Gesneriaceae, the gesneriad family, is a

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 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 coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...

(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 Tropics, tropical and Subtropics, subtropical Old World genera, found in Africa, Asia and ...

) and the

New World The term "New World" is used to describe the majority of lands of Earth's Western Hemisphere, particularly the Americas, and sometimes Oceania."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: ...

(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 flowering plants in the family Gesneriaceae. It contains 62 species which are native to islands of the Caribbean. The genus is classified in the tribe Gesnerieae along with the genera '' Bellonia'', '' Pheidonocarpa'', ...

'', which honours

Swiss Swiss most commonly refers to: * the adjectival form of Switzerland * Swiss people Swiss may also refer to: Places * Swiss, Missouri * Swiss, North Carolina * Swiss, West Virginia * Swiss, Wisconsin Other uses * Swiss Café, an old café located ...

naturalist Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...

and

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

Conrad Gessner Conrad Gessner (; ; 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 talents and supported him t ...

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 of ...

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 ...

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 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, e.g., including only woody plants with secondary growth, only ...

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 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 ...

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 spir ...

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 th ...

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 Scr ...

) 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 of berries in the cul ...

. The

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

(''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 or ICNafp) 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 tho ...

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 illustration, botanical illustrator. Biography Richard was born at Versailles (city), Versailles. Between 1781 and 1789 he collected botanical s ...

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 e ...

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'', 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 studie ...

Robert Brown Robert Brown may refer to: Robert Brown (born 1965), British Director, Animator and author Entertainers and artists * Washboard Sam or Robert Brown (1910–1966), American musician and singer * Robert W. Brown (1917–2009), American printmaker ...

B.L. Burtt BL (or similar) may refer to: Arts and entertainment * Boys' love, a Japanese term for fiction featuring romantic relationships between male characters * BL Publishing, a division of the wargames manufacturing company, Games Workshop * ''Boston ...

, C.B. Clarke,

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 b ...

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 20 years he served as director of the Ro ...

William Jackson Hooker Sir William Jackson Hooker (6 July 178512 August 1865) was an English botany, botanist and botanical illustrator, who became the first director of Kew Gardens, Kew when in 1841 it was recommended to be placed under state ownership as a botan ...

Karl Fritsch Karl Fritsch (24 February 1864 – 17 January 1934) was an Austrian botanist. He was a specialist on the Gesneriaceae and the taxonomy of monocots. Biography Fritsch was born in Vienna, the son of meteorologist Karl Fritsch, and educated ...

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,

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. & S ...

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,

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

, and

Hans Wiehler Hans Joachim Wiehler (8 July 1930 in Klecie, Pomeranian Voivodeship, 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 ...

. The Gesneriad Society 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...

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 (ILS) (also referred to as hemiplasy, deep coalescence, retention of ancestral polymorphism, or trans-species polymorphism) is a phenomenon in evolutionary biology and population genetics that results in discordance bet ...

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, or ...

'' 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, which is native to China and Japan. It has been placed as the only member of the monotypic tribe Rehmannieae, but molecular phylogenetic ...

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

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

. 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 Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...

). Gesnerioideae seedlings have normal

cotyledon A cotyledon ( ; ; "a cavity, small cup, any cup-shaped hollow", gen. (), ) is a "seed leaf" – a significant part of the embryo within the seed of a plant – and is formally defined as "the embryonic leaf in seed-bearing plants, one or mor ...

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 ''Monophyllaea'' is a genus of plants in the family Gesneriaceae. All the species only have one leaf. Species 38 species are accepted. * ''Monophyllaea albicalyx'' * ''Monophyllaea andersonii'' * ''Monophyllaea anthocrena'' * ''Monophyllae ...

'', some ''

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

''). Gesnerioideae flowers usually have four fertile

stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...

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 taxonomic database published by the Royal Botanic Gardens, Kew. History Following the Convention on Biological Diversity, the Royal Botanic Gardens in Kew launched Plants of the World Online i ...

(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. (now treated as a synonym of ''Sinningia''), ''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 ...

'') 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 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 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 temperate forests of Argentina and Chile. The sole species in the genus, ''Asteranthera ovata'', is an evergreen scrambling vine A vine is any plan ...

'', ''

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 – us ...

'' 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 her ...

''. Among Old World genera, ''

Aeschynanthus ''Aeschynanthus'' is a genus of about 150 species of evergreen subtropics, subtropical and tropics, tropical plants in the family (biology), family Gesneriaceae. They are usually trailing epiphytes with brightly colored flowers that are pollinate ...

'' 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 temperate forests of Argentina and Chile. The sole species in the genus, ''Asteranthera ovata'', is an evergreen scrambling vine A vine is any plan ...

'' File:Columnea microphylla (14604475109).jpg, '' Columnea microphylla'' File:Sinningia sceptrum — João de Deus Medeiros 001.jpg, '' Sinningia sceptrum'' File:IMG 7323-Aeschynanthus speciosus.jpg, ''

Aeschynanthus speciosus ''Aeschynanthus speciosus'', also known as the basket plant, is a species of flowering plant in the family Gesneriaceae. There are many species of ''Aeschynanthus'', all of which have long, trailing stems and bright flowers. The large, fleshy, d ...

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 th ...

s, both as garden plants and as

houseplant A houseplant, also known as a pot plant, potted plant, or indoor plant, is an ornamental plant cultivated indoors. for aesthetic or practical purposes. These plants are commonly found in House, homes, Office, offices, and various indoor spaces, w ...

s. Such genera include: ''

'', '' Achimenes'', ''

'', ''

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 hori ...

tuber Tubers are a type of enlarged structure that plants use as storage organs for nutrients, derived from stems or roots. Tubers help plants perennate (survive winter or dry months), provide energy and nutrients, and are a means of asexual reproduc ...

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.
{{Authority control Lamiales families