yuan & Richard G. Olmstead (2008). "A species-level phylogenetic study of the Verbena complex (Verbenaceae) indicates two independent intergeneric chloroplast transfers". Molecular Phylogenetics and Evolution. 48 (1): 23–33. Bibcode:2008MolPE..48...23Y. doi:10.1016/j.ympev.2008.04.004. PMID 18495498.
- ^ RHS A-Z encyclopedia of garden plants. United Kingdom: Dorling Kindersley. 2008. p. 1136. ISBN 978-1405332965.
- ^ "RHS Plant Selector Verbena 'Silver Anne' (G) AGM / RHS Gardening". Apps.rhs.org.uk. Archived from the original on 2012-12-24. Retrieved 2013-05-06.
- ^ "RHS Plant Selector Verbena 'Sissinghurst' (G) AGM / RHS Gardening". Apps.rhs.org.uk. Archived from the original on 2012-12-24. Retrieved 2013-05-06.
- ^ a b "Vervain". Drugs.com. 2009. Retrieved 7 March 2018.
- ^ D. S. Vohra (1 June 2004). Bach Flower Remedies: A Comprehensive Study. B. Jain Publishers. p. 3. ISBN 978-81-7021-271-3. Retrieved 2 September 2013.
- ^ "Flower remedies". Cancer Research UK. Retrieved 11 November 2016.
- ^ "Verbena". frontend. Archived from the original on 2023-06-08. Retrieved 2022-06-22.
- ^ Quattrocchi, Umberto (2000). CRC World Dictionary of Plant Names, Volume 4: R-Z. Taylor & Francis US. p. 2787. ISBN 978-0-8493-2678-3.
- ^ Gledhill, D. (2008). The Names of Plants (4th ed.). Cambridge University Press. p. 399. ISBN 978-0-521-86645-3.
- ^ Pliny the Elder. Naturalis Historia. pp. Liber XXV, Section LIX.
- ^ a b Pliny the Elder. Natural History. pp. Book 25, Section 59.
- ^ Watts, D. C. (2007). Dictionary of Plant Lore. Elsevier. p. 197. ISBN 9780080546025.
- ^ "Aconitum uncinatum (Appalachian Blue Monkshood, Blue Rocket, Devil's Helmet, Eastern Blue Monkshood, Eastern Monkshood, Monkshood, Southern Blue Monkshood, Wild Monkshood, Wolf's Bane) | North Carolina Extension Gardener Plant Toolbox". plants.ces.ncsu.edu. Retrieved 2022-06-22.
- ^ Valiente, Doreen (1994). An ABC of Witchcraft Past and Present. The Crowood Press. ISBN 9780719826917.
- ^ Pitois, Christian (1952). The History and Practice of Magic, Volume 2, Forge Press. p. 402.
- ^ Pitois, (1952) pp. 336, 339, 414
- ^ Gerina Dunwich (September 2019). Herbal Magick: A Guide to Herbal Enchantments, Folklore, and Divination. Weiser Books. ISBN 9781633411586.
- ^ "New World Tapestry". Mayflower 400. 17 February 2020. Archived from the original on 16 July 2021. Retrieved 16 July 2021.
- ^ "Language of Flowers - Flower Meanings, Flower Sentiments". www.languageofflowers.com. Retrieved 2016-11-26.
