Antirrhineae: Difference between revisions
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=== Subdivision=== |
=== Subdivision=== |
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As the Antirrhineae have long been considered a distinct group, there has also been a long debate about recognition of distinct subdivisions. In 1909, [[Georges C.Chr. Rouy|Rouy]]<ref>[[Georges C.Chr. Rouy|Rouy, G.C.C.]] (1909): 'Conspectus' de tribus et des genres de la famille de Scrophulariacées. ''Revue générale de botanique'' '''21''': 194-207.<!-- fulltext probably available on BHL --></ref> separated the snapdragon-like [[subtribe]] Linarieae from the open-mouthed Anarrhineae and the [[monotypic]] [[Rhodochitoneae]]<!-- this should redirect to Rhodochiton -->, the latter due to their [[petal]]-like [[sepal|calyx]]. By the mid-20th century, [[Werner Rothmaler|Rothmaler]] identified five subtribes on [[morphology (biology)|morphological]] grounds:<ref>[[Werner Rothmaler|Rothmaler, W.]] (1943): Zur Gliederung der Antirrhineae. ''Feddes Repertorium Specierum Novarum Regni Vegetabilis'' '''52''': 16–39.</ref> <ref>[[Werner Rothmaler|Rothmaler, W.]] (1956): Taxonomische Monographie der Gattung ''[[Antirrhinum]]''. ''Feddes Repertorium Specierum Novarum Regni Vegetabilis'' '''136''': 1–124.</ref> |
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* Anarrhinae ([[monotypic]]) – also Simbuletinae |
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* Gambeliinae (3 genera)<!-- Galvezia, "Saccularia", Gambelia ? --> |
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* Linariinae (11 genera) |
* Linariinae (11 genera) |
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* Maurandyinae (5) |
* Maurandyinae (5 genera) – including Rhodochitoneae |
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* Gambeliinae (3) |
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* Anarrhinae (1) |
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[[image:Mohavea_confertiflora_group.jpg|thumb|right|260px|[[Ghost flower]] (''Mohavea confertiflora'', a close snapdragon relative) mimics the [[sand blazing star]] (''Mentzelia involucrata'', an entirely unrelated [[asterid]]), confusing its [[pollinator]] ''[[Xeralictus]]'' beetles. This [[convergence (evolution)|convergence]] has long confounded Antirrhineae systematics.]] |
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However subsequent [[phylogenetic analyses]] have produced quite different lineages, such as the six of Vargas (2004), which they labelled as groups:<ref name=Vargas/> |
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* Anarrhinum |
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* Maurandya |
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* Antirrhinum |
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* Schweinfurthia |
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21st-century [[phylogenetic analyses]] indicate quite different lineages. Mainly, the earlier authors seem to have [[lumpers and splitters|overlumped]] the snapdragons-like forms (including toadflaxes) which actually do not seem to be closely related, while overemphasizing the morphological diversity of the true snapdragon relatives. But as early as 1982<ref>[[Franz Speta|Speta, E.]] (1982): Drei neue Antirrhineen-Gattungen aus dem Orient: ''[[Holzneria]]'', ''[[Huebelia]]'' und ''[[Albraunia]]'' (Scrophulariaceae). ''Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie'' '''103'''(1): 9-45.</ref> [[Franz Speta|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''<ref>Ghebrehiwet, Medhanie; Bremer, Birgitta & Thulin, Mats (2000): Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ''ndhF'' sequence data. ''[[Plant Systematics and Evolution|Plant Syst. Evol.]]'' '''220''' (3-4): 223–239. {{doi|10.1007/bf00985047}}</ref> still arrived at a scheme similar to Rothmaler's, but already could tell that ''[[Mohavea]]'' was a snapdragon with extremely modified flowers. On the other hand, the close relationship between the fairly dissimilar-looking ''[[Maurandya]]'' and ''[[Rhodochiton]]'' proposed by Rothmaler was confirmed. Vargas ''et al'' in 2004<ref name=Vargas/> found six probable [[clade]]s based on ITS and [[ndhF]] sequences of 22 genera, which they labelled as: |
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* ''Antirrhinum'' group (at least 7 genera) |
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* ''Gambelia'' group (at least 3 genera) |
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| ⚫ | |||
