Difference between revisions of "Phrymaceae"

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{{taxobox
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#redirect [[:Category:Phrymaceae]]
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|name = Lopseed family
 
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|image = Mimulus tilingii 23134.JPG
 
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|image_caption = ''[[Mimulus tilingii]]''
 
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|regnum = [[Plantae]]
 
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|unranked_divisio = [[Angiosperms]]
 
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|unranked_classis = [[Eudicots]]
 
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|unranked_ordo = [[Asterids]]
 
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|ordo = [[Lamiales]]
 
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|familia = '''Phrymaceae'''
 
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|familia_authority = [[Johannes Conrad Schauer|Schauer]]
 
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|subdivision_ranks = Genera
 
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|subdivision = About 11; see text.
 
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|}}
 
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'''Phrymaceae''', also known as the lopseed family, is a small [[Family (biology)|family]] of [[flowering plant]]s in the [[Order (biology)|order]] [[Lamiales]].<ref name="apweb">Peter F. Stevens (2001 onwards). "Phrymaceae" At: [[Angiosperm Phylogeny Website]]. At: Botanical Databases At: [[Missouri Botanical Garden]] Website. (see ''External links'' below)</ref> It has a nearly [[cosmopolitan distribution]], but is concentrated in two [[Center of diversity|centers of diversity]], one in [[Australia]], the other in western [[North America]].<ref name="barker2012">{{cite journal | last1 = Barker| first1 =  W. L. (Bill)| last2 = Nesom| first2 =  Guy L. | last3 = Beardsley| first3 =  Paul M. | last4 = Fraga| first4 =  Naomi S.| year= 2012| title =A Taxonomic Conspectus of Phyrmaceae: A Narrowed Circumscription for ''Mimulus'', New and Resurrected Genera, and New Names and Combinations| url = http://www.phytoneuron.net/PhytoN-Phrymaceae.pdf | journal = Phytoneuron | volume =39 | issue = | pages = 1–60 | issn=2153-733X}}</ref>  Members of this family occur in diverse [[habitat]]s, including [[desert]]s, [[river bank]]s and [[mountain]]s.
 
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Phrymaceae is a family of mostly [[Herbaceous plant|herbs]] and a few [[subshrub]]s, bearing tubular, [[bilaterally symmetric]] flowers. They can be [[annual plant|annuals]] or [[perennial plant|perennials]].<ref name=barker2012/> Some of the Australian [[genera]] are [[Aquatic plant|aquatic]] or semiaquatic. One of these, ''Glossostigma'', is among the smallest of flowering plants, larger than the aquatic ''[[Lemna]]'' but similar in size to the [[Terrestrial plant|terrestrial]] ''[[Lepuropetalon]]''. The smallest members of Phrymaceae are only a few [[centimeter]]s long, while the largest are [[wood]]y [[shrub]]s to 4 [[Meter|m]] tall. The floral structure of Phrymaceae is variable, to such an extent that a [[Plant morphology|morphological]] assessment is difficult. [[Reproduction]] is also variable, being brought about by different [[mating system]]s which may be [[Sexual reproduction|sexual]] or [[Asexual reproduction|asexual]], and may involve [[outcrossing]], [[Autogamy|self-fertilization]], or [[Mixed mating model|mixed mating]]. Some are [[pollinate]]d by [[insect]]s, others by [[hummingbird]]s. The most common [[fruit]] type in this family is a [[dehiscent]] capsule containing numerous [[seed]]s, but exceptions exist such as an [[achene]], in ''[[Phryma|Phryma leptostachya]]'', or a [[berry (botany)|berry]]-like fruit in ''Leucocarpus''.
 
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About 16 [[species]] are in [[Gardening|cultivation]].<ref name="rhs">[[Anthony Huxley]], Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. {{ISBN|978-0-333-47494-5}} (set).</ref> They are known [[Horticulture|horticulturally]] as "Mimulus" and were formerly [[Biological classification|placed]] in the [[genus]] ''[[Mimulus]]'' when it was [[Circumscription (taxonomy)|defined]] [[sensu lato|broadly]] to include about 150 species. ''Mimulus'', as a [[botanical name]], rather than a [[common name]] or [[Cultivated plant taxonomy|horticultural name]], now represents a genus of only seven species. Most of its former species have been transferred to ''Diplacus'' or ''Erythranthe''.<ref name=barker2012/> Six of the horticultural species are of special importance. These are ''[[Mimulus aurantiacus|Diplacus aurantiacus]], [[Diplacus puniceus]], [[Mimulus cardinalis|Erythranthe cardinalis]], [[Mimulus guttatus|Erythranthe guttata]], [[Erythranthe lutea]],'' and ''[[Erythranthe cuprea]]''.
 
