Glandularia

Mock vervains
Glandularia bipinnatifida has chloroplasts acquired from a North American true vervain species via horizontal transfer.
Scientific classification
Kingdom:	Plantae
(unranked):	Angiosperms
(unranked):	Eudicots
(unranked):	Asterids
Order:	Lamiales
Family:	Verbenaceae
Genus:	Glandularia J.F.Gmel.
Species
Many, see text

Glandularia canadensis, a member of the northern lineage.
Its chloroplasts derive from a South American Verbena

Glandularia peruviana, a member of the southern lineage

Glandularia pulchella

Glandularia, called mock vervains or mock verbenas, is a genus of annual and perennial herbaceous flowering plants in the vervain family, Verbenaceae. They are native to the Americas.

Closely related to the true vervains^[1] and sometimes still included with them in Verbena, horizontal chloroplast transfer occurred at least twice and possibly thrice between these genera, which are otherwise rather too distinct to warrant unification. The discovery of the signal in the chloroplast genome was announced in 2008 by University of Washington researchers. Somehow, chloroplasts from V. orcuttiana, Swamp Verbena (V. hastata) or a close relative of these had admixed into the G. bipinnatifida germline. Although hybridization runs rampant in the true and mock vervains – the ancestors of the well-known Garden Vervain are quite obscure –, it does not seem to have been the cause of the cross-species gene transfer.^[2]

In addition, the researchers discovered the signal of one more transfer event. This had introduced chloroplasts from an ancestral member of the Verbena lineage nowadays found in South America into Glandularia Although all members of the present genus can be distinguished to have a chromosome count of 5, the South American species are diploid, while polyploid hybrid Glandularia are very widespread from northern Central America northwards. The second genetic introgression must have occurred before the genus spread north, as species with the Verbena-like chloroplasts are found all over the Americas. Since the new chloroplast genes replaced the old ones, it may be that the possibly hybridogenic G. bipinnatifida actually underwent horizontal chloroplast transfer twice in its evolutionary history.^[2]

Selected species^[3]

Glandularia alejandrana B.L.Turner Glandularia amoena (Paxton) Umber Glandularia araucana (Phil.) Botta Glandularia aristigera (S.Moore) Tronc. Glandularia aurantiaca (Speg.) Botta Glandularia bajacalifornica (Moldenke) Umber Glandularia ballsii (Moldenke) Botta Glandularia bipinnatifida (Nutt.) Nutt.[verification needed] Glandularia brachyrhynchos G.L.Nesom & Vorobik Glandularia cabrerae (Moldenke) Botta Glandularia canadensis (L.) Nutt. Glandularia cheitmaniana (Moldenke) Botta & Poggio Glandularia chiracahensis Umber Glandularia crithmifolia (Gillies & Hook. ex Hook.) Schnack & Covas Glandularia delticola (Small) Umber Glandularia dissecta (Willd. ex Spreng.) Schnack & Covas Glandularia elegans (Kunth) Umber Glandularia flava (Gillies & Hook. ex Hook.) Schnack & Covas Glandularia glandulifera (Moldenke) Ragonese Glandularia gooddingii (Briq.) Solbrig Glandularia gooddingii var. gooddingii Glandularia gooddingii var. nepetifolia Tidestrom Glandularia guaranitica Tronc. Glandularia hasslerana (Briq.) Tronc. Glandularia herteri (Moldenke) Tronc. Glandularia hookeriana Covas & Schnack Glandularia × hybrida (hort. ex Groenl. & Rümpler) G. L. Nesom & Pruski – Hybrid Verbena, Garden Vervain (often placed in Verbena) Glandularia incisa (Hook.) Tronc. Glandularia kuntzeana (Moldenke) Tronc. Glandularia laciniata (L.) Schnack & Covas Glandularia lilacina (Greene) Umber Glandularia lilacina cv. 'De La Mina'. Glandularia lilloana (Moldenke) Botta

Glandularia lipozygoides (Walp.) L.E.Navas Glandularia macrosperma (Speg.) Tronc. Glandularia maritima (Small) Small Glandularia marrubioides (Cham.) Tronc. Glandularia megapotamica (Spreng.) Cabrera & G.Dawson Glandularia mendocina (Phil.) Covas & Schnack Glandularia microphylla (Kunth) Cabrera Glandularia nana (Moldenke) Tronc. Glandularia origenes (Phil.) Schnack & Covas Glandularia parodii Covas & Schnack Glandularia perakii Covas & Schnack Glandularia peruviana (L.) Small Glandularia phlogiflora (Cham.) Schnack & Covas Glandularia platensis (Spreng.) Schnack & Covas Glandularia polyantha Umber Glandularia pulchella (Sweet) Tronc. Glandularia pulchra (Moldenke) Botta Glandularia pumila (Rydb.) Umber Glandularia quadrangulata (A.Heller) Umber Glandularia radicata (Moldenke) Tronc. ex Mulgura Glandularia santiaguensis Covas & Schnack Glandularia scrobiculata (Griseb.) Tronc. Glandularia selloi (Spreng.) Tronc. Glandularia sessilis (Cham.) Tronc. Glandularia spectabilis (Moldenke) Botta Glandularia stellarioides (Cham.) Schnack & Covas Glandularia subincana Tronc. Glandularia sulphurea (D.Don) Schnack & Covas Glandularia tampensis (Nash) Small[verification needed] Glandularia tenera (Spreng.) Cabrera Glandularia teucriifolia (M.Martens & Galeotti) Umber Glandularia tristachya (Tronc. & Burkart) Schnack & Covas Glandularia tumidula (L.M.Perry) Umber Glandularia turneri G.L.Nesom Glandularia venturii (Moldenke) Botta Glandularia verecunda Umber

Footnotes

1. ^ Botta et al. (1995)
2. ^ ^{a b} Yuan & Olmstead (2008)
3. ^ USDA (1999)

References

• Botta, S.M.: Martinez, S. & Mulgura de Romero, M.E. (1995): Novedades nomenclaturales en Verbenaceae ["Nomenclatural revisions in Verbenaceae"]. Hickenia 2: 127-128.
• United States Department of Agriculture (USDA) (1999): Germplasm Resources Information Network - Glandularia. Version of 1999-AUG-30. Retrieved 2008-AUG-07.
• United States Department of Agriculture (USDA) [2008]: USDA Plants Profile: Verbena bonariensis. Retrieved 2008-AUG-07.
• Yuan, Yao-Wu & Olmstead, Richard G. (2008): A species-level phylogenetic study of the Verbena complex (Verbenaceae) indicates two independent intergeneric chloroplast transfers. Mol. Phylogenet. Evol. 48(1): 23-33. doi:10.1016/j.ympev.2008.04.004 (HTML abstract)