* ''Maurandya'' group (at least 6 genera) |
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| ⚫ | |||
Using ITS data from all 29 currently included genera, the 2013 phylogenetic study of Fernández-Mazuecos ''et al.''<ref>Fernández-Mazuecos, M.; Blanco-Pastor, J.L. & Vargas, P. (2013): A Phylogeny of Toadflaxes (''[[Linaria]]'' Mill.) Based on Nuclear Internal Transcribed Spacer Sequences: Systematic and Evolutionary Consequences. ''[[International Journal of Plant Sciences|Int. J. Plant Sci.]]'' '''174''': 234-249. {{doi|10.1086/668790}}</ref> identified six similar clades, and were able to assign all the genera in the tribe to one of these clades which they labelled as: |
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Fernández-Mazuecos ''et al.'' (2013), also using phylogenetics, identified six [[clades]] within the tribe, similar to those of Vargas ''et al.'', which they labelled as: |
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* '' |
* ''Anarrhinum'' clade (2 genera) |
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* '' |
* ''Antirrhinum'' clade (9 genera) |
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* ''Chaenorhinum'' (3) |
* ''Chaenorhinum'' clade (3 genera) |
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* '' |
* ''Cymbalaria'' clade (9 genera) |
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* ''Galvezia |
* ''Galvezia'' clade (3 genera) |
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* ''Linaria'' (2) |
* ''Linaria'' clade (2 genera) |
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The most striking difference between the 2003 and 2013 results is the precisely inverted placement of ''[[Galvezia]]'' and ''[[Gambelia (plant)|Gambelia]]'' (one as core of a distinct clade<ref>''[[Gambelia]]'' was not included in the 2003 study, but the ''[[Pseudorontium]]''-''[[Schweinfurthia]]'' clade is labeled "''Gambelia'' group". ''[[Galvezia]] fruticosa'' was included in both studies.</ref>, the other close to ''[[Antirrhinum]]'' as part of the ''[[Sairocarpus]]'' complex). Given the similar datasets, a [[clerical error]] confusing the two similar names might be suspected, but just as well a disparity between [[nuclear DNA|nuclear]] (ITS) and [[chloroplast]] (ndhF) evolution or some other [[divergent evolution|divergence]] may be the cause. 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, quite in line with Rothmaler's Gambeliinae. |
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They were able to assign all the genera in the tribe to one of these clades. |
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[[Saccularia]] |
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=== Genera === |
=== Genera === |
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Genera (29) usually included are shown here by clade: |
Genera (29) usually included are shown here by clade: |
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* ''Cymbalaria'' (9 genera) |
* ''Cymbalaria'' clade / Maurandyinae (9 genera) |
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**''[[Asarina]]'' <small>Mill.</small> |
**''[[Asarina]]'' <small>Mill.</small> * |
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**''[[Cymbalaria]]'' <small>Hill</small> |
**''[[Cymbalaria]]'' <small>Hill</small> * |
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**''[[Epixiphium]]'' <small>(Engelm. ex A.Gray) Munz</small> |
**''[[Epixiphium]]'' <small>(Engelm. ex A.Gray) Munz</small> ! |
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**''[[Holmgrenanthe]]'' <small>Elisens</small> |
**''[[Holmgrenanthe]]'' <small>Elisens</small> ! |
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**''[[Lophospermum]]'' <small>D.Don</small> |
**''[[Lophospermum]]'' <small>D.Don</small> * |
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**''[[Mabrya]]'' <small>Elisens</small> |
**''[[Mabrya]]'' <small>Elisens</small> ! |
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**''[[Maurandella]]'' |
**''[[Maurandella]]'' * |
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**''[[Maurandya]]'' <small>Ortega</small> |
**''[[Maurandya]]'' <small>Ortega</small> (*) |
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**''[[Rhodochiton]]'' <small>Zucc. ex Otto & A. Dietr.</small> |
**''[[Rhodochiton]]'' <small>Zucc. ex Otto & A. Dietr.</small> * |
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* ''Anarrhinum'' (2 genera) |
* ''Anarrhinum'' (2 genera) – Anarrhinae |
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**''[[Anarrhinum]]'' <small>Desf.</small> |