External links
[edit]
Media related to Verbena at Wikimedia Commons
Verbena
View on GrokipediaTaxonomy and naming
Etymology
The genus name Verbena derives from the Latin verbenaca, a diminutive form of verbēna, which referred to sacred boughs or twigs—typically of olive, myrtle, or laurel—carried by Roman priests during religious rituals and used to purify temples or as offerings.[7][8] In English, the common name "vervain" (applied especially to Verbena officinalis) traces back to Middle English verveyne, borrowed from Old French verveine, a variant of verbena that emerged in medieval herbal traditions, where the plant was valued for its purported healing properties in poultices and infusions.[9][10] Cultural variations in common names reflect regional folklore; in Spanish-speaking regions, "verbena" retains the Latin root and evokes both the plant and festive gatherings, while in German, "Eisenkraut" (literally "iron herb") stems from beliefs in its magical resistance to iron tools and protective role against witchcraft in Germanic traditions.[11][12] The Linnaean system introduced in the 18th century standardized "Verbena" as the scientific genus name, with Carl Linnaeus formally describing it in his 1753 Species Plantarum, shifting from earlier vernacular and pre-Linnaean classifications to a binomial nomenclature that preserved the ancient Latin term for this group of verbenaceous plants.[14]Classification
The genus Verbena belongs to the family Verbenaceae in the order Lamiales, with 69 accepted species (comprising around 350 taxa including infraspecific categories, as of 2024), of annual and perennial herbaceous or semi-woody plants primarily native to the Americas. Species counts vary by taxonomic treatment; recent estimates recognize the strict sense excluding Glandularia.[3] Carl Linnaeus first established the genus Verbena in 1753, including 14 species based on morphological similarities such as leaf arrangement and inflorescence structure.[15] Subsequent revisions in the 19th and 20th centuries refined its circumscription; for instance, George Bentham in 1839 and later taxonomists like Hilda Umber in 1979 separated several North American species into the distinct genus Glandularia due to differences in seed morphology, chromosome number, and fruit structure.[16] These changes addressed earlier lumping of taxa that shared superficial traits but diverged in reproductive and genetic features, reducing older estimates of Verbena species from around 150 to a more precise count while expanding recognition of related genera.[17] Molecular phylogenetic analyses using DNA sequences, such as those from chloroplast genes conducted in the early 21st century, have demonstrated that Verbena is monophyletic and forms a sister group to Glandularia within the Verbena complex.[18] These studies, including comprehensive sampling by Yuan and Olmstead (2008), support infrageneric divisions into series (e.g., Verbena ser. Verbena and ser. Pseudoverbena), based on cladistic patterns of morphological and genetic variation, though broader Verbenaceae phylogeny reveals polyphyly in neighboring genera like Junellia.[19] Key synapomorphies defining Verbena include opposite or whorled leaves and elongate, spike-like inflorescences, which contrast with the rounded, capitulate heads typical of related genera such as Lantana.[18] These traits, combined with a didynamous corolla and nutlet fruits, provide diagnostic characters for systematic placement within Verbenaceae.[19]Description
Morphology
Verbena species exhibit a range of growth habits, primarily as annual or perennial herbs or subshrubs, typically reaching heights of 10 to 300 cm.[2] The stems are characteristically quadrangular in cross-section, erect to ascending or sometimes procumbent, and often bear short, stiff hairs or glandular pubescence.[20] Leaves are arranged oppositely and decussately, occasionally in whorls of three, and are simple with blades that are entire to pinnatisect, featuring toothed to serrate margins; they measure 1 to 10 cm in length, are lanceolate to ovate in shape, and commonly possess stalked glandular hairs.[20][2] The inflorescence consists of terminal spikes or racemes, sometimes forming panicle-like structures, with bracts subtending the flowers; the axis elongates significantly in fruit.[20] Flowers are small, sessile or subsessile, bisexual, and zygomorphic, with five merous parts including a 4- to 5-lobed calyx that becomes accrescent in fruit and a tubular to infundibuliform corolla.[20] Corolla lobes spread widely, displaying colors such as purple, blue, pink, or white, while the four didynamous stamens are included or exserted, and the single style ends in an unlobed or two-lobed stigma.[20] The superior ovary is four-locular.[20] The fruit is a schizocarp that separates into four one-seeded nutlets at maturity, often remaining enveloped by the persistent calyx; the nutlets may have a scabridulous or granulate inner face and are sometimes winged or appendaged.[20][21] These nutlets produce mucilage when wetted, becoming moist and sticky, which facilitates adhesion to animal fur or soil surfaces for dispersal.[22]Distribution and habitat
