Plantaginaceae

{{Short description|Family of flowering plants in the order Lamiales}} {{Distinguish|Platanaceae}} {{Automatic taxobox |image = Scoparia dulcis at Kadavoor.jpg |image_caption = ''Scoparia dulcis'' |taxon = Plantaginaceae |authority = Juss.<ref name="APGIII-2009">{{Cite journal |last=Angiosperm Phylogeny Group |year=2009 |title=An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III |journal=Botanical Journal of the Linnean Society |volume=161 |issue=2 |pages=105–121 |url=http://onlinelibrary.wiley.com/doi/10.1046/j.1095-8339.2003.t01-1-00158.x/pdf | format= PDF |access-date=2013-07-06 |doi=10.1111/j.1095-8339.2009.00996.x |doi-access=free |hdl=10654/18083 |hdl-access=free }}</ref> |type_genus = ''Plantago'' |type_genus_authority = L. |subdivision_ranks = Tribes |subdivision = * Angelonieae * Antirrhineae * Callitricheae * Cheloneae * Digitalideae * Globularieae * Gratioleae * Hemiphragmeae * Plantagineae * Russelieae * Sibthorpieae * Veroniceae |synonyms = * Antirrhinaceae Pers. * Aragoaceae D.Don * Callitrichaceae Link nom. cons. * Chelonaceae Martinov * Digitalaceae Martinov * Ellisiophyllaceae Honda * Globulariaceae DC. nom. cons. * Gratiolaceae Martinov * Hippuridaceae Vest nom. cons. * Littorellaceae Gray * Psylliaceae Horan. * Sibthorpiaceae D.Don * Veronicaceae Cassel |synonyms_ref = <ref name="GRIN-2003">{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?888 |title=Family: ''Plantaginaceae'' Juss., nom. cons. |work=Germplasm Resources Information Network |date=2003-01-17 |access-date=2011-04-28 |url-status=dead |archive-url=https://web.archive.org/web/20110704093834/http://www.ars-grin.gov/cgi-bin/npgs/html/family.pl?888 |archive-date=2011-07-04 }}</ref> }}

'''Plantaginaceae''', the '''plantain family''' or '''veronica family''', is a large, diverse family of flowering plants in the order Lamiales that includes common flowers such as snapdragon and foxglove. It is unrelated to the banana-like fruit also called "plantain". In older classifications, Plantaginaceae was the only family of the order '''Plantaginales''', but numerous phylogenetic studies, summarized by the Angiosperm Phylogeny Group, have demonstrated that this taxon should be included within Lamiales.

==Overview== The plantain family as traditionally circumscribed consisted of only three genera: ''Bougueria'', ''Littorella'', and ''Plantago''. However phylogenetic research has indicated that Plantaginaceae ''sensu stricto'' (in the strict sense) were nested within Scrophulariaceae (but forming a group that did not include the type genus of that family, ''Scrophularia''). Although Veronicaceae (1782) is the oldest family name for this group, Plantaginaceae (1789) is a conserved name under the International Code of Botanical Nomenclature (ICBN) and thus has priority over any earlier family name for a family including ''Plantago''. Furthermore, the ICBN does not consider family names published before 1789 to be names eligible for conservation, thus ruling out Veronicaceae. The name '''Antirrhinaceae''' has been proposed for conservation over Plantaginaceae. In the meantime, the Angiosperm Phylogeny Group has accepted the name Plantaginaceae. However, Olmstead (2003) chose to use the name Veronicaceae, a later synonym.<ref>{{Cite journal|last=Olmstead|first=Richard G.|date=2002|title=Whatever happened to the Scrophulariaceae?|url=http://courses.washington.edu/bot113/Spring/WebReadings/PdfReadings/Scrophulariaceae.pdf|journal=Fremontia|volume=30|pages=13–22}}</ref> The Plantaginaceae is additionally called the veronica family in English.<ref>{{cite book |last1=Chase |first1=Mark |editor1-last=Henry |editor1-first=Robert J. |title=Plant Diversity and Evolution : Genotypic and Phenotypic Variation in Higher Plants |date=2005 |publisher=CABI Pub. |location=Wallingford, Oxfordshire, UK; Cambridge, Massachusetts |page=20 | isbn=978-0-85199-904-3 |chapter-url=https://archive.org/details/plantdiversityev0000henr/page/20 |access-date=8 January 2025 |chapter=2. Relationships between the families of flowering plants}}</ref>