**''[[Anarrhinum]]'' <small>Desf.</small> * |
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**''[[Kickxia]]'' <small>Dumort.</small> |
**''[[Kickxia]]'' <small>Dumort.</small> * |
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* ''Chaenorhinum'' (3 genera) |
* ''Chaenorhinum'' (3 genera) – newly recognized |
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**''[[Albraunia]]'' <small>Speta</small> |
**''[[Albraunia]]'' <small>Speta</small> * |
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**''[[Chaenorhinum]]'' <small>(DC.) Rchb.</small> |
**''[[Chaenorhinum]]'' <small>(DC.) Rchb.</small> * |
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**''[[Holzneria]]'' <small>Speta</small> |
**''[[Holzneria]]'' <small>Speta</small> * |
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* ''Antirrhinum'' clade / Mohaveinae or Gambeliinae (disputed)<ref>Antirrhininae <small>Speta</small> is a [[junnior synonym]]</ref> (9 genera) |
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**''[[Acanthorrhinum]]'' <small>Rothm.</small> |
**''[[Acanthorrhinum]]'' <small>Rothm.</small> * |
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**''[[Antirrhinum]]'' <small>L.</small> |
**''[[Antirrhinum]]'' <small>L.</small> * |
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**''[[Gambelia (plant)|Gambelia]]'' <small>Nutt.</small> |
**''[[Gambelia (plant)|Gambelia]]'' <small>Nutt.</small> ! |
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**''[[Howelliella]]'' <small>Rothm.</small> |
**''[[Howelliella]]'' <small>Rothm.</small> * |
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**''[[Misopates]]'' <small>Raf.</small> |
**''[[Misopates]]'' <small>Raf.</small> * |
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**''[[Mohavea]]'' <small>A.Gray</small> |
**''[[Mohavea]]'' <small>A.Gray</small> * |
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**''[[Neogaerrhinum]]'' <small>Rothm.</small> |
**''[[Neogaerrhinum]]'' <small>Rothm.</small> * |
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**''[[Pseudomisopates]]'' <small>Güemes</small> |
**''[[Pseudomisopates]]'' <small>Güemes</small> * |
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**''[[Sairocarpus]]'' <small>D.A.Sutton</small> |
**''[[Sairocarpus]]'' <small>D.A.Sutton</small> * |
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* ''Galvezia |
* ''Galvezia'' clade or Gambeliinae (disputed) (3 genera) |
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**''[[Galvezia]]'' <small>Dombey ex Juss.</small> |
**''[[Galvezia]]'' <small>Dombey ex Juss.</small> (disputed) |
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**''[[Pseudorontium]]'' <small>(A.Gray) Rothm.</small> |
**''[[Pseudorontium]]'' <small>(A.Gray) Rothm.</small> |
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**''[[Schweinfurthia]]'' <small>A.Braun</small> |
**''[[Schweinfurthia]]'' <small>A.Braun</small> |
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* ''Linaria'' (2 genera) |
* ''Linaria'' clade / Linariinae ''[[sensu stricto]]'' (2 genera) |
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**''[[Linaria]]'' <small>Mill.</small> |
**''[[Linaria]]'' <small>Mill.</small> * |
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**''[[Nuttallanthus]]'' <small>D.A.Sutton</small> |
**''[[Nuttallanthus]]'' <small>D.A.Sutton</small> ! |
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Although [[Germplasm Resources Information Network|GRIN]] includes ''[[Lafuentea]]'' <small>Lag.</small> in the tribe,<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2283 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Antirrhineae'' |work=Germplasm Resources Information Network |accessdate=2011-04-28}}</ref> in this analysis it was a sister to the Antirrhineae. as also noted by Albach (2005)<ref name="albach">{{Cite journal | doi = 10.3732/ajb.92.2.297 | last1 = Albach | first1 = D. C. | last2 = Meudt | first2 = H. M. | last3 = Oxelman | first3 = B. | year = 2005 | title = Piecing together the "new" Plantaginaceae | url = http://www.amjbot.org/cgi/content/full/92/2/297 | journal = American Journal of Botany | volume = 92 | issue = 2| pages = 297–315 | pmid = 21652407 }}</ref> For the time being it should be considered an [[Outgroup (cladistics)|outgroup]]. |
