Lophospermum

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Asterids
Cladus: Lamiids
Ordo: Lamiales

Familia: Plantaginaceae
Tribus: Antirrhineae
Subtribus: Maurandyinae
Genus: Lophospermum
Species: L. breedlovei – L. chiapense – L. erubescens – L. nubicola – L. purpurascens – L. purpusii – L. scandens – L. turneri
Name

Lophospermum D.Don ex R.Taylor, (1826)
Type species: Lophospermum scandens D.Don

Synonyms

Heterotypic

Rhodochiton Zucc. ex Otto & A. Dietr., Verh. Vereins. Beförd. Gartenbaues Königl. Preuss. Staaten 10: 152. 1834. However, contemporary evidence segregates the two genera and this circumscription is followed on Wikispecies.

References

Don, D., 1826. Philos. Mag. J. 67: 222.

Ghebrehiwet, M., Bremer, B. & Thulin, M. (2000) Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data, Plant Systematics and Evolution, 220: 223–239. Available on line [1]. Accessed 2014 29 May. Provides evidence that Lophospermum and Rhodochiton are sister taxa and are not cogeneric.

Christenhuiz, M.J.M. (2010) Nomenclatural corrections in Mesoamerican Plantaginaceae and a new species of Tetranema from Honduras, Phytotaxa 14: 56–60. Available on line [2]. Accessed 2014 29 May. Provides evidence that Lophospermum and Rhodochiton are sister taxa and are not cogeneric.

The Plant List 2013. Lophospermum in The Plant List Version 1.1. Published online. Accessed: 2014 May 29.

Farr, E. R. & Zijlstra, G. eds. (1996-) Index Nominum Genericorum (Plantarum). 2009 Dec 23 [3]. Shows Lophospermum and Rhodochiton as separate genera.

Elisens, W.J. 1985. Monograph of the Maurandyinae (Scrophulariaceae-Antirrhineae). Syst. Bot. Monogr. 5:66. Shows Lophospermum and Rhodochiton as cogeneric.

Sutton, D.A. 1988. A revision of the tribe Antirrhineae. (Rev Antirrh) 497. Shows Lophospermum and Rhodochiton as cogeneric.

Vernacular names
English: Creeping Gloxinia

Lophospermum is a genus of herbaceous perennial climbers or scramblers, native to mountainous regions of Mexico and Guatemala. Those that climb use twining leaf stalks. Their flowers are tubular, in shades of red, violet and purple, the larger flowers being pollinated by hummingbirds. Now placed in the greatly expanded family Plantaginaceae, the genus was traditionally placed in the Scrophulariaceae. The close relationship with some other genera, particularly Maurandya and Rhodochiton, has led to confusion over the names of some species.

Lophospermum erubescens and Lophospermum scandens are cultivated as ornamental plants, as climbers or trailers. Various Lophospermum cultivars are grown, often under trade names such as "Lofos®".

Description
Close up of flowers of a Lophospermum grown from commercial seed; probably Lophospermum scandens

Lophospermum species are herbaceous perennial climbers with fibrous roots. They climb by means of twining leaf stalks (petioles) rather than tendrils or twining stems. The long stems are branched, becoming woody at the base with age. In some species the stems grow from a woody caudex – a swollen, bulb-like structure at the base of the stem. The leaves are triangular or heart-shaped with a pointed apex and toothed edges (crenate or dentate). Both stems and leaves may have a purplish colour.[1]

Species generally flower and fruit over a long season; for example, from April to the following January in the case of Lophospermum erubescens. The flowers are borne singly on stems (peduncles) which are either held horizontally or grow upwards. The calyx has sepals that are either free or somewhat joined at the base, and overlap or curve outwards. They may be green or tinted with red or purple. The flowers have five petals in shades of red, violet or dark purple, joined at the base to form a tube. The free lobes of the upper two petals are differentiated from those of the lower three petals: for example, the upper two may curve back and the lower three point forwards. There are two prominent folds (plicae) running along the length of the base of the flower tube, usually yellowish in colour with hairs whose length helps to distinguish the species. There are four fertile stamens, of two different lengths, and one infertile stamen. The stamens and style are held inside the flower.[1]

The ovary has two chambers (locules). After fertilization, an ovoid or globe-shaped capsule forms filled with brown seeds, each with a circular "wing" around it.[1]
Taxonomy

The genus Lophospermum was first described in March 1826 by David Don in a paper read to the Linnean Society of London. His account was summarized in the same month in the Philosophical Magazine and Journal and subsequently published in 1827 in the Transactions of the Linnean Society.[2] Don described Lophospermum as closely related to Antirrhinum and Maurandya but distinguished by its bell-shaped (campanulate) flowers and winged seeds. Lophospermum means "with crested seeds".[3] Initially two species were described, Lophospermum scandens and Lophospermum physalodes.[4] (The latter is now Melasma physalodes.[5])

