Gesneriaceae, the gesneriad family, is a

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. They include all forbs (flowering plants without a woody stem), grasses and grass-like plants, a vast majority of ...

s consisting of about 152 genera and ca. 3,540 species in the tropics and subtropics of the

Old World The "Old World" is a term for Afro-Eurasia that originated in Europe , after Europeans became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia, which were previously thought of by th ...

(almost all

Didymocarpoideae The Didymocarpoideae are a subfamily of plants in the family Gesneriaceae. It was formerly the subfamily Cyrtandroideae. This subfamily consists mostly of tropical and subtropical Old World genera, found in Africa, Asia and the Pacific. One spe ...

) and the

New World The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. ...

(most

Gesnerioideae The Gesnerioideae are a subfamily of plants in the family Gesneriaceae: based on the type genus ''Gesneria''. Although genera typically originate in the New World, some species have become widely distributed as ornamental plants. Description Ges ...

), with a very small number extending to temperate areas. Many species have colorful and showy flowers and are cultivated as ornamental plants.

Etymology

The family name is based on the genus ''

Gesneria ''Gesneria'' is a genus of approximately 50 species in the flowering plant family Gesneriaceae. Except for two or three odd South American species, all are native to islands of the Caribbean. The genus is classified in the tribe Gesnerieae alo ...

'', which honours

Swiss Swiss may refer to: * the adjectival form of Switzerland * Swiss people Places * Swiss, Missouri *Swiss, North Carolina * Swiss, West Virginia * Swiss, Wisconsin Other uses * Swiss-system tournament, in various games and sports *Swiss Internati ...

naturalist and

humanist Humanism is a philosophical stance that emphasizes the individual and social potential and agency of human beings. It considers human beings the starting point for serious moral and philosophical inquiry. The meaning of the term "humani ...

Conrad Gessner Conrad Gessner (; la, Conradus Gesnerus 26 March 1516 – 13 December 1565) was a Swiss physician, naturalist, bibliographer, and philologist. Born into a poor family in Zürich, Switzerland, his father and teachers quickly realised his ta ...

Description

Most species are

herbaceous Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition ...

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 wide ...

subshrub A subshrub ( Latin ''suffrutex'') or dwarf shrub is a short shrub, and is a woody plant. Prostrate shrub is a related term. "Subshrub" is often used interchangeably with "bush".Jackson, Benjamin, Daydon; A Glossary of Botanic Terms with their ...

s but a few are woody shrubs or small

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

s. The

phyllotaxy In botany, phyllotaxis () or phyllotaxy is the arrangement of leaves on a plant stem. Phyllotactic spirals form a distinctive class of patterns in nature. Leaf arrangement The basic arrangements of leaves on a stem are opposite and alterna ...

is usually opposite and

decussate Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term chiasma is named aft ...

, but leaves have a spiral or alternate arrangement in some groups. As with other members of the

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, B ...

the flowers have a (usually)

zygomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spiral ...

corolla whose petals are fused into a tube and there is no one character that separates a gesneriad from any other member of Lamiales. Gesneriads differ from related families of the Lamiales in having an unusual inflorescence structure, the "pair-flowered cyme", but some gesneriads lack this characteristic, and some other Lamiales (

Calceolariaceae Calceolariaceae is a family of flowering plants in the order Lamiales that has been recently segregated from Scrophulariaceae. The family includes three genera, ''Calceolaria'', '' Porodittia'', and '' Jovellana'', but analysis suggests that the ...

and some

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 S ...

) share it. The ovary can be superior, half-inferior or fully inferior, and the fruit a dry or fleshy capsule 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 ...

. The

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 are always small and numerous. Gesneriaceae have traditionally been separated from Scrophulariaceae by having a

unilocular A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...

rather than bilocular ovary, with parietal rather than axile placentation.

Taxonomy

"Gesneriaceae" is a

conserved name A conserved name or ''nomen conservandum'' (plural ''nomina conservanda'', abbreviated as ''nom. cons.'') is a scientific name that has specific nomenclatural protection. That is, the name is retained, even though it violates one or more rules whic ...

(''nom. cons.''), meaning that although alternative, less well used names for the family were published earlier, the

International Code of Nomenclature for algae, fungi, and plants The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "trad ...

specifies this as the name to be used. It was published by

Louis Claude Richard Louis Claude Marie Richard (19 September 1754 – 6 June 1821) was a French botanist and botanical illustrator. Richard was born at Versailles. Between 1781 and 1789 he collected botanical specimens in Central America and the West Indies. On his ...

and

Antoine Laurent de Jussieu Antoine Laurent de Jussieu (; 12 April 1748 – 17 September 1836) was a French botanist, notable as the first to publish a natural classification of flowering plants; much of his system remains in use today. His classification was based on an ...

in 1816. In 1829,

Barthélemy Dumortier Barthélemy, or Barthélémy is a French name, a cognate of Bartholomew. Notable people with this name include: Given name * Barthélemy (explorer), French youth who accompanied the explorer de La Salle in 1687 * Barthélémy Bisengimana, Cong ...

divided the family into two tribes, based on the number of stamens. However, the only genus he placed in his two-stamen tribe, ''

Columellia ''Columellia'' is a group of plant species in the Columelliaceae described as a genus in 1794. ''Columellia'' is native to the Andes Mountains of western South America ( Colombia, Ecuador, Peru, Bolivia).Serrano, M. & J. Terán. 1998 000 Tr ...

