Lamium galeobdolon

Yellow archangel
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Lamiales
Family:	Lamiaceae
Genus:	Lamium
Species:	L. galeobdolon
Binomial name
Lamium galeobdolon (L.) L.
Synonyms
Lamium galeobdolon Galeopsis galeobdolon L. (basionym) Neckeria galeobdolon (L.) Forsyth f. Galeobdolon galeobdolon (L.) H.Karst., nom. illeg. Lamiastrum galeobdolon (L.) Ehrend. & Polatschek Leonurus galeobdolon (L.) Scop. Pollichia galeobdolon (L.) Schrank L. g. subsp. argentatum Galeobdolon argentatum Smejkal (basionym) Lamiastrum argentatum (Smejkal) Soják Lamiastrum galeobdolon subsp. argentatum (Smejkal) Stace Lamium argentatum (Smejkal) Henker ex G.H.Loos Lamium galeobdolon f. argentatum (Smejkal) Mennema L. g. subsp. flavidum Lamium flavidum F.Herm. (basionym) Galeobdolon flavidum (F.Herm.) Holub Lamiastrum flavidum (F.Herm.) Ehrend. Lamium pallidum F.Herm. L. g. subsp. galeobdolon Cardiaca sylvatica Lam. Galeobdolon luteum Huds. Galeobdolon luteum var. albomaculatum Salter ex J.Dix Galeobdolon luteum var. regulare Gray Galeobdolon umbrosum Wibel Galeobdolon vulgare (Pers.) Pers. Galeopsis lutea Gilib. Lamium cavernianum Losa Lamium luteum Krock. Lamium vulgare (Pers.) Fritsch Pollichia longicaulis Krock. Pollichia vulgaris Pers. L. g. subsp. montanum Pollichia montana Pers. (basionym) Galeobdolon endtmannii (G.H.Loos) Holub Galeobdolon luteum proles montanum (Pers.) Rouy Galeobdolon luteum var. florentinum Silva Tar. Galeobdolon luteum var. montanum (Pers.) Gray Galeobdolon luteum subsp. montanum (Pers.) R.R.Mill Galeobdolon montanum (Pers.) Rchb. Galeopsis galeobdolon var. montanum (Pers.) Lej. Lamiastrum galeobdolon subsp. montanum (Pers.) Ehrend. & Polatschek Lamiastrum montanum (Pers.) Ehrend. Lamium endtmannii G.H.Loos Lamium galeobdolon var. montanum (Pers.) Pers. Lamium galeobdolon f. montanum (Pers.) Bolzon Lamium montanum subsp. endtmannii (G.H.Loos) G.H.Loos Lamium montanum var. florentinum (Silva Tar.) Buttler & Schippm. Lamium montanum (Pers.) Hoffm. ex Kabath

Lamium galeobdolon, the yellow archangel, is a species of flowering plant in the mint family Lamiaceae. It is native to Europe and western Asia but it is widely introduced in North America and elsewhere. It is the only species in the genus Lamium with yellow flowers. Another common name for this species is golden dead-nettle. In New Zealand, it is called the aluminium plant or artillery plant. The common names archangel and dead-nettle have been in use for hundreds of years, dating back to at least the 16th century.

Lamium galeobdolon comprises four closely-related subspecies, one of which, the variegated yellow archangel (Lamium galeobdolon subsp. argentatum) from central Europe, is widely present as an invasive subspecies in several European countries outside of its native range and also in New Zealand, Canada, and the United States.

Description

Lamium galeobdolon sensu lato is a perennial plant with square stems growing from 15 to 60 cm (6 to 24 in) tall. The paired opposite leaves are stalked with toothed margins. The leaves are 4 to 7 cm (2 to 3 in) long, and may or may not have silvery markings (an important character used to distinguish subspecies). The inflorescence is a pseudowhorl of 4–16 flowers (called a verticillaster) clustered around the axil of a leaf-pair. Each flower has bilateral (zygomorphic) symmetry. The calyx is five-lobed and the corolla is yellow with a prominent hood. The flower's lower lip has three lobes with the central lobe often streaked with orange. There are two short stamens and two long ones. Flowering is in late spring to early summer, typically May–June in Britain. A pair of fused carpels give rise to a four-chambered schizocarp.

