Phrymaceae, also known as the lopseed family, is a small

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Ancient Greek, Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed with ...

s in the

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

Lamiales The Lamiales (also known as the mint order) are an order of flowering plants in the asterids clade of the Eudicots. Under the APG IV system of flowering plant classification the order consists of 24 families, and includes about 23,810 species ...

.Peter F. Stevens (2001 onwards). "Phrymaceae" At: Angiosperm Phylogeny Website. At: Botanical Databases At:

Missouri Botanical Garden The Missouri Botanical Garden is a botanical garden located at 4344 Shaw Boulevard in St. Louis, Missouri. It is also known informally as Shaw's Garden for founder and philanthropy, philanthropist Henry Shaw (philanthropist), Henry Shaw. I ...

Website. (see ''External links'' below) It has a nearly

cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...

, but is concentrated in two centers of diversity, one in

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

, the other in western

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

. Members of this family occur in diverse

habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...

s, including

desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...

s, river banks and

mountain A mountain is an elevated portion of the Earth's crust, generally with steep sides that show significant exposed bedrock. Although definitions vary, a mountain may differ from a plateau in having a limited summit area, and is usually higher t ...

s. Phrymaceae is a family of mostly

herbs Herbs are a widely distributed and widespread group of plants, excluding vegetables, with savory or aromatic properties that are used for flavoring and garnish (food), garnishing food, for medicinal purposes, or for fragrances. Culinary use typi ...

and a few

subshrub A subshrub (Latin ''suffrutex'') or undershrub is either a small shrub (e.g. prostrate shrubs) or a perennial that is largely herbaceous but slightly woody at the base (e.g. garden pink and florist's chrysanthemum). The term is often interch ...

s, bearing tubular, bilaterally symmetric flowers. They can be annuals or

perennials In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

. Some of the Australian

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

are 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 '' Lepuropetalon''. The smallest members of Phrymaceae are only a few

centimeter upright=1.35, Different lengths as in respect to the electromagnetic spectrum, measured by the metre and its derived scales. The microwave is in-between 1 meter to 1 millimeter. A centimetre (International spelling) or centimeter (American ...

s long, while the largest are

wood Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...

shrub A shrub or bush is a small to medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees by their multiple ...

s to 4 m tall. The floral structure of Phrymaceae is variable, to such an extent that a morphological assessment is difficult.

Reproduction Reproduction (or procreation or breeding) is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: Asexual reproduction, asexual and Sexual ...

is also variable, being brought about by different

mating system A mating system is a way in which a group is structured in relation to sexual behaviour. The precise meaning depends upon the context. With respect to animals, the term describes which males and females mating, mate under which circumstances. Reco ...

s which may be sexual or asexual, and may involve

outcrossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing in animals Out ...

self-fertilization Autogamy or self-fertilization refers to the Cell fusion, fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a Reproduction, reproductive mechanism employed by many flowering pl ...

, or mixed mating. Some are pollinated by

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

s, others by

hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...

s. The most common

fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...

type in this family is a

dehiscent Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part. Structures that op ...

capsule containing numerous

seed In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...

s, but exceptions exist such as an

achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple fruit, simple dry fruits, dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and Dehi ...

, in ''

Phryma leptostachya ''Phryma leptostachya'', or lopseed, is a perennial herb of the genus ''Phryma''. When distinguished from '' Phryma oblongifolia'' and ''Phryma nana'', it is native to eastern North America. The plant stands about 0.3 to 1.0 meters tall, and th ...

'', or a

berry A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone or pit although many pips or seeds may be present. Common examples of berries in the cul ...

-like fruit in ''Leucocarpus''. About 16

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

are in cultivation.

Anthony Huxley Anthony Julian Huxley (2 December 1920 – 26 December 1992) was a British botanist and writer. An elected council member of the Royal Horticultural Society, he became its vice president in 1991. He edited '' Amateur Gardening'' from 1967 to 19 ...

, Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. (set). They are known horticulturally as "Mimulus" and were formerly placed in the

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

Mimulus Mimulus , also known as monkeyflowers, is a plant genus in the family (biology), family Phrymaceae, which was traditionally placed in family Scrophulariaceae. The genus now contains only seven species, two native to eastern North America and th ...

'' when it was

defined A definition is a statement of the meaning of a term (a word, phrase, or other set of symbols). Definitions can be classified into two large categories: intensional definitions (which try to give the sense of a term), and extensional definitio ...

broadly Vice Media Group LLC is a Canadian-American digital media and broadcasting company. Vice Media encompasses four main business areas: Vice Studios Group (film and TV production); Vice TV (a joint venture with A&E Networks, also known as Vicelan ...

to include about 150 species. ''Mimulus'', as a

botanical name A botanical name is a formal scientific name conforming to the ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) and, if it concerns a plant cultigen, the additional cultivar or cultivar group, Group epithets must conform t ...

, rather than a

common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...

or horticultural name, now represents a genus of only seven species. Most of its former species have been transferred to ''Diplacus'' or ''Erythranthe''. Six of the horticultural species are of special importance. These are ''

Diplacus aurantiacus ''Diplacus'' is a genus of flowering plants in the family Phrymaceae, which was traditionally placed in family Scrophulariaceae. It includes 49 species native to the western United States and northwestern Mexico. Most prefer dry and rocky areas. ...

, Diplacus puniceus,

Erythranthe cardinalis ''Erythranthe cardinalis'', the scarlet monkeyflower, is a flowering perennial in the family Phrymaceae. Together with other species in ''Mimulus'' section ''Erythranthe'', it serves as a model system for studying pollinator-based reproductive is ...

, Erythranthe guttata, Erythranthe lutea,'' and '' Erythranthe cuprea''. Phrymaceae has recently become a model system for

evolutionary Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certa ...

studies.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. . (See ''External links'' below). Within the order Lamiales, Phrymaceae is a

member Member may refer to: * Military jury, referred to as "Members" in military jargon * Element (mathematics), an object that belongs to a mathematical set * In object-oriented programming, a member of a class ** Field (computer science), entries in ...

of an unnamed

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

of five families.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. (See ''External links'' below). This clade has the topology of a phylogenetic grade and can therefore be represented as .Nancy F. Refulio-Rodriguez and Richard G. Olmstead. 2014. "Phylogeny of Lamiidae". ''American Journal of Botany'' 101(2):287-299. Two of these families,

Mazaceae Mazaceae is a family of plants in the order Lamiales. The family was described by James L. Reveal in 2011. Genera in this family were most recently previously included in Phrymaceae and in older classifications were placed in Scrophulariaceae. ...

and Rehmanniaceae are not part of the

APG III system The APG III system of flowering plant classification is the third version of a modern, mostly molecular-based, system of plant taxonomy being developed by the Angiosperm Phylogeny Group (APG). Published in 2009, it was superseded in 2016 by a f ...

. They were not formally validated until 2011.James L. Reveal.2011. page 47. In: "Summary of recent systems of angiosperm classification". ''Kew Bulletin'' 66(1):5-48. The composition of Phrymaceae and the delimitation of genera changed radically from 2002 to 2012 as a result of

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

studies.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. . (See ''External links'' below).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. 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. Previously, Phrymaceae had been

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...

with ''Phryma leptostachya'' as its only

. It was limited in

geographic Geography (from Ancient Greek ; combining 'Earth' and 'write', literally 'Earth writing') is the study of the lands, features, inhabitants, and phenomena of Earth. Geography is an all-encompassing discipline that seeks an understanding o ...

range Range may refer to: Geography * Range (geographic), a chain of hills or mountains; a somewhat linear, complex mountainous or hilly area (cordillera, sierra) ** Mountain range, a group of mountains bordered by lowlands * Range, a term used to i ...

to eastern North America and eastern

China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...

