Antirrhineae

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Antirrhineae
A nectar spur and a bright yellow "tongue" spot, as in these Linaria reflexa flowers, are widespread traits among the Antirrhineae
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Lamiales
Family: Plantaginaceae
Tribe: Antirrhineae
Dumort.[1]
Genera

About 30, see text

The Antirrhineae are one of the 12

snapdragons
.

Description and uses

common snapdragons (Antirrhinum majus) grown in Jerusalem

Most Antirrhineae are

glycosides.[2]

The Antirrhineae are not noted as food- or fodder plants, probably due to the iridoid content making them less than palatable. However, the tribe does not seem to contain highly poisonous plants, either; rather, use in

snapdragon", Antirrhinum majus
is probably the single most widely known member of the Antirrhineae by far.

Antirrhinum majus

invasive nuisance weed in agriculture, like other toadflaxes and snapdragons, its attractive flowers make it useful as a wildflower. At present however, the other members of this family are generally only of minor or localized interest.[3]

Taxonomy

The Antirrhineae include about 30 genera with roughly 320 species, of which 150 are in genus Linaria. The type genus is Antirrhinum L.[4][5]

Antirrhineae are probably most closely related to the

foxgloves (Digitalis), and speedwell
(Veronica).

Subdivision

Developing capsule fruit in the unusual calyx of Rhodochiton atrosanguineus

As the Antirrhineae have long been considered a distinct group, there has also been a long debate about recognition of distinct subdivisions. In 1909,

monotypic Rhodochitoneae, the latter due to their petal-like calyx. By the mid-20th century, Rothmaler on morphological grounds identified five subtribes containing 21 genera:[7][8]

  • Anarrhinum group / Anarrhinae / Simbuletinae (
    monotypic
    )
  • Gambelia group / Gambeliinae (3 genera)
  • Linaria group / Linariinae (11 genera)
  • Maurandya group / Maurandyinae (5 genera) – including Rhodochitoneae
  • Mohavea group / Mohaveinae (monotypic)
asterid Mentzelia involucrata, misleading its pollinator Xeralictus
beetles, as well as botanists researching Antirrhineae systematics

Some 21st-century

phylogenetic analyses indicate some quite different lineages. Mainly, the earlier authors seem to have overlumped the snapdragon-like forms (including toadflaxes) which actually do not seem to be closely related, while overemphasizing the morphological diversity of the true snapdragon relatives. As early as 1982,[9] Speta had realized that the typical toadflaxes (including Nuttallanthus) were a lineage well apart from the snapdragons and similar genera, and established the Antirrhininae for the latter. In 2000, combining internal transcribed spacer (ITS) and morphological data from 16 genera, Ghebrehiwet et al[10] confirmed Rothmaler's proposal of a close relationship between the fairly dissimilar-looking Maurandya and Rhodochiton and the distinctness of their lineage from the bulk of the subfamily. However, they found the "Linarieae" hard to resolve, but could already tell that Kickxia should be moved to the Anarrhinae, and Asarina and Cymbalaria to the Maurandyinae. In addition, Mohavea was recognized as a snapdragon relative with extremely modified flowers, refuting a monotypic Mohaveinae. Vargas et al in 2004[2] found six probable clades based on ITS and ndhF
sequences of 22 genera, which they labelled as:

However, they also noted profound morphological diversity amongst the Antirrhinum group, in accordance with the proliferation of segregated genera.

Using ITS data from all 29 then-recognized genera, the 2013 study of Fernández-Mazuecos et al.[5] identified six similar clades. They were able to assign all these genera to one of these clades, which they labelled as:

  • Anarrhinum clade (2 genera)
  • Antirrhinum clade (9 genera)
  • Chaenorhinum clade (3 genera)
  • Cymbalaria clade (9 genera)
  • Galvezia clade (3 genera)
    showy island snapdragon
    (Gambelia/Galvezia speciosa)
  • Linaria clade (2 genera)

The most striking difference between the 2004 and 2013 results is the precisely inverted placement of

hybrid introgression, or a disparity between nuclear (ITS) and chloroplast (ndhF) evolution or some other divergence[13] seems a more likely cause. The erratic behavior of the two genera was noted by Vargas et al but not discussed in Fernández-Mazuecos et al.; adding to the confusion, Ghebrehiwet et al found a strongly supported Galvezia fruticosa-Gambelia speciosa clade excluding Pseudorontium and Schweinfurthia
and equidistant from the true snapdragons and toadflaxes, entirely in line with Rothmaler's Gambeliinae. Furthermore, several species were historically moved between the two genera without authors noting anything suggesting against a very close relationship between them.