In the strict sense, following recent taxonomic revisions, the genus Verbena comprises approximately 70 species.[3] It is predominantly native to the Americas, with the majority occurring across tropical, subtropical, and temperate regions from southern South America northward to Canada.[19] The center of diversity for Verbena sensu stricto lies in North and Central America (particularly Mexico and the United States).[19] The family Verbenaceae originated and initially diversified in wet forest habitats of South America.[23] In North America, species are widespread east of the Rocky Mountains and in the Southwest, often in open or semi-open landscapes.[19] One notable exception within the genus is Verbena officinalis, which is native to Eurasia and North Africa but has been naturalized worldwide, including in the Americas, Europe, Asia, Africa, and Australia, likely since ancient times due to its historical use in traditional medicine and rituals.[24] This species has established populations in temperate to subtropical zones globally, often escaping cultivation to occupy waste areas and roadsides.[25] Verbena species generally prefer open habitats such as grasslands, roadsides, disturbed soils, and rocky slopes, where they can thrive in full sun with minimal competition.[26] Many exhibit strong tolerance to drought and nutrient-poor soils, enabling persistence in arid or semi-arid conditions once established.[27] Altitudinal distribution varies widely, from sea level in coastal and lowland areas to elevations exceeding 3,000 meters in some Andean species, such as Verbena demissa in montane Ecuador.[19] Several Verbena species have been introduced outside their native ranges and can become invasive. For instance, V. bonariensis, native to South America, has naturalized widely and is considered an environmental weed in Australia and New Zealand, where it invades grasslands, roadsides, and disturbed sites, potentially displacing native vegetation.[28][29]Ecology
Reproduction
Verbena species typically exhibit a flowering phenology extending from spring through autumn, with specific blooming periods varying by species and geographic location. For instance, Verbena urticifolia flowers from May to November, while Verbena hastata blooms from June to October.[30][31] The flowers feature spatial separation of anthers and stigmas, favoring cross-pollination.[32] Pollination in Verbena is predominantly entomophilous, mediated by a range of insects including bumblebees, digger bees, small carpenter bees, butterflies, wasps, and flies.[33][34] These species are facultatively xenogamous, capable of both cross- and self-pollination, though outcrossing is preferred due to low self-pollination rates, often supplemented by incidental pollination from thrips.[32] Seed production is prolific, enabling effective self-sowing in suitable habitats, with established plants generating numerous seeds per individual.[35] Dispersal mechanisms include anemochory, where lightweight nutlets are carried by wind, and zoochory, facilitated by adhesion to animal fur via sticky surfaces or mucilage on the seeds.[36] Asexual reproduction is uncommon in Verbena but occurs in some perennial species through the rooting of prostrate stems in contact with moist soil.[37]Interactions with other organisms
Verbena species experience herbivory from various organisms, including deer and insects, though they demonstrate notable resistance due to physical and chemical defenses. Deer typically avoid Verbena owing to its coarse foliage and bitter taste, rating it as "seldom severely damaged" in assessments of browse preference, but may graze it under food scarcity. Insects such as generalist herbivores occasionally feed on leaves and stems, yet the genus's production of iridoid glycosides, including compounds like verbenalin and hastatoside in Verbena officinalis, acts as a feeding deterrent by imparting a bitter flavor and potentially releasing toxic aglycones upon hydrolysis.[38][39][40][41][42] Additionally, volatile oils present in Verbena foliage contribute to deterrence against browsers, functioning as repellents that disrupt insect feeding and oviposition behaviors. These secondary metabolites, common across the Verbenaceae family, enhance overall plant resilience in natural settings without broadly harming beneficial insects.[43][44] Mutualistic interactions in Verbena primarily involve pollination and seed dispersal, with limited evidence of other symbioses like nitrogen fixation. The nectar-rich flowers attract diverse pollinators, including bees, butterflies, and occasionally hummingbirds, supporting reproductive success through these partnerships. Seeds, often small and sticky, facilitate dispersal by birds that consume them—such as songbirds like cardinals and sparrows—and by ants in some habitats, where adhesion promotes transport away from parent plants.[45][46][47] No nitrogen-fixing associations occur, as the genus lacks symbiotic bacteria, but it forms arbuscular mycorrhizal associations that enhance nutrient acquisition.[48] Some species, like V. hastata, serve as larval hosts for butterflies such as the verbena moth (Crambodes erutalis).