. ''Phryma'' had been previously placed by Cronquist in

Verbenaceae The Verbenaceae ( ), the verbena family or vervain family, is a family of mainly tropical flowering plants. It contains trees, shrubs, and herbs notable for heads, spikes, or clusters of small flowers, many of which have an aromatic smell. T ...

. Research on

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

relationships revealed that several genera, traditionally included in

Scrophulariaceae The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scr ...

, were actually more closely related to ''Phryma'' than to ''

Scrophularia The genus ''Scrophularia'' of the family Scrophulariaceae comprises about 200 species of herbaceous flowering plants commonly known as figworts. Species of ''Scrophularia'' all share square stems, opposite leaves and open two-lipped flowers fo ...

''.Bengt Oxelman, Per Kornhall, Richard G. Olmstead & Birgitta Bremer. 2005. "Further disintegration of the Scrophulariaceaea". ''Taxon'' 54(2): 411-425. These genera became part of an expanded Phrymaceae. ''

Mazus ''Mazus'' is a genus of low-growing perennial plants. It has been placed in various plant families including Phrymaceae, Scrophulariaceae, and recently in the family Mazaceae. page 47 Consisting of around 40 species, this genus is generally fo ...

'' and '' Lancea'' were included in Phrymaceae for a short time before further studies indicated that they, along with

Dodartia ''Dodartia'' is a genus of flowering plants in the family Mazaceae. It has only one currently accepted species, ''Dodartia orientalis'', native to Ukraine, Russia, Anatolia, the Caucasus, Central Asia, Iran, Afghanistan, Pakistan, the Altai, nort ...

should be segregated as a new family, Mazaceae. As currently understood, Phrymaceae consists of about 210 species in 13 genera. ''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

(about 130 species) and

(about 30 species). Phrymaceae consists of four

s, all of which have strong statistical support in

cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

analyses of

DNA sequence A nucleic acid sequence is a succession of bases within the nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. This succession is denoted by a series of a set of five different letters that indicate the order of the nu ...

s. No relationships among these four clades have been strongly supported by the bootstrap or

posterior probability The posterior probability is a type of conditional probability that results from updating the prior probability with information summarized by the likelihood via an application of Bayes' rule. From an epistemological perspective, the posteri ...

assessments of clade support in any of the

dataset A data set (or dataset) is a collection of data. In the case of tabular data, a data set corresponds to one or more database tables, where every column of a table represents a particular variable, and each row corresponds to a given record o ...

s that have been produced so far. One of the four main clades consists of a single species, ''

''. Another consists of ''Mimulus''

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...

(seven species) and six genera that have an Australian

distribution Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations *Probability distribution, the probability of a particular value or value range of a varia ...

. The other two clades have an American-Asian

disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...

.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. One of these includes the large genus ''Diplacus'', while the other includes the other large genus, ''Erythranthe''.

Estimate Estimation (or estimating) is the process of finding an estimate or approximation, which is a value that is usable for some purpose even if input data may be incomplete, uncertain, or unstable. The value is nonetheless usable because it is de ...

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 segregates of ''Mimulus'', the number of species assigned to ''Mimulus'' sensu lato has ranged from about 90David J. Mabberley. 2008. ''Mabberley's Plant-Book'' third edition (2008). Cambridge University Press: UK. . to about 150.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. A 2008

paper Paper is a thin sheet material produced by mechanically or chemically processing cellulose fibres derived from wood, Textile, rags, poaceae, grasses, Feces#Other uses, herbivore dung, or other vegetable sources in water. Once the water is dra ...

indicates that the actual number of species is well over 150. In 2012, a revision of Phrymaceae recognized 188 species in the family, but noted that 17 species from Australia and five from North America would be named and described in future

publications To publish is to make Content (media), content available to the general public.Berne Conv ...

. Ten of those unnamed species will be in '' Peplidium'', raising the number of species in that genus from four to 14. Illustration Mimulus guttatus0

Description

The following description is

excerpt {{Short pages monitor