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Phrymaceae has recently become a [[Conceptual model|model system]] for [[Introduction to evolution|evolutionary]] [[Research|studies]].<ref name=wu2008>Carrie A. Wu, David B. Lowry, Arielle M. Cooley, Kevin M. Wright, Y.W. Lee, and John H. Willis. 2008. "Mimulus is an emerging model system for the integration of ecological and genomic studies". ''Heredity'' 100(2):220-230. {{doi|10.1038/sj.hdy.6801018}}. (See ''External links'' below).</ref>
 
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Within the order Lamiales, Phrymaceae is a [[Element (mathematics)|member]] of an [[Undescribed taxon|unnamed]] [[clade]] of five families.<ref name=schaferhoff2010>Bastian Schäferhoff, Andreas Fleischmann, Eberhard Fischer, Dick C. Albach, Thomas Borsch, Günther Heubl, and Kai F. Müller. 2010. "Towards resolving Lamiales relationships: insights from rapidly evolving chloroplast sequences". BioMed Central Evolutionary Biology 10:352. {{doi|10.1186/1471-2148-10-352}} (See ''External links'' below).</ref> This clade has the topology of a [[phylogenetic grade]] and can therefore be represented as {Mazaceae [Phrymaceae (Paulowniaceae <Orobanchaceae + Lamiaceae>)]}.<ref name=refrod2014>Nancy F. Refulio-Rodriguez and Richard G. Olmstead. 2014. "Phylogeny of Lamiidae". ''American Journal of Botany'' 101(2):287-299. {{doi|10.3732/ajb.1300394}}</ref> Two of these families, [[Mazaceae]] and [[Rehmanniaceae]] are not part of the [[APG III system]].<ref name=APGIII2009>{{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 |accessdate=2013-07-06 |doi=10.1111/j.1095-8339.2009.00996.x }}</ref> They were not formally [[valid publication|validated]] until 2011.<ref name=reveal2011>James L. Reveal.2011. page 47. In: "Summary of recent systems of angiosperm classification". ''Kew Bulletin'' '''66'''(1):5-48.</ref>
 
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The composition of Phrymaceae and the delimitation of genera changed radically from 2002 to 2012 as a result of [[molecular phylogenetic]] [[Research|studies]].<ref name=beardsley2002>Paul M. Beardsley and Richard G. Olmstead. 2002. "Redefining Phrymaceae: the placement of ''Mimulus'', tribe Mimuleae, and ''Phryma''". ''American Journal of Botany'' '''89'''(7):1093-1102. {{doi|10.3732/ajb.89.7.1093}}. (See ''External links'' below).</ref><ref name=beardsley2004>Paul M. Beardsley, Steve E. Schoenig, Justen B. Whittall, and Richard G. Olmstead. 2004. "Patterns of evolution in western North American ''Mimulus'' (Phrymaceae)". ''American Journal of Botany'' '''91'''(3):474-489. {{doi|10.3732/ajb.91.3.474}}</ref><ref name=beardsley2005>Paul M. Beardsley and William R. Barker. 2005. "Patterns of evolution in Australian ''Mimulus'' and related genera (Phrymaceae ~ Scrophulariaceae): a molecular phylogeny using chloroplast and nuclear sequence data". ''Australian Systematic Botany'' '''18'''(1):61-73. {{doi|10.1071/SB04034}}</ref> Previously, Phrymaceae had been [[monotypic]] with ''Phryma leptostachya'' as its only [[species]]. It was limited in [[Biogeography|geographic]] [[Range (biology)|range]] to eastern North America and eastern [[China]]. ''Phryma'' had been previously placed by [[Arthur Cronquist|Cronquist]] in [[Verbenaceae]]. Research on [[phylogeny|phylogenetic]] [[Plant taxonomy|relationships]] revealed that several genera, traditionally included in [[Scrophulariaceae]], were actually more [[Affinity (taxonomy)|closely related]] to ''Phryma'' than to ''[[Scrophularia]]''.<ref name=oxelman2005>Bengt Oxelman, Per Kornhall, Richard G. Olmstead & Birgitta Bremer. 2005. "Further disintegration of the Scrophulariaceaea". ''Taxon'' '''54'''(2): 411-425.</ref> These genera became part of an [[sensu lato|expanded]] Phrymaceae. ''[[Mazus]]'' and ''[[Lancea (plant)|Lancea]]'' were included in Phrymaceae for a short time before further studies indicated that they, along with [[Dodartia]] should be [[Segregate (taxonomy)|segregated]] as a new family, Mazaceae.
 