A group of genera including ''Lindernia'' has now been segregated<ref name="Albach-2005" /><ref name="Oxelman-2005">{{Cite journal | doi = 10.2307/25065369 | last1 = Oxelman | first1 = B. | last2 = Kornhall | first2 = P. | last3 = Olmstead | first3 = R. G. | last4 = Bremer | first4 = B. | year = 2005 | title = Further disintegration of Scrophulariaceae | journal = Taxon | volume = 54 | issue = 2| pages = 411–425 | jstor=25065369 | bibcode = 2005Taxon..54..411O }}</ref> as the family Linderniaceae,<ref>{{cite journal |last1=Rahmanzadeh |first1=R. |first2=K. |last2=Müller |first3=E. |last3=Fischer |first4=D. |last4=Bartels |first5=T. |last5=Borsch |date=2005 |title=The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales) |periodical=Plant Biology |volume=7 |issue=1 |pages=67–78 |pmid=15666207|doi=10.1055/s-2004-830444 |bibcode=2005PlBio...7...67R |s2cid=260252656 }}</ref> which is recognized by Haston ''et al.'' 2007 (also known as LAPG II) as "Post-APG II family."

Plantaginaceae ''sensu lato'' (in the broad sense) are a diverse, cosmopolitan family, occurring mostly in temperate zones. The group consists of herbs, shrubs and also a few aquatic plants with roots (such as the genus ''Callitriche''). As the family is so diverse, its circumscription is difficult to establish.<ref name="Albach-2005">{{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 | journal = American Journal of Botany | volume = 92 | issue = 2| pages = 297–315 | pmid = 21652407 | bibcode = 2005AmJB...92..297A }}</ref>

The leaves are spiral to opposite and simple to compound. Unusual in Lamiales is the absence of vertical partitions in the heads of the glandular hairs.

The structure and form of the flowers is variable. Some genera are 4-merous (i.e., with 4 sepals and 4 petals), such as ''Aragoa'' (but this one has 5 sepals); others are 5- to 8-merous, such as ''Sibthorpia''. The flowers of most genera are polysymmetric. The corolla is often two-lipped. In some taxa, the androecium is formed before the corolla.

The fruit is a loculicidal capsule, dehiscing through the partitions between the cells.<ref>{{cite journal| last =Haston, E.| first = Richardson, J. E., Stevens, P. F., Chase, M. W., Harris, D. J. | title = A linear sequence of Angiosperm Phylogeny Group II families| journal = Taxon| volume = 56| issue = 1| year = 2007| pages = 7–12| doi =10.2307/25065731 | jstor = 25065731 | bibcode = 2007Taxon..56....7H | doi-access = free}}</ref>

==Genera== [[File:Bacopa monnieri W IMG 1612.jpg|thumb|upright=0.86|''Bacopa monnieri'' in Hyderabad, India. ]] [[File:Achetaria azurea at Kadavoor.jpg|thumb|''Matourea azurea'' in Kerala]] The enlarged Plantaginaceae consists of 107 genera<ref name="POWO">[https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30001136-2 Plantaginaceae Juss.] ''Plants of the World Online''. Retrieved 14 May 2024.</ref> and about 1,900 species.<ref name="Byng-2016">{{cite journal |author1=Christenhusz, M. J. M. |author2=Byng, J. W. | year = 2016 | title = The number of known plants species in the world and its annual increase | journal = Phytotaxa | volume = 261 | pages = 201–217 | url = http://biotaxa.org/Phytotaxa/article/download/phytotaxa.261.3.1/20598 | doi = 10.11646/phytotaxa.261.3.1 | issue = 3 |bibcode=2016Phytx.261..201C | doi-access = free }}</ref> The largest genus is ''Veronica'', with about 450 species. ''Veronica'' also includes the genera ''Hebe'', ''Parahebe'' and ''Synthyris'', formerly often treated as distinct. All genera of Plantaginaceae were formerly included in Scrophulariaceae except where otherwise stated.