[37] Several Verbena species exhibit invasiveness in non-native regions, leading to ecological disruptions through competition with indigenous plants. For instance, Verbena bonariensis has naturalized across parts of North America and Australia, forming dense stands that outcompete native forbs for light, water, and nutrients in grasslands and disturbed areas, thereby reducing biodiversity. Similarly, Verbena brasiliensis invades floodplains and pastures in Africa and Asia, displacing local flora and altering community structure by rapid vegetative spread and prolific seed production. These impacts highlight the genus's potential to shift ecosystem dynamics in introduced habitats.[49] In native ecosystems, Verbena often serves as a pioneer species during ecological succession, colonizing disturbed soils and aiding community development. Species like Verbena hastata and Verbena lasiostachys thrive in open, eroded sites, where their fibrous root systems stabilize soil against erosion and facilitate nutrient cycling for subsequent colonizers. This role is particularly evident in prairies and riparian zones, where Verbena hastens recovery post-disturbance by providing early ground cover and habitat structure.[50][51][34]Cultivation
Growing conditions
Verbena plants thrive in well-drained soils with a slightly acidic to neutral pH ranging from 5.8 to 7.2, as heavy clay or compacted soils can lead to root rot while sandy soils benefit from organic amendments to retain moisture.[52][53] Full sun exposure of at least 6-8 hours per day is essential for prolific flowering, though they tolerate light shade but may produce fewer blooms in partial conditions.[52][54] Hardiness varies by species and cultivar, with most perennial verbenas thriving in USDA zones 4-11; many cultivated ornamental varieties are well-suited to zones 7-11 as perennials, where they exhibit good heat tolerance up to 35°C (95°F) and become drought-resistant once established, requiring minimal supplemental water in average conditions.[55][56][52] In cooler zones, they are typically grown as annuals, mirroring the sunny, dry habitats of their native American origins.[57] Moderate watering is recommended to keep soil evenly moist without sogginess, particularly during establishment, while overwatering should be avoided to prevent disease.[52] Fertilization should be moderate using a balanced, slow-release formula applied monthly during the growing season to support blooming without excess nitrogen, which can cause leggy growth and reduced flowers.[58][59] Hybrid verbenas, such as V. × hybrida varieties like those from the Superbena or Lanai series, have been selectively bred for more compact habits and enhanced performance in containers, maintaining dense growth in full sun with similar low-maintenance needs.[55][54]Propagation and maintenance
Verbena plants can be propagated through seeds, which are typically sown indoors in early spring, such as February to March, at temperatures between 18-24°C, with germination occurring in 14-28 days.[60] Seeds should be surface-sown on moist, well-draining soil without covering, as they require light for germination, and for some species, a period of cold stratification or light scarification may improve success rates.[61] Vegetative propagation is often preferred for maintaining specific traits, using stem cuttings taken in late spring or summer from non-flowering shoots about 3 inches long, stripped of lower leaves and rooted in a moist, gritty medium under high humidity, such as covered with a plastic bag, typically rooting in 2-6 weeks.[61][60] For perennial species, division can be performed in spring by separating established clumps at the roots and replanting immediately in suitable conditions.[60] Ongoing maintenance involves regular deadheading of spent flowers to encourage continuous blooming throughout the season, using clean shears to trim faded blooms back to a leaf node or healthy stem.[53] Pruning is recommended after the main flowering period or in spring for perennials, cutting back stems to promote bushier growth and new shoots, while trailing varieties may need trimming several times during the growing season.[60] In colder climates, overwintering tender varieties requires protection such as mulching roots with organic material in autumn or bringing potted plants indoors, whereas hardy types like Verbena bonariensis need only minimal mulching to insulate against frost; hardier species such as V. officinalis may require little to no protection in zones 3-8.[60] Popular cultivars for propagation include trailing types such as 'Superbena', which root readily from cuttings in 3-3.5 weeks and are ideal for hanging baskets due to their vigorous, cascading habit.[55][62]Pests and diseases
Common pests