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As currently understood, Phrymaceae consists of about 210 species in 13 genera.<ref name=barker2012/> ''Erythranthe'' (111 species) and ''Diplacus'' (46 species) are much larger than the other genera. Phrymaceae is distributed nearly worldwide but with the majority of species in western [[North America]] (about 130 species) and [[Australia]] (about 30 species). Phrymaceae consists of four [[clade]]s, all of which have strong [[Resampling (statistics)|statistical support]] in [[cladistic]] analyses of [[DNA sequence]]s. No relationships among these four clades have been strongly supported by the [[Bootstrapping (statistics)|bootstrap]] or [[posterior probability]] assessments of clade support in any of the [[dataset]]s that have been produced so far. One of the four main clades consists of a single species, ''[[Phryma leptostachya]]''. Another consists of ''Mimulus'' [[sensu stricto]] (seven species) and six genera that have an Australian [[Species distribution|distribution]]. The other two clades have an American-Asian [[disjunct distribution]].<ref name=wen2010>Jun Wen, Stephanie M. Ickert-Bond, Ze-Long Nie, and Rong Li. 2010. "Timing and modes of evolution of eastern Asian - North American biogeographic disjunctions in seed plants". In: Long, M., Gu, H. and Zhou, Z., Darwin's Heritage Today : Proceedings of the Darwin 2010 Beijing International Conference. Beijing: Higher Education Press, pp.252-269.</ref> One of these includes the large genus ''Diplacus'', while the other of these includes the other large genus, ''Erythranthe''.
 
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[[Estimate]]s of the number of species in Phrymaceae have varied widely because of a lack of clear differences between species in certain genera, especially ''Diplacus'' and ''Erythranthe''. When these two genera have been treated as [[Segregate (taxonomy)|segregates]] of ''Mimulus'', the number of species assigned to ''Mimulus'' sensu lato has ranged from about 90<ref name="mabberley2008">David J. Mabberley. 2008. ''Mabberley's Plant-Book'' third edition (2008). Cambridge University Press: UK. {{ISBN|978-0-521-82071-4}}.</ref> to about 150.<ref name=fischer2004>Eberhard Fischer. 2004. pages 401-405. In: "Scrophulariaceae" pages 333-432. In: Klaus Kubitzki (editor) and Joachim W. Kadereit (volume editor). ''The Families and Genera of Vascular Plants'' volume VII. Springer-Verlag: Berlin; Heidelberg, Germany. {{ISBN|978-3-540-40593-1}}</ref> A 2008 [[Scientific paper|paper]] indicates that the actual number of species is well over 150.<ref name=wu2008/>
 
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In 2012, a [[taxonomic revision|revision]] of Phrymaceae recognized 188 species in the family, but noted that 17 species from Australia and five from North America would be [[Neologism|named]] and [[diagnosis (taxonomy)|described]] in future [[Academic publishing|publications]]. Ten of those unnamed species will be in ''Peplidium'', raising the number of species in that genus from four to 14.<ref name=barker2012/>
 
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[[File:Illustration Mimulus guttatus0.jpg|thumb|right|''Mimulus guttatus'' from Thomé, ''Flora von Deutschland, Österreich und der Schweiz'' 1885]]
 
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== Genera ==
 
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The following genera are from a [[Plant taxonomy|taxonomic]] [[conspectus]] of Phrymaceae published in 2012.<ref name=barker2012/> In that conspectus, ''Eunanus'', ''Tricholoma'', and ''Berendtiella'' were not accepted as they are in some recent works. ''Eunanus'' is reduced to a [[Section (botany)|section]] in ''Diplacus''. ''Tricholoma'' is subsumed within ''Glossostigma''.
 