The genus is placed in the tribe Antirrhineae; within this tribe, it is closely related to Maurandya (including Maurandella), Mabrya and Rhodochiton.[6] It has been included in Maurandya as section Lophospermum.[7] Rhodochiton has been included in Lophospermum as section Rhodochiton.[1][8] Scientific names within these three genera have been confused; for example, an image accepted by Tropicos as Lophospermum erubescens bears the caption Maurandya barclaiana (a variant spelling of Maurandya barclayana).[9]
Phylogeny

A number of molecular phylogenetic studies have shown that subtribe Maurandyinae, defined by Elisen to consist of the five North American genera Holmgrenanthe, Lophospermum, Mabrya, Maurandya and Rhodochiton, forms a monophyletic group, which is related to the Old World genera Cymbalaria and Asarina.[6][10][11] Gehebrehiwet et al. suggested that the Maurandyinae could be expanded to include Cymbalaria and Asarina.[10] Vargas et al. presented the following cladogram in 2013:[11]

Antirrhineae   other clades       Cymbalaria lineage     Cymbalaria     Asarina     Maurandyinae sensu Elisens   Maurandya (including M. antirrhiniflora)       Rhodochiton       Lophospermum     Mabrya               other clades

Old World New World

Vargas et al. concluded that the Antirrhineae evolved in the Old World and subsequently colonized North America more than once, probably in the Miocene epoch (23 to 5 million years ago). One such colonization led to the evolution of the Maurandyinae (in Elisen's sense).[11]
Species
Lophospermum purpusii

As of July 2014, The Plant List accepts seven species (based on Tropicos and the World Checklist of Selected Plant Families):[Note 1]

Lophospermum breedlovei Elisens
Lophospermum chiapense Elisens
Lophospermum erubescens D.Don
Lophospermum purpurascens Elisens
Lophospermum purpusii (Brandegee) Rothm.
Lophospermum scandens D.Don
Lophospermum turneri Elisens

In his 1985 monograph, Wayne J. Elisens included Rhodochiton in Lophospermum as section Rhodochiton:[1]

Lophospermum atrosanguineum = Rhodochiton atrosanguineum (Zucc.) Rothm.
Lophospermum hintonii = Rhodochiton hintonii (Elisens) D.A.Sutton
Lophospermum nubicola = Rhodochiton nubicola (Elisens) D.A.Sutton

Distribution and habitat
Distribution of Lophospermum species: the distribution is discontinuous within the two areas shown.[1]

Species of Lophospermum are native to mountainous regions of Mexico and parts of adjoining Guatemala: the Sierra Madre Oriental, the Sierra Madre del Sur and the Sierra Madre de Chiapas, as well as the Altiplano and the Cordillera Neovolcánica. Most species have small discrete ranges; the exception is L. erubescens which has a wider distribution in oak forests of the Sierra Madre Oriental.[12] They are usually found between 500 and 3,000 m (1,600 and 9,800 ft) in oak or oak-pine forests or deciduous tropical forests (L. purpusii). L. scandens will grow on recent lava flows.[13]

Lophospermum erubescens has become naturalized in many tropical and subtropical areas of the world, including Colombia, Venezuela, Jamaica and Hawaii,[14] as well as Australia (New South Wales and Queensland).[15]
Ecology

Lophospermum erubescens and L. scandens are known to be pollinated by hummingbirds. Species pollinated in this way typically have yellow to red, open throated flowers with long floral tubes,[16] up to 63 mm (2.5 in) long in L. erubescens.[1] The pollinators of the remaining species are unknown, but Elisens suggests that, based on floral colour and morphology, L. purpusii is similarly hummingbird-pollinated, whereas L. breedlovei, L. chiapense, L. purpurascens and L. turneri may be pollinated by bees, as they have differently shaped flowers, with landing platforms and narrower openings to the flower tubes.[1] The nectar composition of L. purpusii is similar to other hummingbird-pollinated flowers, whereas that of L. turneri is quite different, with a very high proportion of sucrose.[16]
Cultivation
Comparison of Lophospermum scandens with Maurandya barclayana:
1 Maurandya flower is shorter with narrower sepals
2 Maurandya leaf is smooth with entire (untoothed) margins
3 Lophospermum flower is longer with broader sepals
4 Lophospermum leaf is somewhat hairy with toothed margins

Lophospermum erubescens has been cultivated as an ornamental climber since at least the 19th century. Joseph Paxton described its cultivation in 1836, saying that it was "a very fine creeper and deserves growing by every lover of plants".[17] Other species that have been cultivated include L. purpusii and L. scandens.[18] Some cultivars of Lophospermum are available; Suntory Flowers (a division of the Japanese firm Suntory) has introduced a number under the registered name "Lofos®", with selling names such as Compact White, Compact Pink, Wine Red (='Sunasaro') and Summer Cream (='Sunasashiro').[19][20] Those which are the subject of US Plant Patents are described as originating from the hybrid Lophospermum scandens × L. erubescens,[21] a hybrid known since the 1840s.[22]