Although [[Germplasm Resources Information Network|GRIN]] includes ''[[Lafuentea]]'' <small>Lag.</small> in the tribe,<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2283 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Antirrhineae'' |work=Germplasm Resources Information Network |accessdate=2011-04-28}}</ref> in this analysis it was a sister to the Antirrhineae. as also noted by Albach (2005)<ref name="albach">{{Cite journal | doi = 10.3732/ajb.92.2.297 | last1 = Albach | first1 = D. C. | last2 = Meudt | first2 = H. M. | last3 = Oxelman | first3 = B. | year = 2005 | title = Piecing together the "new" Plantaginaceae | url = http://www.amjbot.org/cgi/content/full/92/2/297 | journal = American Journal of Botany | volume = 92 | issue = 2| pages = 297–315 | pmid = 21652407 }}</ref> For the time being it should be considered an [[Outgroup (cladistics)|outgroup]]. |
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Halleria |
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Chelone |
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Tetranema |
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Veronica |
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Isoplexis |
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Plantago |
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Globularia |
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Digitalis |
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==References== |
==References== |
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Revision as of 02:02, 7 March 2016
| Antirrhineae | |
|---|---|
| |
| Antirrhinum majus | |
| Scientific classification | |
| Kingdom: | |
| (unranked): | |
| (unranked): | |
| (unranked): | |
| Order: | |
| Family: | |
| Tribe: | Antirrhineae |
| Genera | |
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see text | |
Antirrhineae are one of the twelve tribes of the Plantaginaceae family. The tribe contains about 29 genera in that family, of which the largest is Linaria, and about 320 species of which 150 are in Linaria. The type genus is Antirrhinum L. (Fernández-Mazuecos 2013).
Description
The flowers are tubular with a basal appendix (spur, gibbous or saccate). Other characters include poricidal capsule dehiscence and the possession of iridoid glycosides.[2]
Taxonomy
Subdivision
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[3] 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 identified five subtribes on morphological grounds:[4] [5]
- Anarrhinae (monotypic) – also Simbuletinae
- Gambeliinae (3 genera)
- Linariinae (11 genera)
- Maurandyinae (5 genera) – including Rhodochitoneae
- Mohaveinae (monotypic)
21st-century phylogenetic analyses indicate quite different lineages. Mainly, the earlier authors seem to have overlumped the snapdragons-like forms (including toadflaxes) which actually do not seem to be closely related, while overemphasizing the morphological diversity of the true snapdragon relatives. But as early as 1982[6] 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 ITS and morphological data from 16 genera, Ghebrehiwet et al[7] still arrived at a scheme similar to Rothmaler's, but already could tell that Mohavea was a snapdragon with extremely modified flowers. On the other hand, the close relationship between the fairly dissimilar-looking Maurandya and Rhodochiton proposed by Rothmaler was confirmed. Vargas et al in 2004[2] 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)
- Chaenorrhinum group (3 genera)
- Gambelia group (at least 3 genera)
- Linaria group (1-2 genera)
- Maurandya group (at least 6 genera)
However, they also noted profound morphological diversity amongst the Antirrhinum group, in accordance with the proliferation of segregate genera.
Using ITS data from all 29 currently included genera, the 2013 phylogenetic study of Fernández-Mazuecos et al.[8] identified six similar clades, and were able to assign all the genera in the tribe 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 2003 and 2013 results is the precisely inverted placement of Galvezia and Gambelia (one as core of a distinct clade[9], the other close to Antirrhinum as part of the Sairocarpus complex). Given the similar datasets, a clerical error confusing the two similar names might be suspected, but just as well a disparity between nuclear (ITS) and chloroplast (ndhF) evolution or some other divergence may be the cause. 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, quite in line with Rothmaler's Gambeliinae.
Genera
Genera (29) usually included are shown here by clade:
- Cymbalaria clade / Maurandyinae (9 genera)
- Asarina Mill. *
- Cymbalaria Hill *
- Epixiphium (Engelm. ex A.Gray) Munz !
- Holmgrenanthe Elisens !
- Lophospermum D.Don *
- Mabrya Elisens !