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<!--''[[genus name]]'' {{Au|author,date}} (synonyms) &ndash; common name &ndash; number of species, range -->
 
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{|
 
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|- valign=top
 
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* ''[[Diplacus]]'' {{Au|[[Thomas Nuttall|Nuttall]], 1838}}
 
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* ''[[Elacholoma]]'' {{Au|[[Ferdinand von Mueller|F.Mueller]] & [[Ralph Tate|Tate]], 1895}} (the stigma lobes are relatively long and are receptive over most of their length)
 
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* ''[[Erythranthe]]'' {{Au|[[Édouard Spach|Spach]], 1840}}
 
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* ''[[Glossostigma]]'' {{Au|[[Robert Wight|Wight]] & [[George Arnott Walker-Arnott|Arnott]], 1836}} (with three or four lobes in the calyx instead five; contains one large and one vestigial stigma lobe)
 
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* ''[[Hemichaena]]'' {{Au|[[George Bentham|Bentham]], 1841}}
 
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* ''[[Leucocarpus]]'' {{Au|[[David Don|D.Don]], 1830}}
 
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* ''[[Microcarpaea]]'' {{Au|[[Robert Brown (Scottish botanist from Montrose)|Robert Brown]], 1810}}
 
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* ''[[Mimetanthe]]'' {{Au|[[Edward Lee Greene|Greene]], 1885}}
 
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* ''[[Mimulus]]'' {{Au|[[Linnaeus|L.]], 1753}}
 
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* ''[[Peplidium]]'' {{Au|[[Alire Raffeneau Delile|Delile]], 1813}}  (contains one large and one vestigial stigma lobe)
 
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* ''[[Phryma]]'' {{Au|L., 1753}}
 
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* ''[[Thyridia (plant)|Thyridia]]'' {{Au|W.R.Barker & Beardsley, 2012}}
 
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* ''[[Uvedalia]]'' {{Au|[[Robert Brown (Scottish botanist from Montrose)|Robert Brown]], 1810}}
 
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|}
 
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== Description ==
 
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The following description is [[excerpt]]ed from the conspectus published in 2012.<ref name=barker2012/>
 
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Mostly [[Annual plant|annual]] or [[Perennial plant|perennial]] [[Herbaceous plant|herbs]], a few [[subshrub]]s. [[Leaf|Leaves]] [[Phyllotaxis|opposite]], sometimes [[Gland (botany)|glandular]]-[[Glossary of botanical terms#punctate|punctate]]. [[Flower]]s [[inferior ovary|hypogynous]]; usually in [[raceme]]s, rarely solitary or in [[Glossary of botanical terms#axil|axillary]] clusters of 2 or 3. [[Calyx (botany)|Calyx]] tubular, toothed, usually ribbed below teeth. [[Corolla (flower)|Corolla]] [[Floral symmetry|zygomorphic]], or rarely, sub-actinomorphic; 5-[[Glossary of botanical terms#lobe|lobed]], or rarely reduced to 3 or 4 lobes. [[Stamen]]s 4, [[Glossary of botanical terms#didynamy|didynamous]], or rarely 2. [[Androecium|Filaments]] [[Glossary of botanical terms#inserted|inserted]] on corolla tube. [[Carpel]]s 2, bearing many [[ovule]]s; or rarely, carpel 1 with a single ovule. [[Stigma (botany)|Stigmas]] 2-lobed, [[Glossary of botanical terms#sensitive|sensitive]] except in ''[[Elacholoma hornii]]'' which has a [[Glossary of botanical terms#linear|linear]] stigma. [[Fruit]] a [[Dehiscence (botany)|loculicidal]] [[Capsule (fruit)|capsule]], rarely a [[schizocarp]] or [[berry (botany)|berry]]; borne in a [[Glossary of botanical terms#persistent|persistent]] calyx. [[Seed]]s small; many or only 1 (''Phryma'').
 
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The family Phrymaceae is mainly defined by the following three [[Phenotypic trait|characteristics]]:  
 
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* Tubular, toothed [[sepal|calyces]] (with five [[Glossary of botanical terms#lobe|lobes]]).
 
