page_title: Antirrhineae
section_title: Subdivision
hierarchical_section_title: Antirrhineae / Taxonomy / Subdivision
caption_attribution_description: English: Galvezia speciosa 'Firecracker' — native to the coastal California chaparral and woodlands. In The Water Conservation Garden At Cuyamaca College, located in El Cajon, southern California. Identified by sign.
context_page_description: The Antirrhineae are one of the 12 tribes of the family Plantaginaceae. It contains the toadflax relatives, such as snapdragons. They are probably most closely related to the turtlehead tribe and/or a large and badly resolved core group of their family including plants as diverse as water-starworts, foxgloves, and speedwell. The Antirrhineae include about 30 genera with roughly 320 species, of which 150 are in genus Linaria. The type genus is Antirrhinum L.
context_section_description: As the Antirrhineae have long been considered a distinct group, there has also been a long debate about recognition of distinct subdivisions. In 1909, Rouy separated the snapdragon-like subtribe Linarieae from the open-mouthed Anarrhineae and the 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: 
  Anarrhinum-Gruppe / Anarrhinae / Simbuletinae (monotypic)
  Gambelia-Gruppe / Gambeliinae (3 genera)
  Linaria-Gruppe / Linariinae (11 genera)
  Maurandya-Gruppe / Maurandyinae (5 genera) – including Rhodochitoneae
  Mohavea-Gruppe / Mohaveinae (monotypic)
  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, 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 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 found six probable clades based on ITS and ndhF sequences of 22 genera, which they labelled as:
  Anarrhinum group (2 genera)
  Antirrhinum group (at least 7 genera)
  Chaenorhinum group (3 genera)
  Gambelia group (at least 3 genera)
  Linaria group (1 or 2 genera)
  Maurandya group (at least 6 genera)
  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. 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)
  Linaria clade (2 genera)
  The most striking difference between the 2004 and 2013 results is the precisely inverted placement of Galvezia and Gambelia (one as core of a distinct clade, the other close to Antirrhinum as part of the Sairocarpus complex). Given the highly similar datasets and analyses, a clerical error confusing the two generic names might be suspected, but considering that the 2013 study included two species of each genus which congruently resolved as sister taxa, hybrid introgression, or a disparity between nuclear (ITS) and chloroplast (ndhF) evolution or some other divergence 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 phylogeny, at present, resolution is insufficient. In the more recent studies, the Chaenorhinum clade is fairly robustly resolved as sister to the true snapdragons, but otherwise not much is clear. Except for the Antirrhinum-Chaenorhinum and Linaria clades, all main lineages have resolved as basa
caption: Ornamental cultivar 'Firecracker' of the disputed showy island snapdragon (Gambelia/Galvezia speciosa)
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