{| |- valign=top | ; Tribe Angelonieae * ''Angelonia'' Humb. & Bonpl. * ''Basistemon'' Turcz. * ''Melosperma'' Benth. * ''Monttea'' Gay * ''Ourisia'' Comm. ex Juss.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2294 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Angelonieae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052612/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2294 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Antirrhineae * ''Acanthorrhinum'' Rothm. * ''Albraunia'' Speta * ''Anarrhinum'' Desf. * ''Antirrhinum'' L. * ''Asarina'' Mill. * ''Chaenorhinum'' (DC.) Rchb. * ''Cymbalaria'' Hill * ''Epixiphium'' (Engelm. ex A.Gray) Munz * ''Gadoria'' {{small|Güemes & Mota}} * ''Galvezia'' Dombey ex Juss. * ''Gambelia'' Nutt. * ''Holmgrenanthe'' Elisens * ''Holzneria'' Speta * ''Howelliella'' Rothm. * ''Kickxia'' Dumort. * ''Linaria'' Mill. * ''Lophospermum'' D.Don * ''Mabrya'' Elisens * ''Maurandella'' {{small|(A.Gray) Rothm.}} * ''Maurandya'' Ortega * ''Misopates'' Raf. * ''Mohavea'' A.Gray * ''Nanorrhinum'' {{small|Betsche}} * ''Neogaerrhinum'' Rothm. * ''Nuttallanthus'' D.A.Sutton * ''Pseudomisopates'' Güemes * ''Pseudorontium'' (A.Gray) Rothm. * ''Rhodochiton'' Zucc. ex Otto & A. Dietr. * ''Sairocarpus'' D.A.Sutton * ''Schweinfurthia'' A.Braun<ref name="GRIN">{{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 |access-date=2011-04-28}}</ref> ; Tribe Callitricheae * ''Callitriche'' L. * ''Hippuris'' L.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2290 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Callitricheae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052606/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2290 |archive-date=2015-09-24 |url-status=dead }}</ref> | ; Tribe Cheloneae * ''Brookea'' Benth. * ''Chelone'' L. * ''Chionophila'' Benth. * ''Collinsia'' Nutt. * ''Keckiella'' Straw * ''Nothochelone'' (A.Gray) Straw * ''Pennellianthus'' Crosswh.<ref>{{cite journal |last1=Crosswhite |first1=Frank S. |last2=Kawano |first2=Shoichi |title=''Pennellianthus'' (Scrophulariaceae)--A New Genus of Japan and USSR |url=https://www.jstor.org/stable/2423949 |journal=The American Midland Naturalist |access-date=11 November 2021 |pages=358–367 |doi=10.2307/2423949 |date=1970|volume=83 |issue=2 |jstor=2423949 |url-access=subscription }}</ref> * ''Penstemon'' Schmidel * ''Tonella'' Nutt. ex A.Gray * ''Uroskinnera'' Lindl.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2287 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Cheloneae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052557/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2287 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Digitalideae * ''Digitalis'' L. * ''Erinus'' L.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2285 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Digitalideae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052554/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2285 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Globularieae * ''Campylanthus'' Roth * ''Globularia'' L. * ''Poskea'' Vatke<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2292 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Globularieae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052609/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2292 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Gratioleae * ''Adenosma'' R.Br. * ''Bacopa'' Aubl. * ''Benjaminia'' Mart. ex Benj. * ''Boelckea'' Rossow * ''Cheilophyllum'' Pennell ex Britton * ''Chodaphyton'' {{small|Minod}} * ''Darcya'' B.L.Turner & C.C.Cowan * ''Deinostema'' T.Yamaz. * ''Dizygostemon'' (Benth.) Radlk. ex Wettst. * ''Dopatrium'' Buch.-Ham. ex Benth. * ''Fonkia'' Phil. * ''Geochorda'' Cham. & Schltdl. * ''Gratiola'' L. * ''Hydrotriche'' Zucc. * ''Ildefonsia'' Gardner * ''Lapaea'' {{small|Scatigna & V.C.Souza}} * ''Leucospora'' Nutt. * ''Limnophila'' R.Br. * ''Matourea'' Aubl. * ''Mecardonia'' Ruiz & Pav. * ''Philcoxia'' P.Taylor & V.C.Souza * ''Schistophragma'' Benth. ex Endl. * ''Scoparia'' L. * ''Stemodia'' L. * ''Tetraulacium'' Turcz.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2289 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Gratioleae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052601/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2289 |archive-date=2015-09-24 |url-status=dead }}</ref> *''Umbraria'' {{small|Scatigna & V.C.Souza}} | ; Tribe Hemiphragmeae * ''Hemiphragma'' Wall.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2293 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Hemiphragmeae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052611/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2293 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Plantagineae * ''Aragoa'' Kunth * ''Littorella'' P.J.Bergius * ''Plantago'' L.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2286 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Plantagineae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052555/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2286 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Russelieae * ''Russelia'' Jacq. * ''Tetranema'' Benth.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2288 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Russelieae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052559/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2288 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Sibthorpieae * ''Ellisiophyllum'' Maxim. * ''Sibthorpia'' L.<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2291 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Sipthorpieae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052608/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2291 |archive-date=2015-09-24 |url-status=dead }}</ref> ; Tribe Veroniceae * ''Kashmiria'' D.Y.Hong * ''Lagotis'' Gaertn. * ''Neopicrorhiza'' D.Y.Hong * ''Paederota'' L. * ''Picrorhiza'' Royle ex Benth. * ''Scrofella'' Maxim. * ''Veronica'' L. * ''Veronicastrum'' Heist. ex Fabr. * ''Wulfenia'' Jacq. * ''Wulfeniopsis'' D.Y.Hong<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2284 |title=GRIN Genera of ''Plantaginaceae'' tribe ''Veroniceae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20150924052549/http://www.ars-grin.gov/cgi-bin/npgs/html/gnlist.pl?2284 |archive-date=2015-09-24 |url-status=dead }}</ref>