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* [[Stigma (botany)|Stigmas]] with two [[Glossary of botanical terms#lamella|lamellas]] that have [[Glossary of botanical terms#sensitive|sensitive]] inner surfaces, that close together on contact with a [[pollinator]].
 
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* [[Capsule (fruit)|Capsules]] that are [[Locule|loculicidally]] [[Dehiscence (botany)|dehiscent]], splitting between the partitions of the [[Ovary (botany)|ovary]].
 
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== History ==
 
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The family Phrymaceae was established in 1847 by [[Johannes Conrad Schauer]] as a guest author in the ''[[Prodromus Systematis Naturalis Regni Vegetabilis|Prodromus]]'' of [[Alphonse Pyramus de Candolle]].<ref name="ipniphrymaceae">''Phrymaceae'' in [[International Plant Names Index]]. (see ''External links'' below).</ref><ref name=schauer1847>Johannes Conrad Schauer. 1847. "Phrymaceae" pages 520-521. In: Alphonse Pyramus de Candolle (editor). ''Prodromus Systematis Naturalis Regni Vegetabilis [...]'' volume 11.  Sumptibus Victoris Masson. Paris, France. (See ''External links'' below).</ref> The family has often been called "Phrymataceae", even in modern times,<ref name="takhtajan2009">Armen L. Takhtajan (Takhtadzhian). ''Flowering Plants'' second edition (2009). page 557. Springer Science+Business Media. {{ISBN|978-1-4020-9608-2}}. {{ISBN|978-1-4020-9609-9}} {{doi|10.1007/978-1-4020-9609-9}}</ref> but the [[Correct name (botany)|correct name]] for the family is Phrymaceae.<ref name=barker2012/>
 
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Until 2002, Phrymaceae was usually defined as consisting of only a single, anomalous species, ''Phryma leptostachya''. Whenever Phrymaceae was not recognized, ''Phryma'' was usually placed in the family [[Verbenaceae]], but sometimes in [[Lamiaceae]].<ref name="heywood2007">Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada. (2007). {{ISBN|978-1-55407-206-4}}.</ref> ''Mimulus'' and its relatives were usually placed in some version of [[Scrophulariaceae]] that was much larger than the currently accepted circumscription of that family.<ref name=tank2006>David C. Tank, Paul M. Beardsley, Scot A. Kelchner, and Richard G. Olmstead. 2006. "Review of the systematics of Scrophulariaceae s.l. and their current disposition". ''Australian Systematic Botany'' '''19'''(6):289-307. {{doi|10.1071/SB05009}} (See ''External links'' below).</ref>
 
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In 2002, a [[molecular phylogenetic]] study showed that ''Phryma'' formed a strongly supported clade with ''Mimulus'' and its various relatives. [[Chloroplast]] [[DNA]] showed ''Phryma'' to be [[Paraphyly|embedded]] within a broadly defined ''Mimulus'', but this result was not strongly supported, and was contradicted by [[data]] from the [[Internal transcribed spacer|ITS]] and [[External transcribed spacer|ETS]] regions of the [[nuclear genome]].<ref name=beardsley2002/>
 
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In 2004, in the most recent comprehensive [[Treatise|treatment]] of families and genera in Lamiales, Phrymaceae consisted of ''Phryma'' only.<ref name=cantino2004>Philip D. Cantino. 2004. "Phrymaceae". pages 323-326. In: Klaus Kubitzki (editor) and Joachim W. Kadereit (volume editor). ''The Families and Genera of Vascular Plants'' volume VII. Springer-Verlag: Berlin; Heidelberg, Germany. {{ISBN|978-3-540-40593-1}}</ref> In that treatment, it was suggested that ''Mimulus'' and its relatives (8 genera) might be transferred from Scrophulariaceae to Phrymaceae. It was also suggested that 11 other genera in Scrophulariaceae might be transferred in the same way. The 11 "additional genera" were ''[[Dodartia]], [[Mazus]], [[Lancea (plant)|Lancea]], [[Bythophyton]], [[Encopella]], [[Hemianthus]], [[Micranthemum]], [[Bryodes]], [[Dintera]], [[Psammetes]],'' and ''[[Mimulicalyx]]''.<ref name=fischer2004/>
 