'', is now placed in the separate family

Columelliaceae Columelliaceae is a family of trees and shrubs native to the Andes of South America. In the APG II taxonomy it is placed in the order Lamiales, but a 2008 study suggested that the family is sister to the Bruniaceae, and the Angiosperm Phylogeny ...

. Dumortier's publication has been treated as the first for the family by some sources. Botanists who have made significant contributions to the systematics of the family are

George Bentham George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studi ...

, Robert Brown, B.L. Burtt,

C.B. Clarke Charles Baron Clarke (17 June 1832 – 25 August 1906) was a British botanist. He was born at Andover, the eldest son of Turner Poulter Clarke. He was educated at King's College School, London, and at Trinity and Queens' Colleges, Cambridge ...

Olive Mary Hilliard Olive Mary Hilliard ( Hillary, 4 July 1925 – 30 November 2022) was a South African botanist and taxonomist. Hilliard authored 372 land plant species names, the fifth-highest number of such names authored by any female scientist. Hilliard was bo ...

Joseph Dalton Hooker Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of ...

, William Jackson Hooker,

Karl Fritsch Karl Fritsch (24 February 1864 – 17 January 1934) was an Austrian botanist. He was born in Vienna and educated mainly at the University of Vienna, obtaining his PhD degree in 1886 and his Habilitation Habilitation is the highest univ ...

Elmer Drew Merrill Elmer Drew Merrill (October 15, 1876 – February 25, 1956) was an American botanist and taxonomist. He spent more than twenty years in the Philippines where he became a recognized authority on the flora of the Asia-Pacific region. Through ...

Harold E. Moore, Jr. Harold Emery Moore, Jr. (July 7, 1917 – October 27, 1980) was an American botanist especially known for his work on the systematics of the palm family. He served as Director of the L. H. Bailey Hortorium at Cornell University, and was appoint ...

John L. Clark John is a common English name and surname: * John (given name) * John (surname) John may also refer to: New Testament Works * Gospel of John, a title often shortened to John * First Epistle of John, often shortened to 1 John * Seco ...

Conrad Vernon Morton Conrad Vernon Morton (24 October 1905 – 29 July 1972) was an American botanist who did notable writings on Ferns. He was also a specialist in Gesneriaceae and Solanaceae for the Smithsonian Institution from 1928. In 1938, botanists Standl. & ...

Henry Nicholas Ridley Henry Nicholas Ridley CMG (1911), MA (Oxon), FRS, FLS, F.R.H.S. (10 December 1855 – 24 October 1956) was an English botanist, geologist and naturalist who lived much of his life in Singapore. He was instrumental in promoting rubber trees i ...

, Laurence Skog,

W.T. Wang WT or Wt may refer to: * Weight People * Watertender, a former petty officer rating in the United States Navy * Weapons technician, a former petty officer rating in the United States Navy * Robert Wight (1796–1872), in botanical taxonomy, a ...

Anton Weber Anton may refer to: People *Anton (given name), including a list of people with the given name *Anton (surname) Places *Anton Municipality, Bulgaria **Anton, Sofia Province, a village *Antón District, Panama **Antón, a town and capital of th ...

, and

Hans Wiehler Hans Joachim Wiehler (8 July 1930 in Klettendorf – 2003) was a German botanist who specialized in the plant family Gesneriaceae. In 1954 he received a Bachelor of Arts degree from Goshen College in Goshen, Indiana and a Bachelor of Divinity de ...

. The

Gesneriad Society The Gesneriad Society, Inc., is an international horticultural society devoted to the promotion, cultivation, and study of gesneriads (plant family Gesneriaceae). The organization was founded in 1951 as the American Gloxinia Society by Elvin Mc ...

is an international horticultural society devoted to the promotion, cultivation, and study of Gesneriaceae.