- Maurandella *
- Maurandya Ortega (*)
- Rhodochiton Zucc. ex Otto & A. Dietr. *
- Anarrhinum (2 genera) – Anarrhinae
- Anarrhinum Desf. *
- Kickxia Dumort. *
- Chaenorhinum (3 genera) – newly recognized
- Albraunia Speta *
- Chaenorhinum (DC.) Rchb. *
- Holzneria Speta *
- Antirrhinum clade / Mohaveinae or Gambeliinae (disputed)[10] (9 genera)
- Acanthorrhinum Rothm. *
- Antirrhinum L. *
- Gambelia Nutt. !
- Howelliella Rothm. *
- Misopates Raf. *
- Mohavea A.Gray *
- Neogaerrhinum Rothm. *
- Pseudomisopates Güemes *
- Sairocarpus D.A.Sutton *
- Galvezia clade or Gambeliinae (disputed) (3 genera)
- Galvezia Dombey ex Juss. (disputed)
- Pseudorontium (A.Gray) Rothm.
- Schweinfurthia A.Braun
- Linaria clade / Linariinae sensu stricto (2 genera)
- Linaria Mill. *
- Nuttallanthus D.A.Sutton !
Although GRIN includes Lafuentea Lag. in the tribe,[11] in this analysis it was a sister to the Antirrhineae. as also noted by Albach (2005)[12] For the time being it should be considered an outgroup.
Halleria Chelone Tetranema Veronica Isoplexis Plantago Globularia Digitalis
References
- ^ Fl. Belg.: 34 (1827)
- ^ a b Vargas P, JA Rosselló, R Oyama, J Güemes. 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 Syst Evol 249:151–172.
- ^ Rouy, G.C.C. (1909): 'Conspectus' de tribus et des genres de la famille de Scrophulariacées. Revue générale de botanique 21: 194-207.
- ^ Rothmaler, W. (1943): Zur Gliederung der Antirrhineae. Feddes Repertorium Specierum Novarum Regni Vegetabilis 52: 16–39.
- ^ Rothmaler, W. (1956): Taxonomische Monographie der Gattung Antirrhinum. Feddes Repertorium Specierum Novarum Regni Vegetabilis 136: 1–124.
- ^ Speta, E. (1982): Drei neue Antirrhineen-Gattungen aus dem Orient: Holzneria, Huebelia und Albraunia (Scrophulariaceae). Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 103(1): 9-45.
- ^ Ghebrehiwet, Medhanie; Bremer, Birgitta & Thulin, Mats (2000): Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological and ndhF sequence data. Plant Syst. Evol. 220 (3-4): 223–239. doi:10.1007/bf00985047
- ^ Fernández-Mazuecos, M.; Blanco-Pastor, J.L. & Vargas, P. (2013): A Phylogeny of Toadflaxes (Linaria Mill.) Based on Nuclear Internal Transcribed Spacer Sequences: Systematic and Evolutionary Consequences. Int. J. Plant Sci. 174: 234-249. doi:10.1086/668790
- ^ Gambelia was not included in the 2003 study, but the Pseudorontium-Schweinfurthia clade is labeled "Gambelia group". Galvezia fruticosa was included in both studies.
- ^ Antirrhininae Speta is a junnior synonym
- ^ "GRIN Genera of Plantaginaceae tribe Antirrhineae". Germplasm Resources Information Network. Retrieved 2011-04-28.
- ^ Albach, D. C.; Meudt, H. M.; Oxelman, B. (2005). "Piecing together the "new" Plantaginaceae". American Journal of Botany. 92 (2): 297–315. doi:10.3732/ajb.92.2.297. PMID 21652407.
Wikimedia Commons has media related to Antirrhineae.
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Bibliography
- Mario Fernández-Mazuecos, José Luis Blanco-Pastor, and Pablo Vargas. A Phylogeny of Toadflaxes (Linaria Mill.) Based on Nuclear Internal Transcribed Spacer Sequences: Systematic and Evolutionary Consequences. International Journal of Plant Sciences, Vol. 174, No. 2 (February 2013), pp. 234-249 The University of Chicago Press. Article DOI: 10.1086/668790
- Sutton DA. (1988) A revision of the tribe Antirrhineae. Oxford University Press, Oxford.
- Oyama, R. K.; Baum, D. A. (2004). "Phylogenetic relationships of North American Antirrhinum (Veronicaceae)". American Journal of Botany. 91 (6): 918–925. doi:10.3732/ajb.91.6.918. PMID 21653448.