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''Dodartia'', ''Mazus'', and ''Lancea'' make Phrymaceae paraphyletic if they are included within it.<ref name=refrod2014/><ref name=albach2009>Dirk C. Albach, Kun Yan, Søren R. Jensen, and Hong-Qing Li. 2009. "Phylogenetic placement of ''Triaenophora'' (formerly Scrophulariaceae) with some implications for the phylogeny of Lamiales". ''Taxon'' '''58'''(3):749-756.</ref> They now constitute the related family Mazaceae.<ref name=reveal2011/>
 
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The [[Monotypic taxon|monotypic]] genera [[Bythophyton]] and [[Encopella]] might properly belong to [[Plantaginaceae]] [[Tribe (biology)|tribe]] [[Gratioliae]].<ref name=albach2005>Dirk C. Albach, Heidi M. Meudt, and Bengt Oxelman. 2005. "Piecing together the "new" Plantaginaceae". ''American Journal of Botany'' 92(2):297-315. {{doi|10.3732/ajb.92.2.297}}</ref> This [[hypothesis]] has never been [[Hypothesis testing|tested]] by molecular phylogenetics.
 
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''Hemianthus'' is so similar to ''Micranthemum'' that its recognition as a separate genus is doubtful.<ref name="mabberley2008"/> ''Micranthemum'' and ''Bryodes'' have been shown to be members of [[Linderniaceae]].<ref name=schaferhoff2010/>
 
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The [[Africa]]n [[monotypic]] genera ''Dintera'' and ''Psammetes'' are little known and their [[Affinity (taxonomy)|affinities]] remain obscure. ''Mimulicalyx'' has 2 species, both [[Endemism|endemic]] to [[China]]. Their familial placement remains uncertain.
 
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Thus ''Bythophyton, Encopella, Dintera, Psammetes,'' and ''Mimulicalyx'' might be considered as possible members of Phrymaceae since they have not been unequivocally placed elsewhere. Instead of recognizing Phrymaceae and several of the other Lamiales families of [[APG III]], some authors have chosen to maintain a large [[polyphyletic]] Scrophulariaceae until there is a clear understanding of how it should be "disintegrated".<ref name="heywood2007"/>
 
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== References ==
 
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{{reflist}}
 
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== External links ==
 
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{{commons category|Phrymaceae}}
 
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* [http://www.mobot.org/MOBOT/Research/APweb/maps/phrymaceae.gif Distribution map]
 
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* [http://biology.duke.edu/willislab/pub_pdfs/Wuetal07Heredity.pdf ''Mimulus'' as a model system]
 
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* [http://www.nature.com/hdy/journal/v100/n2/pdf/6801018a.pdf ''Mimulus'' as a model system]
 
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* [http://www.plantsystematics.org/reveal/pbio/fam/MagnoliidaeOnLine.html classification scheme for extant flowering plants]
 
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* [http://www.biomedcentral.com/content/pdf/1471-2148-10-352.pdf Lamiales relationships (pdf)]
 
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* [http://www.amjbot.org/cgi/content/full/89/7/1093 Beardsley et alii.2002. AJB]
 
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* [http://www.amjbot.org/content/91/3/474.full Beardsley et alii.2004. AJB]
 
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* [http://www.ipni.org/ipni/advPlantNameSearch.do?find_family=Phrymaceae+&find_genus=&find_species=&find_infrafamily=&find_infragenus=&find_infraspecies=&find_authorAbbrev=&find_includePublicationAuthors=on&find_includePublicationAuthors=off&find_includeBasionymAuthors=on&find_includeBasionymAuthors=off&find_publicationTitle=&find_isAPNIRecord=on&find_isAPNIRecord=false&find_isGCIRecord=on&find_isGCIRecord=false&find_isIKRecord=on&find_isIKRecord=false&find_rankToReturn=all&output_format=normal&find_sortByFamily=on&find_sortByFamily=off&query_type=by_query&back_page=plantsearch Phrymaceae]
 
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* [http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=6163 Phrymaceae (page 520)]
 
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* [http://phylodiversity.net/dtank/Tank_Lab/Publications_files/Aust.%20Syst.%20Bot.%202006%20Tank.pdf Review of the systematics of Scrophulariaceae s.l.]
 
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* [http://www.tolweb.org/Lamiales/20726 Lamiales]
 
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{{taxonbar}}
 
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[[Category:Phrymaceae| 01]]
 
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[[Category:Lamiales families]]
 

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