Phylogeny

From about 1997 onwards,

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 led to extensive changes in the classification of the family Gesneriaceae and its genera, many of which have been re-circumscribed or synonymized. New species are still being discovered, particularly in Asia, and may further change generic boundaries. A consensus phylogeny used to build classifications of the family in 2013 and 2020 is shown below (to the level of tribes). The family

is shown as the sister to Gesneriaceae. A

phylogenomic Phylogenomics is the intersection of the fields of evolution and genomics. The term has been used in multiple ways to refer to analysis that involves genome data and evolutionary reconstructions. It is a group of techniques within the larger fields ...

study published in 2021 which used 418 nuclear genes confirmed the

monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...

of all the subfamilies and tribes. It resolved ''Peltanthera'' as sister to a clade of Calceolariaceae and Gesneriaceae. Within the Gesnerioideae, Napeantheae rather than Titanotricheae was found to be sister to the remaining tribes. The position of Titanotricheae varied according to the method used to build the cladogram, which the authors suggested was due to

incomplete lineage sorting Incomplete lineage sorting, also termed hemiplasy, deep coalescence, retention of ancestral polymorphism, or trans-species polymorphism, describes a phenomenon in population genetics when ancestral gene copies fail to coalesce (looking backwards i ...

following rapid divergence. The phylogenetic position of ''Titanotrichum'' remains unsettled. The genus ''Sanango'' has not always been included in Gesneriaceae. However, molecular phylogenetic studies published up to and including 2021 suggest that it does belong in the family as the most basal member, and it is placed in its own subfamily. The studies also show the genus ''

Peltanthera ''Peltanthera'' is a genus of flowering plants containing a single species, ''Peltanthera floribunda''. The genus was originally placed in family Loganiaceae and has since been variously placed in Buddlejaceae, Scrophulariaceae, Gesneriaceae ...

'' to be outside the family, although some sources still place it within the Gesneriaceae. The genus ''

Rehmannia ''Rehmannia'' is a genus of seven species of flowering plants in the order Lamiales and family Orobanchaceae, endemic to China. It has been placed as the only member of the monotypic tribe Rehmannieae, but molecular phylogenetic studies suggest ...

'' has also sometimes been included in the family but is now referred to the family

Orobanchaceae Orobanchaceae, the broomrapes, is a family of mostly parasitic plants of the order Lamiales, with about 90 genera and more than 2000 species. Many of these genera (e.g., '' Pedicularis'', '' Rhinanthus'', '' Striga'') were formerly included in ...

. No single morphological feature absolutely divides two main subfamilies (i.e. forms a uniform

synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...

). Gesnerioideae seedlings have normal

cotyledon A cotyledon (; ; ; , gen. (), ) is a significant part of the embryo within the seed of a plant, and is defined as "the embryonic leaf in seed-bearing plants, one or more of which are the first to appear from a germinating seed." The num ...

s of the same size and shape (isocotylous). The cotyledons of Didymocarpoideae are usually, but not always, eventually different in size and shape (anisocotylous). One cotyledon ceases to grow and withers away, while the other continues to grow, and may even form a very large leaf that is the only one the plant has ('' Monophyllaea'', some ''

Streptocarpus ''Streptocarpus'' ("twisted fruit" from Greek στρεπτός (''streptos'') "twisted" and καρπός (''carpos'') "fruit") is an Afrotropical genus of flowering plants in the family Gesneriaceae. The genus is native to Afromontane biotopes ...

''). Gesnerioideae flowers usually have four fertile

stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...

s, rarely two or five. Didymocarpoideae flowers usually have two fertile stamens, less often four, rarely one or five.

Subfamilies and genera

On the basis of

, morphological and biogeographical differences, the family has been divided into two major subfamilies: subfamily

(formerly Cyrtandroideae) with all but one species from the

, and subfamily

native from the Americas west through the Pacific to Australia and southeastern China. The genus ''Sanango'' is placed in its own subfamily, Sanangoideae. The two main subfamilies are further divided into tribes and subtribes. Genera accepted by

Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants ...

(PoWO) are listed below, together with their placement in a subfamily and tribe by Weber ''et al.'' (2020). Three genera are listed by PoWO but not by Weber ''et al.'': '' Coptocheile'' Hoffmanns. (doubtfully placed in Gesneriaceae), ''Parakohleria'' Wiehler (now included in ''

Pearcea ''Pearcea'' is a genus of tropical herbaceous plants in the family Gesneriaceae native to western South America. It is classified in tribe Gloxinieae and is closely related to the genus ''Kohleria'', in which some of its species were previously i ...

'') and ''

'' Benth. (excluded from Gesneriaceae by molecular phylogenetic studies).

Ecology

About half of the New World species (i.e. the subfamily Gesnerioideae) are co-adapted to

bird pollination Ornithophily or bird pollination is the pollination of flowering plants by birds. This sometimes (but not always) coevolutionary association is derived from insect pollination (entomophily) and is particularly well developed in some parts of the w ...