Regarding internal

basal
in one recent study or another, but the proposed interrelationships between the clades/subtribes other than Antirrhinum-Chaenorhinum are at most tenuously supported in all of them. As it seems, however, Rothmaler's general concept of an ancestral radiation of basal lineages and a subsequent diversification of the toadflax-snapdragon group was essentially correct, even though he overlumped the latter.

Genera

As recognized around 1900 already, the daisy-leaved toadflax (Anarrhinum bellidifolium) is not truly a toadflax

As of 2013, 29

genera are included in the Antirrhineae. Listed by clade, they are:[5][2]
Fairy snapdragon (Chaenorhinum origanifolium)
Cymbalaria pallida
The enigmatic Lafuentea rotundifolia flowering near the Castillo de San Julián in Spain
  • Linaria clade / Linariinae
    sensu stricto

Linnean taxonomy) of the Antirrhineae, far from Digitalis, Oreosolen, Ourisia or Rehmannia; in fact, these five genera are apparently all distinct from each other at least at tribal level (Oreosolen and Rehmannia are even outside the Plantaginaceae). However, the results were not unequivocal and support for including Lafuentea in the Antirrhineae not very robust. Besides, the interrelationships of the Plantaginaceae tribes are also not at all robustly resolved, and some fairly close relatives of the Antirrhineae – such as Campylanthus or the singular Hemiphragma – have similarly uncertain positions as Lafuentea with regard to their presumed closest relatives.[4] Thus, while Lafuentea seems to be a "living fossil" from near the origin of the Antirrhineae and is certainly highly useful as an outgroup
in cladistic studies of their internal relations, whether its placement within this tribe is correct requires further study.

References

  1. ^ Dumortier, B.C.J. (1827): Florula Belgica: 34.
  2. ^
    doi:10.1007/s00606-004-0216-1
    .
  3. PMID 21653448
    .
  4. ^
    PMID 21652407
    .
  5. ^
    doi:10.1086/668790
    .
  6. ^ Rouy, G. (1909): 'Conspectus' de tribus et des genres de la famille de Scrophulariacées. Revue générale de botanique 21: 194-207.
  7. ^ Rothmaler, W. (1943). "Zur Gliederung der Antirrhineae". Feddes Repertorium Specierum Novarum Regni Vegetabilis. 52: 16–39.
  8. ^ Rothmaler, W. (1956). "Taxonomische Monographie der Gattung Antirrhinum". Feddes Repertorium Specierum Novarum Regni Vegetabilis. 136: 1–124.
  9. ^ Speta, E. (1982). "Drei neue Antirrhineen-Gattungen aus dem Orient: Holzneria, Huebelia und Albraunia (Scrophulariaceae)". Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. 103 (1): 9–45.
  10. doi:10.1007/bf00985047
    .
  11. ^ "Chaenorrhinum" in Vargas et al (2004) is a lapsus.
  12. ^ Gambelia ITS was not included in the 2004 study, but the Pseudorontium-Schweinfurthia clade is labeled "Gambelia group". Galvezia fruticosa was included in both studies.
  13. ^ Compare the "suspicious" position of Antirrhinum in the rps16 intron analysis of Albach et al. (2005).
  14. doi:10.1111/njb.01000
  15. junior synonym
    .
  16. ^ GRIN (2011): GRIN Genera of Plantaginaceae tribe Antirrhineae.[dead link] Retrieved 2011-04-28.

Further reading

Wikimedia Commons has media related to Antirrhineae.
Wikispecies has information related to Antirrhineae.
  • Sutton DA. (1988) A revision of the tribe Antirrhineae. Oxford University Press, Oxford.

External links

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