Cultivated Verbena plants are susceptible to several insect and animal pests that can damage foliage and overall vigor. Aphids and whiteflies are common sap-sucking insects that feed on plant juices, leading to curled and distorted leaves, yellowing, and reduced growth.[57] These pests can be managed non-chemically by introducing natural predators such as ladybugs, which consume large numbers of aphids and whiteflies, or by applying insecticidal soaps that disrupt their cell membranes without harming beneficial insects.[63][64] Spider mites, particularly the two-spotted spider mite, thrive in dry, hot conditions and cause stippling or fine white dots on leaves due to their piercing-sucking mouthparts, potentially leading to webbing and leaf drop if unchecked.[57][65] Prevention and control involve increasing humidity around plants through regular misting or grouping with other moisture-retaining species, as higher humidity discourages mite reproduction.[66] Japanese beetles and various caterpillars act as defoliators, skeletonizing leaves by chewing on foliage and flowers, which can weaken plants and reduce blooming.[67][68] Effective non-chemical strategies include hand-picking beetles and caterpillars early in the morning when they are less active, followed by dropping them into soapy water, or applying neem oil sprays that interfere with their feeding and molting processes.[69] Deer may occasionally browse on Verbena in garden settings, especially when food is scarce, nibbling on tender shoots and flowers despite the plant's general resistance due to its fuzzy leaves and scent.[70][71] Repellents such as commercial deer sprays containing putrescent egg solids or fencing around plantings provide reliable protection without chemical residues.[72]Diseases and disorders
Verbena plants in cultivation are susceptible to several fungal pathogens that thrive in specific environmental conditions. Powdery mildew, caused by the fungus Golovinomyces ambrosiae, commonly affects plants in humid environments, manifesting as white powdery growth on leaves and petals.[73][74] This disease can be managed through applications of appropriate fungicides and by ensuring adequate air circulation to reduce humidity.[73] Botrytis blight, also known as gray mold and caused by Botrytis cinerea, produces tan spots on leaves, crown rot in established plants, and gray fuzzy spores on infected tissue, favored by cool, wet conditions.[73] Management includes improving air circulation, reducing overhead watering, and applying fungicides preventively. Root rot, primarily induced by Pythium species, results from overwatering and poorly drained soils, leading to dark brown roots, wilting, and yellowing foliage.[73] Prevention involves using well-draining media and avoiding excessive moisture, with fungicides applied as needed for control.[73] Bacterial and viral infections also pose risks to cultivated Verbena. Bacterial leaf spot, caused by Pseudomonas or Xanthomonas species, produces water-soaked lesions on leaves that may turn necrotic, often spreading in wet conditions.[75] Copper-based bactericides effectively suppress the pathogen, though sanitation practices are essential to limit spread.[75] Viral diseases, such as those caused by alfalfa mosaic virus or potyviruses, result in mosaic patterns, mottling, and leaf distortion, with transmission occurring mechanically via contaminated tools or pruning equipment during propagation.[76] Selecting resistant varieties is recommended, as no curative treatments exist, and infected plants should be removed to prevent further dissemination.[76] Abiotic disorders further compromise Verbena health in suboptimal growing conditions. Iron chlorosis, prevalent in alkaline soils where iron availability is reduced, causes interveinal yellowing of young leaves while veins remain green.[77] This can be corrected by applying iron chelates to the soil or as foliar sprays to enhance uptake.[77] Sunscald occurs in extreme heat, leading to bleached or scorched foliage on exposed plants, particularly when combined with drought stress. Providing partial shade during peak temperatures helps mitigate this non-pathogenic damage. Integrated management strategies are crucial for preventing disease outbreaks in Verbena cultivation. Crop rotation disrupts pathogen life cycles by avoiding repeated planting in the same soil, while rigorous sanitation—such as removing infected debris and sterilizing tools—reduces inoculum sources.[78] These practices, combined with monitoring for early symptoms, support overall plant health without relying solely on chemical interventions.[79]Uses
Ornamental applications
Verbena species and hybrids are widely valued in horticulture for their versatility as border plants, groundcovers, and container fillers, owing to their cascading growth habits and extended blooming periods that can last up to three months or more under optimal conditions.[70] These plants thrive in full sun and well-drained soils, making them ideal for edging pathways, filling gaps in rock gardens, or trailing over the sides of hanging baskets and window boxes, where their low-growing forms—often reaching just 6 to 12 inches in height—provide a spilling effect that enhances visual interest without overwhelming surrounding features.[57][80] Among the most popular ornamental varieties are the V. × hybrida series, renowned for their vivid clusters of flowers in shades of red, pink, purple, and white, which bloom continuously from late spring through early fall when deadheaded regularly.[70] Trailing cultivars such as Verbena 'Homestead Purple' (V. canadensis 'Homestead Purple') exemplify this appeal, offering bright purple blooms over a long season and rapid coverage in sunny, dry sites, often selected for their vigorous spreading habit in mixed plantings.[81][82] In garden design, Verbena pairs effectively with salvias and petunias to create pollinator-friendly landscapes, attracting butterflies and bees while adding complementary colors and textures to beds or containers.[83] Several hybrids have earned recognition through All-America Selections awards, such as EnduraScape™ Pink Bicolor and Verbena Imagination, for their heat and drought tolerance, contributing to their growing use in sustainable, low-water gardening trends like xeriscapes.[84][85]Medicinal and other uses
