| Lopseed family | |
|---|---|
 | |
| Erythranthe tilingii | |
Scientific classification | |
| Kingdom: | Plantae |
| Clade: | Tracheophytes |
| Clade: | Angiosperms |
| Clade: | Eudicots |
| Clade: | Asterids |
| Order: | Lamiales |
| Family: | Phrymaceae Schauer |
| Genera | |
About 11; see text. | |
Phrymaceae, also known as the lopseed family, is a smallfamily offlowering plants in theorderLamiales.[1] It has a nearlycosmopolitan distribution, but is concentrated in twocenters of diversity, one inAustralia, the other in westernNorth America.[2] Members of this family occur in diversehabitats, includingdeserts,river banks andmountains.
Phrymaceae is a family of mostlyherbs and a fewsubshrubs, bearing tubular,bilaterally symmetric flowers. They can beannuals orperennials.[2] Some of the Australiangenera areaquatic or semiaquatic. One of these,Glossostigma, is among the smallest of flowering plants, larger than the aquaticLemna but similar in size to theterrestrialLepuropetalon. The smallest members of Phrymaceae are only a fewcentimeters long, while the largest arewoodyshrubs to 4m tall. The floral structure of Phrymaceae is variable, to such an extent that amorphological assessment is difficult.Reproduction is also variable, being brought about by differentmating systems which may besexual orasexual, and may involveoutcrossing,self-fertilization, ormixed mating. Some arepollinated byinsects, others byhummingbirds. The most commonfruit type in this family is adehiscent capsule containing numerousseeds, but exceptions exist such as anachene, inPhryma leptostachya, or aberry-like fruit inLeucocarpus.
About 16species are incultivation.[3] They are knownhorticulturally as "Mimulus" and were formerlyplaced in thegenusMimulus when it wasdefinedbroadly to include about 150 species.Mimulus, as abotanical name, rather than acommon name orhorticultural name, now represents a genus of only seven species. Most of its former species have been transferred toDiplacus orErythranthe.[2] Six of the horticultural species are of special importance. These areDiplacus aurantiacus,Diplacus puniceus,Erythranthe cardinalis,Erythranthe guttata,Erythranthe lutea, andErythranthe cuprea.
Phrymaceae has recently become amodel system forevolutionarystudies.[4]
Within the order Lamiales, Phrymaceae is amember of anunnamedclade of five families.[5] This clade has the topology of aphylogenetic grade and can therefore be represented as {Mazaceae [Phrymaceae (Paulowniaceae <Orobanchaceae + Lamiaceae>)]}.[6] Two of these families,Mazaceae andRehmanniaceae are not part of theAPG III system.[7] They were not formallyvalidated until 2011.[8]
The composition of Phrymaceae and the delimitation of genera changed radically from 2002 to 2012 as a result ofmolecular phylogeneticstudies.[9][10][11] Previously, Phrymaceae had beenmonotypic withPhryma leptostachya as its onlyspecies. It was limited ingeographicrange to eastern North America and easternChina.Phryma had been previously placed byCronquist inVerbenaceae. Research onphylogeneticrelationships revealed that several genera, traditionally included inScrophulariaceae, were actually moreclosely related toPhryma than toScrophularia.[12] These genera became part of anexpanded Phrymaceae.Mazus andLancea were included in Phrymaceae for a short time before further studies indicated that they, along withDodartia should besegregated as a new family, Mazaceae.
As currently understood, Phrymaceae consists of about 210 species in 13 genera.[2]Erythranthe (111 species) andDiplacus (46 species) are much larger than the other genera. Phrymaceae is distributed nearly worldwide but with the majority of species in westernNorth America (about 130 species) andAustralia (about 30 species). Phrymaceae consists of fourclades, all of which have strongstatistical support incladistic analyses ofDNA sequences. No relationships among these four clades have been strongly supported by thebootstrap orposterior probability assessments of clade support in any of thedatasets that have been produced so far. One of the four main clades consists of a single species,Phryma leptostachya. Another consists ofMimulussensu stricto (seven species) and six genera that have an Australiandistribution. The other two clades have an American-Asiandisjunct distribution.[13] One of these includes the large genusDiplacus, while the other of these includes the other large genus,Erythranthe.