Identification

Subspecies of Lamium galeobdolon sensu lato are the only taxa in the genus with yellow flowers. All of the subspecies are stoloniferous except subsp. flavidum. The invasive subsp. argentatum is readily distinguished by its silvery white variegated leaves. Lamium galeobdolon subsp. galeobdolon and subsp. montanum can be difficult to distinguish but assuming the plant in question is in full flower, subsp. galeobdolon has a maximum of six (rarely seven) flowers per verticillaster while subsp. montanum averages ten flowers per verticillaster. The hairiness of the lower stems is also a useful clue, with subsp. galeobdolon having hairs on the four ridges only, while subsp. montanum is more uniformly hairy across the faces as well as the ridges.

Lamium galeobdolon subsp. flavidum is notable for the lack of stolons, frequently branched fertile stems with upright flowering lateral shoots, and 10–16 small flowers per verticillaster. An identification key for the remaining taxa follows Rosenbaumová, Plačková & Suda (2004), Streeter et al. (2009), and Walker & Mountford (2020):

Identification Key Lamium galeobdolon sensu lato in central and western Europe
1a. Leaves and bracts (except for the uppermost) with a distinct silvery pattern (two flexuous bands along the midrib) persisting year around, bracts with wide apical teeth; flowers relatively large	Lamium galeobdolon subsp. argentatum
1b. Leaves generally without silvery pattern (if present, then never forming two continuous flexuous bands along the midrib); apical teeth of bracts mostly narrower; flowers relatively smaller	2
2a. The uppermost bracts ovate, generally less than twice as long as wide, with more-or-less equally distributed marginal teeth, apical tooth short and obtuse, lowermost bracts narrower or only slightly wider than the uppermost ones; the uppermost leaves shortly petiolate; verticillasters few-flowered (maximum number of flowers usually 4–7); hairs on the lower half of the stem almost entirely confined to the angles	Lamium galeobdolon subsp. galeobdolon
2b. The uppermost bracts lanceolate, more than twice as long as wide, marginal teeth towards the apex more separated, apical tooth long and acute; lowermost bracts mostly distinctly wider than the uppermost ones; the uppermost leaves with longer petioles; maximum number of flowers in verticillasters generally (6–)10–14; hairs on the lower half of the stem more or less equal distribution on the faces and the angles	Lamium galeobdolon subsp. montanum

Other taxa in the group can have marked leaves but those of subsp. argentatum are distinctive:

Permanent silvery pattern on leaf lamina in L. argentatum always forms two flexuous bands along the midrib, mostly not interrupted by the lateral veins. They are distinctive particularly from the late autumn to the early spring when a contrast brownish-maroon colouring develops on the abaxial side and extends to the midrib-zone of the upper leaf side. Although variegated leaves were frequently observed in all other taxa in our study, they never matched the description above. Silvery-grey markings constituted merely separate splashes or flecks (small-sized in L. montanum and L. flavidum, larger in L. galeobdolon), not merged into continuous bands. Thus, a combination of two distinct silvery flexuous bands together with a strong chocolate-maroon zone along the midrib in winter can be regarded as specific L. argentatum markers.

— Rosenbaumová, Plačková & Suda (2004)

In particular, a cultivar (of subsp. flavidum - ? - ) known as 'Herman's Pride' has silver-spotted leaves, but even though the markings are unlike those of subsp. argentatum, the two taxa are sometimes confused.

The base chromosome number of taxa in genus Lamium is x = 9. Lamium galeobdolon subsp. flavidum and subsp. galeobdolon are diploid (2n = 2x = 18) while subsp. argentatum and subsp. montanum are tetraploid (2n = 4x = 36). A few triploid individuals have been reported. The triploids are thought to be natural hybrids between diploid subsp. galeobdolon and tetraploid subsp. argentatum.