, particularly by

hummingbird Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics ar ...

s in the Americas. Bird-pollinated species typically have two-lipped flowers in shades of red; examples are found in the genera ''

Asteranthera ''Asteranthera'' is a monotypic plant genus in the family Gesneriaceae, native to the humid forests of Argentina and Chile. The sole species in the genus, ''Asteranthera ovata'', is an evergreen scrambling vine A vine ( Latin ''vīnea'' "gr ...

'', ''

Columnea ''Columnea'' is a genus of around 200 species of epiphytic herbs and shrubs in the family Gesneriaceae, native to the tropics of the Americas and the Caribbean. The tubular or oddly shaped flowers are usually large and brightly colored – ...

'' and ''

Sinningia ''Sinningia'' is a genus of flowering plants in the family Gesneriaceae. It is named after Wilhelm Sinning (1792–1874), a gardener of the Botanische Gärten der Friedrich-Wilhelms-Universität Bonn. There are about 65 species of tuberous herba ...

''. Among Old World genera, ''

Aeschynanthus ''Aeschynanthus'' is a genus of about 150 species of evergreen subtropical and tropical plants in the family Gesneriaceae. They are usually trailing epiphytes with brightly colored flowers that are pollinated by sunbirds. The genus name comes f ...

'' has similar flowers. File:Estrellita del Monte Parque Nacional Hornopirén 02.jpg, ''

Asteranthera ovata ''Asteranthera'' is a monotypic plant genus in the family Gesneriaceae, native to the humid forests of Argentina and Chile. The sole species in the genus, ''Asteranthera ovata'', is an evergreen scrambling vine A vine ( Latin ''vīnea'' "gr ...

'' File:Columnea microphylla (14604475109).jpg, '' Columnea microphylla'' File:Sinningia sceptrum — João de Deus Medeiros 001.jpg, ''

Sinningia sceptrum ''Sinningia'' is a genus of flowering plants in the family Gesneriaceae. It is named after Wilhelm Sinning (1792–1874), a gardener of the Botanische Gärten der Friedrich-Wilhelms-Universität Bonn. There are about 65 species of tuberous herba ...

'' File:IMG 7323-Aeschynanthus speciosus.jpg, '' Aeschynanthus speciosus''

Cultivation

Some genera in the family are grown as

ornamental plant Ornamental plants or garden plants are plants that are primarily grown for their beauty but also for qualities such as scent or how they shape physical space. Many flowering plants and garden varieties tend to be specially bred cultivars that ...

s, both as garden plants and as

houseplant A houseplant, sometimes known as a pot plant, potted plant, or an indoor plant, is an ornamental plant that is grown indoors. As such, they are found in places like residences and offices, mainly for decorative purposes. Common houseplants are us ...

s. Such genera include: ''

'', ''

Achimenes ''Achimenes'' is a genus of about 25 species of tropical and subtropical rhizomatous perennial herbs in the flowering plant family Gesneriaceae. They have a multitude of common names such as magic flowers, widow's tears, Cupid's bower, or hot wat ...

'', ''

Haberlea ''Haberlea'' is a monotypic genus of flowering plants in the family Gesneriaceae. The only member of this genus, ''Haberlea rhodopensis'', is endemic to parts of Bulgaria and a small part of northern Greece, especially in the Rhodope Mountains. C ...

'', ''

Nematanthus ''Nematanthus'' is a genus of flowering plants in the family Gesneriaceae. All of its species are endemic to Brazil. Compared to other gesneriads, ''Nematanthus'' has leaves that are small, succulent, and hard-surfaced. The plant has a trailing ...

'' (syn. ''Hypocyrta''), '' Ramonda'', and ''

'' (Cape primroses, African violets). One of the most familiar members of the family to gardeners are the African violets in ''Streptocarpus'' section ''Saintpaulia''. Gesneriads are divided culturally into three groups on the basis of whether, and how, their stems are modified into storage organs:

rhizomatous In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow h ...

tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growin ...

ous, and "fibrous-rooted", meaning those that lack such storage structures (although all gesneriads have fibrous roots).

References

External links

World Checklist of GesneriaceaeGenera of GesneriaceaeGesneriad Reference Web
(on the Gesneriad Reference Web)
Gesneriaceae
i
Flora of ChinaThe Gesneriad Society
(formerly the American Gloxinia and Gesneriad Society)
Annotated Bibliography of the GesneriaceaePhylogenetic relationships in the Gesnerioideae (Gesneriaceae) based on nrDNA ITS and cpDNA trnL-F and trnE-T spacer region sequences
(link to abstract)
Evolution of Cyrtandra (Gesneriaceae) in the Pacific Ocean: the origin of a supertramp cladeWeber, A. 2004. Gesneriaceae and Scrophulariaceae: Robert Brown and now. ''Telopea'' 10(2): 543-571.Gesneriaceae: All you need to know about gesneriads.
{{Taxonbar, from=Q156686 Lamiales families