Verbena officinalis, commonly known as common vervain, has been utilized in traditional herbal medicine for its diuretic properties, aiding in the promotion of urine production and supporting kidney and bladder health.[86] It is also employed as a sedative to alleviate anxiety, insomnia, and nervous tension, with studies demonstrating anxiolytic and anticonvulsant effects in animal models.[87] Additionally, the plant exhibits anti-inflammatory activity, which has been attributed to its polyphenolic compounds and is used to treat conditions such as respiratory inflammation, skin irritations, and rheumatic disorders.[88] Extracts of V. officinalis promote wound healing, particularly for difficult-to-heal lesions, due to their softening, antibacterial, and tissue-regenerating effects, with verbenalin identified as a key iridoid glycoside contributing to these benefits.[89] In industrial contexts, extracts of Verbena officinalis are integrated into cosmetics for their astringent properties, which help tone the skin, reduce puffiness, and soothe irritations, making them suitable for formulations targeting oily or inflamed skin.[90] Certain Verbena species, such as Verbena hybrida, show potential in phytoremediation, effectively accumulating heavy metals like chromium from contaminated tannery wastewater and soil, offering an eco-friendly approach to environmental cleanup.[91] Despite these uses, V. officinalis can exhibit toxicity in large doses, potentially leading to gastrointestinal upset or neurological effects due to compound accumulation.[92] It is contraindicated during pregnancy, particularly the first and second trimesters, as studies indicate prenatal developmental toxicity and risks to reproductive performance in animal models.[93]Cultural significance
Symbolism and folklore
In ancient Roman tradition, Verbena species, particularly Verbena officinalis known as vervain, held sacred status as verbenae or sagmina, referring to ritual branches plucked from sacred enclosures like the Capitoline Hill. These were carried by priests such as the Flamines and Fetiales during sacrifices, treaty negotiations, and purification rites to invoke divine protection against evil influences and ensure ritual sanctity. The plant symbolized purity and warded off harm, with beliefs that its leaves could protect wearers from injury in battle or misfortune.[94] Druidic lore in Celtic cultures revered vervain as one of the sacred herbs, harvested at specific lunar phases for use in protection charms, love divinations, and purification ceremonies to repel malevolent spirits.[95] In broader European folklore, it was esteemed as a vulnerary herb for staunching wounds and promoting healing, often incorporated into amulets or baths for both physical and spiritual safeguarding. Later traditions extended this to warding off vampires and witchcraft, reflecting its enduring role as a charm against supernatural threats.[96][97] In modern esotericism, vervain maintains associations with the planet Venus, symbolizing love, harmony, and attraction, and is used in rituals to enhance romantic intentions or create protective talismans against negative energies.[98] Across cultures, including Native American practices, species like blue vervain (Verbena hastata) have been used medicinally, with roots and leaves applied as poultices or infusions for wound care.[99]Representation in culture
Verbena, particularly the species known as vervain (Verbena officinalis), has appeared in literature as a symbol of enchantment and protection, drawing from its historical associations with magic and healing. In Victorian-era floriography, the language of flowers, verbena represented enchantment, often used to convey sentiments of fascination or spiritual allure in poetry and correspondence.[100] In Roman mythology, vervain was associated with ancient names such as "Mercury's blood." Christian traditions further elevated vervain's status, associating it with the Passion of Christ; medieval legends claimed it was used to staunch the wounds of Jesus on the cross, earning it the title "herb of the cross" and imbuing it with themes of sacrifice and redemption.[101][102] In art, verbena features prominently in botanical illustrations, notably those by Pierre-Joseph Redouté, the renowned 19th-century French painter known as the "Raphael of flowers." Redouté depicted species like lemon verbena (Aloysia citrodora, formerly Verbena triphylla) and Verbena globiflora in detailed watercolors for works such as Choix des plus belles fleurs (1827–1833), capturing their delicate blooms and leaves to highlight their ornamental elegance. In modern media, verbena appears as a romantic motif in gardening shows and films, symbolizing carefree charm and enduring affection; for instance, its airy purple spikes evoke whimsical, love-filled garden scenes in programs like those on Proven Winners, where it adorns borders and baskets to create intimate, nostalgic atmospheres.[103] Contemporary representations extend to branding and cultural events, where verbena's fresh, citrusy scent inspires perfumes like L'Occitane's Verbena line, launched in 1990 and evoking Provençal summers with notes of lemon verbena for a lively, invigorating profile.[104] Additionally, Spanish verbenas—outdoor festivals blending religious devotion, music, and dance—draw their name from the plant, originally honoring patron saints with garlands of verbena flowers and celebrating wildflower heritage in communal gatherings.Species