Taxonomy

Lamium galeobdolon was first described as Galeopsis galeobdolon by the Swedish botanist Carl Linnaeus in 1753. Linnaeus himself transferred it to the genus Lamium in 1759, and therefore the correct name in Lamium is Lamium galeobdolon (L.) L.

Most authorities with a global scope accept four closely-related taxa, either as subspecies of Lamium galeobdolon, or as full species in genus Lamium, in which case the group of taxa is referred to as Lamium subgen. Galeobdolon. Some authorities segregate the species into other genera (either Galeobdolon or Lamiastrum) but genus Lamium remains monophyletic in any case.

Subspecies	Species complexes
Lamium galeobdolon subsp. argentatum (Smejkal) J.Duvign.	Lamium argentatum (Smejkal) Henker ex G.H.Loos	Galeobdolon argentatum Smejkal	Lamiastrum argentatum (Smejkal) Soják
Lamium galeobdolon subsp. flavidum (F.Herm.) Á.Löve & D.Löve	Lamium flavidum F.Herm.	Galeobdolon flavidum (F.Herm.) Holub	Lamiastrum flavidum (F.Herm.) Ehrend.
Lamium galeobdolon subsp. galeobdolon	Lamium galeobdolon (L.) L. sensu stricto	Galeobdolon galeobdolon (L.) H.Karst.	Lamiastrum galeobdolon (L.) Ehrend. & Polatschek
Lamium galeobdolon subsp. montanum (Pers.) Hayek Basionym: Pollichia montana Pers.	Lamium montanum (Pers.) Hoffm. ex Kabath	Galeobdolon montanum (Pers.) Rchb.	Lamiastrum montanum (Pers.) Ehrend.

Lamium galeobdolon subsp. argentatum was first described as Galeobdolon argentatum by the Czech botanist Miroslav Smejkal in 1975. In a comprehensive and influential treatment of genus Lamium published in 1989, the Dutch botanist Jacob Mennema reduced the species to forma and placed it in synonymy with Lamium galeobdolon subsp. galeobdolon. Hence subsp. argentatum was neglected for a long time and its invasion history is poorly documented.

There are a number of closely related taxa that hybridise with L. galeobdolon and in some cases are not unequivocally accepted as distinct species but considered subspecies or varieties by many authors. Most well known among these is variegated yellow archangel (subsp. argentatum), whose leaves often have variegation, showing as silver patches arranged as a wide semicircle. This, and in particular its large-flowered and even stronger-marked cultivar 'Variegatum', is the taxon most often met with as a garden escapee.

Distribution

Lamium galeobdolon sensu lato is native across Europe, western Asia, and portions of the Middle East. It ranges from Ireland and Spain east across Europe as far as the West Siberian Plain and southeast to Iran. Lamium galeobdolon subsp. flavidum is native to the Alps, northern Italy, and the mountain ranges northeast of the Adriatic Sea. L. galeobdolon subsp. galeobdolon and subsp. montanum have broadly similar distributions, but subsp. montanum extends further west and south than subsp. galeobdolon, which has a more northeast distribution. Within Britain and Ireland, subsp. montanum is a widespread native across England and Wales and locally in southern Scotland and eastern Ireland, while nominate subsp. galeobdolon is restricted to a small area in Lincolnshire in eastern England.

Lamium galeobdolon sensu lato is widely introduced in outside of its native areas in Europe, and also in New Zealand, Canada, and the United States, but authorities disagree about the distribution of introduced subspecies. As of April 2024, Plants of the World Online (POWO) claims that Lamium galeobdolon subsp. argentatum has been introduced in Great Britain and Ireland (in agreement with BSBI data) and Italy, but since subsp. argentatum is reported to be introduced in the Netherlands and Switzerland, the list is incomplete. POWO also claims that Lamium galeobdolon subsp. galeobdolon has been introduced in Madeira, New Zealand (both North Island and South Island), and the United States, but iNaturalist data show subsp. argentatum as much the most widespread subspecies in all these areas (click on the 'Map' tag for each of the subspecies). However, authoritative sources based in North America implicitly refer to Lamium galeobdolon sensu lato, so POWO's claim is unsubstantiated. The list of U.S. states where Lamium galeobdolon sensu lato is said to occur varies dramatically depending on the source.

