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

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

s in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...

Lamiales The order Lamiales (also known as the mint order) are an order in the asterid group of dicotyledonous flowering plants. It includes about 23,810 species, 1,059 genera, and is divided into about 25 families. These families include Acanthaceae, Big ...

.Peter F. Stevens (2001 onwards). "Phrymaceae" At:

Angiosperm Phylogeny Website The Angiosperm Phylogeny Website (or APweb) is a website dedicated to research on angiosperm phylogeny and taxonomy. The site is hosted by the Missouri Botanical Garden website and maintained by researchers, Peter F. Stevens and Hilary M. Davis ...

. 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 philanthropist Henry Shaw. Its herbarium, with more than 6.6 million ...

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

cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The extr ...

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

Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...

, the other in western

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...

. Members of this family occur in diverse

habitat In ecology, the term habitat summarises the array of resources, physical and 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 ...

s, including

desert A desert is a barren area of landscape where little precipitation occurs and, consequently, living conditions are hostile for plant and animal life. The lack of vegetation exposes the unprotected surface of the ground to denudation. About on ...

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 ...

s. Phrymaceae is a family of mostly

herbs In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicina ...

and a few subshrubs, bearing tubular,

bilaterally symmetric Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pl ...

flowers. They can be annuals or

perennials A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also widel ...

. Some of the Australian

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

are aquatic or semiaquatic. One of these, ''Glossostigma'', is among the smallest of flowering plants, larger than the aquatic ''

Lemna ''Lemna'' is a genus of free-floating aquatic plants referred to by the common name "duckweed". They are morphologically divergent members of the arum family Araceae. These rapidly growing plants have found uses as a model system for studies in ...

'' but similar in size to the

terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...

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

centimeter 330px, Different lengths as in respect to the Electromagnetic spectrum, measured by the Metre and its deriveds scales. The Microwave are in-between 1 meter to 1 millimeter. A centimetre (international spelling) or centimeter (American spellin ...

s long, while the largest are

wood Wood is a porous and fibrous structural tissue found in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulose fibers that are strong in tension and embedded in a matrix of lignin ...

shrub A shrub (often also called a 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 tree ...

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. Reproduction is a fundamental feature of all known life; each individual or ...

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 mate under which circumstances. Recognised ...

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 can be a usefu ...

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

, or mixed mating. Some are

pollinate Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...

d by

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

s, others by

hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With about 361 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are ...

s. The most common

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

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 A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...

s, but exceptions exist such as an

achene An achene (; ), also sometimes called akene and occasionally achenium or achenocarp, is a type of simple dry fruit produced by many species of flowering plants. Achenes are monocarpellate (formed from one carpel) and indehiscent (they do not o ...

, 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 t ...

'', 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 are strawberries, rasp ...

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

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

are in cultivation.

Anthony Huxley Anthony Julian Huxley (2 December 1920 – 26 December 1992) was a British botanist. He edited ''Amateur Gardening'' from 1967 to 1971, and was vice-president of the Royal Horticultural Society in 1991. He was the son of Julian Huxley. He was ...

, 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 ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...

Mimulus Mimulus is a plant genus in the family Phrymaceae, which was traditionally placed in family Scrophulariaceae. The genus now contains only seven species, two native to eastern North America and the other five native to Asia, Australia, Africa, ...

'' when it was defined

broadly Vice Media Group LLC is an American-Canadian digital media and broadcasting company. , the Vice Media Group included five main business areas: VICE.com (digital content); VICE STUDIOS (film and TV production) VICE TV (also known as VICELAND); V ...

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 Group epithets must conform to the ''Interna ...

, 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 contrast ...

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

evolutionary Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variati ...

studies Study or studies may refer to: General * Education ** Higher education * Clinical trial * Experiment * Observational study * Research * Study skills, abilities and approaches applied to learning Other * Study (art), a drawing or series of d ...

.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 Anonymity describes situations where the acting person's identity is unknown. Some writers have argued that namelessness, though technically correct, does not capture what is more centrally at stake in contexts of anonymity. The important idea he ...

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...

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 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 fu ...

. 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 ...

.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 "unispe ...

with ''Phryma leptostachya'' as its only

. It was limited in

geographic Geography (from Greek: , ''geographia''. Combination of Greek words ‘Geo’ (The Earth) and ‘Graphien’ (to describe), literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, a ...

range to eastern North America and eastern

China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...

. ''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. The ...

. Research on

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...

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 for ...

''.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'' and '' Lancea'' were included in Phrymaceae for a short time before further studies indicated that they, along with Dodartia 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 (; ) 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 typically shared derived char ...

analyses of

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...

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 posterior p ...

assessments of clade support in any of the datasets 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 vari ...

. 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 of these 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, rags, grasses or other vegetable sources in water, draining the water through fine mesh leaving the fibre evenly distribu ...

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 available to the general public.Berne Conve ...

. 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