Estimates of the number of species in Phrymaceae have varied widely because of a lack of clear differences between species in certain genera, especiallyDiplacus andErythranthe. When these two genera have been treated assegregates ofMimulus, the number of species assigned toMimulus sensu lato has ranged from about 90[14] to about 150.[15] A 2008paper indicates that the actual number of species is well over 150.[4]
In 2012, arevision of Phrymaceae recognized 188 species in the family, but noted that 17 species from Australia and five from North America would benamed anddescribed in futurepublications. Ten of those unnamed species will be inPeplidium, raising the number of species in that genus from four to 14.[2]
The following description isexcerpted from the conspectus published in 2012.[2]
Mostlyannual orperennialherbs, a fewsubshrubs.Leavesopposite, sometimesglandular-punctate.Flowershypogynous; usually inracemes, rarely solitary or inaxillary clusters of 2 or 3.Calyx tubular, toothed, usually ribbed below teeth.Corollazygomorphic, or rarely, sub-actinomorphic; 5-lobed, or rarely reduced to 3 or 4 lobes.Stamens 4,didynamous, or rarely 2.Filamentsinserted on corolla tube.Carpels 2, bearing manyovules; or rarely, carpel 1 with a single ovule.Stigmas 2-lobed,sensitive except inElacholoma hornii which has alinear stigma.Fruit aloculicidalcapsule, rarely aschizocarp orberry; borne in apersistent calyx.Seeds small; many or only 1 (Phryma).
The family Phrymaceae is mainly defined by the following threecharacteristics:
The family Phrymaceae was established in 1847 byJohannes Conrad Schauer as a guest author in theProdromus ofAlphonse Pyramus de Candolle.[16][17] The family has often been called "Phrymataceae", even in modern times,[18] but thecorrect name for the family is Phrymaceae.[2]
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 familyVerbenaceae, but sometimes inLamiaceae.[19]Mimulus and its relatives were usually placed in some version ofScrophulariaceae that was much larger than the currently accepted circumscription of that family.[20]
In 2002, amolecular phylogenetic study showed thatPhryma formed a strongly supported clade withMimulus and its various relatives.ChloroplastDNA showedPhryma to beembedded within a broadly definedMimulus, but this result was not strongly supported, and was contradicted bydata from theITS andETS regions of thenuclear genome.[9]
In 2004, in the most recent comprehensivetreatment of families and genera in Lamiales, Phrymaceae consisted ofPhryma only.[21] In that treatment, it was suggested thatMimulus 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" wereDodartia,Mazus,Lancea,Bythophyton,Encopella,Hemianthus,Micranthemum,Bryodes,Dintera,Psammetes, andMimulicalyx.[15]
Dodartia,Mazus, andLancea make Phrymaceae paraphyletic if they are included within it.[6][22] They now constitute the related family Mazaceae.[8]
Themonotypic generaBythophyton andEncopella might properly belong toPlantaginaceaetribeGratioliae.[23] Thishypothesis has never beentested by molecular phylogenetics.
Hemianthus is so similar toMicranthemum that its recognition as a separate genus is doubtful.[14]Micranthemum andBryodes have been shown to be members ofLinderniaceae.[5]
TheAfricanmonotypic generaDintera andPsammetes are little known and theiraffinities remain obscure.Mimulicalyx has 2 species, bothendemic toChina. Their familial placement remains uncertain.
ThusBythophyton, Encopella, Dintera, Psammetes, andMimulicalyx 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 ofAPG III, some authors have chosen to maintain a largepolyphyletic Scrophulariaceae until there is a clear understanding of how it should be "disintegrated".[19]
Ataxonomicconspectus of Phrymaceae published in 2012 included 13 genera. In that conspectus,Eunanus,Tricholoma, andBerendtiella were not accepted as they are in some recent works.Eunanus is reduced to asection inDiplacus.Tricholoma is subsumed withinGlossostigma.[2] Subsequentmolecular phylogenetic studies showed thatCyrtandromoea also belonged in the family,[24] a placement accepted byPlants of the World Online, which also accepts the monospecific genusMimulicalyx, producing a total of 15 genera.[25]