Ecology

Lamium galeobdolon sensu lato has been widely introduced as a garden plant (often under the name Lamiastrum galeobdolon), especially in New Zealand, Canada, and the United States where it has escaped cultivation and become an invasive species. In New Zealand, it is listed by the 2020 National Pest Plant Accord and therefore banned from sale, propagation, and distribution throughout the country. It is also listed by the Invasive Species Council of British Columbia in Canada. In the U.S. state of Washington, it is listed as a Class B Noxious Weed and therefore banned from sale by state law. It is also a B-listed Noxious Weed in the state of Oregon.

The variegated yellow archangel (Lamium galeobdolon subsp. argentatum) has become an invasive subspecies in several European countries. In the Netherlands, subspecies argentatum was introduced as an ornamental ground cover, and by 1985 it had become naturalised and recorded in more localities than the native subspecies galeobdolon. It is also invasive in Britain where it spreads by stolons at the rate of 1–2 metres (3 ft 3 in – 6 ft 7 in) per growing season. In Switzerland, subspecies argentatum tripled its occurrence in four decades (1980–2020) while exhibiting "a higher growth rate and regeneration capacity" than the native subspecies galeobdolon.

Based on morphological characters, number of chromosomes, pattern of geographical distribution, and phytochemistry, early botanists hypothesized that the tetraploid subsp. montanum originated as a hybrid between the two diploid taxa, subsp. flavidum and subsp. galeobdolon. However, more recent phylogenetic data suggests that subsp. montanum originated from subsp. flavidum alone. Similarly, there is conflicting evidence regarding the origins of the tetraploid subsp. argentatum suggesting that it may or may not have originated from subsp. galeobdolon alone.

Etymology

Both the generic name Lamium and the specific name galeobdolon were used by Pliny the Elder in the first century AD. The name galeobdolon has several possible origins. It may come from the Latin words galeo meaning "to cover with a helmet" and dolon meaning "a fly's sting", or it may come from the Greek words galen meaning "weasel" and bdolos meaning "foetid smell". The latter interpretation is likely the source of the old common name 'yellow weasel-snout'.

The common names archangel and dead-nettle have been in use for hundreds of years. In 1578, the Flemish physician and botanist Rembert Dodoens observed that "Dead nettell groweth every where". The English herbalist John Gerard used the word "archangel" in 1633. Gerard believed the dead-nettles were so-named because their leaves resembled those of the true nettles in the family Urticaceae. The names "Arch-Angel" and "Dead Nettle" appear in the book The English Physitian (also known as the Complete Herbal) published by the English botanist and herbalist Nicholas Culpeper in 1652. Both Dodoens and Gerard believed the name "archangel" referred to members of genus Lamium with "clusters of hooded flowers at the nodes rising along the stems like a choir of robed figures". Others believe the name refers to the wing-like shape of its paired opposite leaves.

Gallery

• 
  Botanical illustration by Johann Georg Sturm (1796)
• 
  Flower cluster with yellow petals, hooded flowers, and unequal stamens
• 
  Leaves with silvery white variegated markings growing in Jūrmala, Latvia
• 
  Cluster of plants with yellow flowers and silvery white variegated leaves
• 
  Plants with yellow flowers growing in Hungary