; Tribe unknown *''Agathelpis'' {{small|Choisy}} *''Anamaria'' {{small|V.C.Souza}} *''Ancistrostylis'' {{small|T.Yamaz.}} *''Bythophyton'' Hook.f. *''Dintera'' Stapf *''Encopella'' Pennell *''Lafuentea'' Lag. *''Trapella'' {{small|Oliv.}} *''Triaenophora'' {{small|Soler.}} *''Trungboa'' {{small|Rauschert}} |}

Although GRIN includes ''Lafuentea'' Lag. in the tribe Antirrhineae,<ref name="GRIN"/> in the phylogenetic analysis of Fernández-Mazuecos ''et al.'' (2013)<ref>{{cite journal |jstor=10.1086/668790 |first1=Mario |last1=Fernández-Mazuecos |first2=José Luis |last2=Blanco-Pastor |first3=Pablo |last3=Vargas |title=A Phylogeny of Toadflaxes (''Linaria'' Mill.) Based on Nuclear Internal Transcribed Spacer Sequences: Systematic and Evolutionary Consequences |journal=International Journal of Plant Sciences |volume=174 |issue=2 |date=February 2013 |pages=234–249 |doi=10.1086/668790|bibcode=2013IJPlS.174..234F |s2cid=85302392 }}</ref> it was a sister to the Antirrhineae, as also noted by Albach (2005).<ref name="Albach-2005"/> For the time being it should be considered an outgroup.

===Excluded genera=== {{div col|colwidth=28em}} * ''Artanema'' D.Don → Linderniaceae * ''Chamaegigas'' Dinter ex Heil → Linderniaceae * ''Craterostigma'' Hochst. → Linderniaceae * ''Limosella'' L. → Scrophulariaceae * ''Lindenbergia'' Lehm. → Orobanchaceae * ''Lindernia'' All. → Linderniaceae * ''Micranthemum'' Michx. → Linderniaceae * ''Microcarpaea'' R.Br. → Phrymaceae * ''Picria'' Lour. → Linderniaceae * ''Rehmannia'' Libosch. ex Fisch. & C.A.Mey. → Orobanchaceae * ''Torenia'' L. → Linderniaceae<ref>{{cite web |url=http://www.ars-grin.gov/cgi-bin/npgs/html/gnothlist.pl?888 |title=GRIN genera sometimes placed in ''Plantaginaceae'' |work=Germplasm Resources Information Network |access-date=2011-04-28 |archive-url=https://web.archive.org/web/20041118031328/http://www.ars-grin.gov/cgi-bin/npgs/html/gnothlist.pl?888 |archive-date=2004-11-18 |url-status=dead }}</ref> {{div col end}}

==References== {{Reflist|3}} * {{cite journal | author = Olmstead, R. G., dePamphilis, C. W., Wolfe, A. D., Young, N. D., Elisons, W. J. & Reeves P. A. | year = 2001 | title = Disintegration of the Scrophulariaceae | journal = American Journal of Botany | volume = 88 | pages = 348–361 | doi = 10.2307/2657024 | pmid = 11222255 | issue = 2 | jstor = 2657024| bibcode = 2001AmJB...88..348O | doi-access = free }}

==Bibliography== * [http://www.rjb.csic.es/jardinbotanico/ficheros/documentos/pdf/pubinv/PVG/Vargas_al_2004_PSE_Antirrhineae.pdf 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.]

==External links== * {{Commons category-inline}} * {{Wikispecies-inline}} * [http://www.mobot.org/MOBOT/research/APweb/welcome.html Treatment of Plantaginaceae in MOBOT]

Category:Plantaginaceae Category:Lamiales families