Diversity and evolution
The genus Verbena encompasses approximately 70 accepted species of annual and perennial herbs or subshrubs in the strict sense, though broader historical classifications recognize up to 250 species (many now placed in related genera such as Glandularia), with the highest species richness concentrated in South America, particularly in regions from Brazil to Peru and Argentina.[3][19] This diversity reflects the genus's adaptation to a range of habitats, including open grasslands, rocky slopes, and disturbed areas, though significant endemism occurs in the Andean highlands of Bolivia, Peru, Ecuador, and Colombia, as well as in the arid and semi-arid landscapes of the southwestern United States, such as Arizona, New Mexico, and Texas.[3] Evolutionary studies indicate that Verbena originated in South America where the Verbenaceae family likely arose amid the diversification of tropical ecosystems.[105] This period marked the beginning of an adaptive radiation in the New World, enabling the genus to colonize diverse ecological niches across the Americas through mechanisms like habitat specialization and dispersal via wind-dispersed seeds, with subsequent migrations leading to limited presence in Africa and Eurasia.[18] Genetic analyses reveal notable diversity within Verbena, including polyploidy prevalent in South American lineages—contrasting with predominantly diploid North American taxa—where chromosome numbers often exceed the base of , contributing to reproductive isolation and morphological variation.[106] Hybridization barriers have been explored through chloroplast DNA sequencing, which has uncovered intergeneric transfers (e.g., between Verbena and Glandularia), highlighting reticulate evolution and incomplete lineage sorting as key processes in the genus's diversification.[107] Several Verbena species face conservation threats primarily from habitat loss due to agriculture, urbanization, and invasive species, with notable examples including V. californica (Red Hills vervain) in California and V. orcuttiana in Baja California, Mexico, both restricted to specialized serpentine or coastal habitats now fragmented by development.[108] These pressures underscore the need for targeted protection to preserve the genus's evolutionary legacy amid ongoing environmental changes.[109]Notable species
Verbena officinalis, commonly known as common vervain or European vervain, is a widespread perennial herb native to Europe, Asia, and parts of Africa, with introductions in the Americas and Oceania. It grows to a height of 30–140 cm, featuring square stems, opposite leaves, and spikes of small lavender flowers. This species thrives in temperate biomes, often in disturbed habitats like roadsides and waste areas, and has been utilized historically as a medicinal plant for treating ailments such as headaches and anxiety, as well as for environmental and social purposes.[110][111] Verbena bonariensis, or tall verbena, is a South American native perennial that forms wiry stems rising to 2–4 feet from a basal clump, topped with clusters of small rose-violet to lavender flowers from June to frost. Valued for its airy habit and pollinator attraction, it has naturalized in parts of the United States and can self-seed aggressively, leading to invasive tendencies in some regions like the southeastern U.S. where it outcompetes native vegetation.[5] Verbena hastata, known as blue vervain or swamp verbena, is a North American native perennial reaching 2–6 feet tall, with square stems bearing opposite, lance-shaped leaves and dense spikes of blue-violet flowers blooming from July to September. It inhabits wet meadows, stream banks, and marshes across eastern North America, from Canada to Mexico, supporting wetland ecosystems and attracting butterflies and bees.[4][37] The Verbena × hybrida group encompasses garden verbenas, hybrid cultivars developed primarily from South American species such as V. teucrioides, V. incisa, V. peruviana, and V. phlogiflora starting in the mid-19th century. These compact, trailing or mounded plants, typically 6–18 inches tall, produce prolific clusters of flowers in various colors and are bred for ornamental use in gardens and containers, offering extended bloom periods and disease resistance.[112] Among endangered species, Verbena simplex (narrowleaf vervain), a U.S. native perennial with slender stems up to 3 feet tall and pale lavender flower spikes, is listed as endangered in states like Massachusetts due to habitat loss from development and agriculture. Restricted to dry prairies, open woodlands, and limestone glades in the Midwest and eastern U.S., it features narrow leaves and blooms in summer, highlighting conservation needs for rare Verbena taxa.[113]References
- https://en.wiktionary.org/wiki/vervain
- https://en.[wiktionary](/page/Wiktionary).org/wiki/Eisenkraut