See also

• List of plants in The English Physitian

Bibliography

• Bendiksby, Mika; Brysting, Anne K.; Thorbek, Lisbeth; Gussarova, Galina; Ryding, Olof (August 2011). "Molecular phylogeny and taxonomy of the genus Lamium L. (Lamiaceae): Disentangling origins of presumed allotetraploids". Taxon. 60 (4): 986–1000. doi:10.1002/tax.604004. JSTOR 41317321.
• DeFelice, Michael S. (2005). "Henbit and the Deadnettles, Lamium spp.: Archangels or Demons?". Weed Technology. 19 (3): 768–774. doi:10.1614/WT-05-072.1. JSTOR 3989505.
• Gilman, Arthur V. (2015). New Flora of Vermont. Memoirs of The New York Botanical Garden, Volume 110. Bronx, New York, USA: The New York Botanical Garden Press. ISBN 978-0-89327-516-7.
• Gledhill, David (2008). The Names of Plants (4th ed.). Cambridge University Press. ISBN 978-0-521-86645-3.
• Haines, Arthur (2011). New England Wild Flower Society's Flora Novae Angliae: A Manual for the Identification of Native and Naturalized Higher Vascular Plants of New England. Illustrated by Elizabeth Farnsworth and Gordon Morrison. Yale University Press. ISBN 978-0-300-17154-9.
• Mennema, J. (1989). A taxonomic revision of Lamium (Lamiaceae) (PDF). Leiden Botanical Series. Vol. 11. Leiden, The Netherlands: E. J. Brill. pp. 1–196. ISBN 90-04-09109-2. Retrieved 16 March 2024.
• Rosenbaumová, R.; Plačková, I.; Suda, J. (February 2004). "Variation in Lamium subg. Galeobdolon (Lamiaceae) — insights from ploidy levels, morphology and isozymes". Plant Systematics and Evolution. 244 (3/4): 219–244. Bibcode:2004PSyEv.244..219R. doi:10.1007/s00606-003-0071-5. JSTOR 23645275.
• Rusterholz, Hans-Peter; Huber, Katharina; Baur, Bruno (31 March 2023). "Invasion of a horticultural plant into forests: Lamium galeobdolon argentatum affects native above-ground vegetation and soil properties". Plants. 12 (7): 1527. doi:10.3390/plants12071527. PMC 10097399. PMID 37050152.  This article incorporates text available under the CC BY 4.0 license.
• Taylor, Brian; Glaister, Jim (2018). "Variegated Yellow Archangel Lamium galeobdolon subsp. argentatum". In Fennell, Mark; Jones, Laura; Wade, Max (eds.). Practical Management of Invasive Non-Native Weeds in Britain and Ireland. Liverpool University Press. pp. 103–104. doi:10.2307/j.ctv34h08r7. ISBN 978-1-85341-165-6. JSTOR j.ctv34h08r7.40.
• Wittmann, Helmut; Strobl, Walter (1986). "Zur Kenntnis der Gattung Galeobdolon Adans. im Bundesland Salzburg (Österreich)" (PDF). Berichte der Bayerischen Botanischen Gesellschaft zur Erforschung der Flora. 57: 163–176. Retrieved 26 March 2024.

External links

• Ellis, R. Gwynn, ed. (September 1987). "Variegated archangels" (PDF). B.S.B.I. News. 46. Cardiff: Dept. of Botany, National Museum of Wales: 9–11. Retrieved 26 March 2024.
• Ellis, R. Gwynn, ed. (April 1987). "Galeobdolon argentatum smejkal and other yellow archangels in Scotland" (PDF). B.S.B.I. News. 45. Cardiff: Dept. of Botany, National Museum of Wales: 45. Retrieved 26 March 2024.
• "Aluminium plant". Bay of Plenty Regional Council Toi Moana. Retrieved 27 March 2024.
• "Artillery plant". Weedbusters. Retrieved 27 March 2024.
• "Lamium galeobdolon". N.C. Cooperative Extension. Retrieved 27 March 2024.
• "Yellow Archangel (Lamiastrum galeobdolon)". Tualatin Soil and Water Conservation District. Retrieved 27 March 2024.
• "Lamiastrum galeobdolon subsp. argentatum, the variegated yellow archangel". Easy Wildflowers. January 27, 2015. Retrieved 29 March 2024.
• "B-Listed Noxious Weed Yellow Archangel reportedly seen for sale in Oregon". Oregon Department of Agriculture. Retrieved 5 April 2024.
• "Lamium galeobdolon - yellow archangel". Royal Horticultural Society.