As noted above, Lophospermum species have in the past been placed in Maurandya and the two genera have regularly been confused, particularly in cultivation. For example, the cultivar Lophospermum 'Magic Dragon' (a cross between Lophospermum 'Red Dragon' and L. erubescens) was patented under the genus name Maurandya.[23] Cultivated species of Lophospermum have longer flowers than those of Maurandya and leaves with toothed rather than entire margins.[1] The Suntory cultivars were patented under the generic name Asarina,[21] now treated as an entirely European genus.[6]

Plants may be grown from seed and treated as annuals. In frost-free climates, or where the roots can be protected from frost, plants may be perennial, regrowing from the base after dying back in the winter.[18] Selected forms and cultivars may be grown from cuttings.[23]
Notes

In addition to these seven species, The Plant List (version 1.1) also gives Lophospermum nubiculum Elisens as an accepted name, citing Tropicos.[24] However, as of 16 July 2014, the record for this name is marked "inactive" in Tropicos if reached via The Plant List, and the name cannot be found in a search. The name Lophospermum nubiculum does not appear in Elisens (1985), and appears to be an orthographic error for Lophospermum nubicola (a synonym of Rhodochiton nubicola).

References

Elisens, Wayne J. (1985), "Monograph of the Maurandyinae (Scrophulariaceae-Antirrhineae)", Systematic Botany Monographs, 5: 1–97, doi:10.2307/25027602, JSTOR 25027602
"IPNI Plant Name Query Results for Lophospermum", The International Plant Names Index, retrieved 2014-07-12
Hyam, R. & Pankhurst, R.J. (1995), Plants and their names : a concise dictionary, Oxford: Oxford University Press, p. 195, ISBN 978-0-19-866189-4
Taylor, R. (1827), "Proceedings of Learned Societies: Linnaean Society", Philosophical Magazine and Journal, 68: 222
"Lophospermum physalodes", The Plant List, retrieved 2014-07-12
Vargas, P; Rosselló, J.A.; Oyama, R. & Güemes, J. (2004), "Molecular evidence for naturalness of genera in the tribe Antirrhineae (Scrophulariaceae) and three independent evolutionary lineages from the New World and the Old", Plant Systematics and Evolution, 249 (3–4): 151–172, doi:10.1007/s00606-004-0216-1, S2CID 20328396
"Lophospermum D.Don", Tropicos.org, Missouri Botanical Garden, retrieved 2014-07-12
"Rhodochiton Zucc. ex Otto & A. Dietr.", Tropicos.org, Missouri Botanical Garden, retrieved 2014-07-12
"Image – Lophospermum erubescens", Tropicos.org, Missouri Botanical Garden, retrieved 2014-07-12
Ghebrehiwet, Medhanie; Bremer, Birgitta & Thulin, Mats Thulin (2000), "Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data", Plant Systematics and Evolution, 220 (3–4): 223–239, doi:10.1007/bf00985047, S2CID 36061550
Vargas, Pablo; Valente, Luis M.; Blanco-Pastor, José Luis; Liberal, Isabel; Guzmán, Beatriz; Cano, Emilio; Forrest, Alan & Fernández-Mazuecos, Mario (2013), "Testing the biogeographical congruence of palaeofloras using molecular phylogenetics: snapdragons and the Madrean–Tethyan flora", Journal of Biogeography, 41 (5): 932–943, doi:10.1111/jbi.12253
Elisens 1985, p. 25
Elisens 1985, p. 68
Elisens 1985, pp. 18, 75
Barker, W.R. & Harden, G.J., "Lophospermum erubescens D.Don", New South Wales Flora Online, retrieved 2014-08-07
Elisens, Wayne J. & Freeman, C. Edward (1988), "Floral Nectar Sugar Composition and Pollinator Type Among New World Genera in Tribe Antirrhineae (Scrophulariaceae)", American Journal of Botany, 75 (7): 971–978, doi:10.2307/2443763, JSTOR 2443763
Paxton, J. (1836), "Select List of Ornamental Creepers", Paxton's Magazine of Botany, and Register of Flowering Plants, 2: 33–37, retrieved 2014-07-13
Sanders, T.W. & Hellyer, A.G.L. (1966), Sanders' Encyclopaedia of Gardening (22nd ed.), London: Collingridge, OCLC 123581294 – entries under Maurandia, pp. 303–304
Lofos® | Suntory Collection Europe, archived from the original on 2014-07-13, retrieved 2014-07-13
"Search for Lophospermum", RHS Horticultural Database, Royal Horticultural Society, retrieved 2014-08-09
"Asarina plant named 'Sun-Asaro'", US Plant Patent, 19 December 2002
Bailey, L.H. (1919), "Maurandia", The Standard Cyclopedia of Horticulture, vol. 4 L-O, vol. 4, New York: Macmillan, pp. 2012–2013, OCLC 313800248
"Maurandya plant named 'Magic Dragon'", US Plant Patent, 19 December 2002
"Search for "Lophospermum"", The Plant List, retrieved 2014-07-09