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Rubiaceae is a family of flowering plants, variously called the coffee family, madder family, or bedstraw family. The group contains many commonly known plants, including the economically important coffee (Coffea), quinine (Cinchona), and gambier (Uncaria), the medicinal ipecacuanha (Carapichea ipecacuanha), and the horticulturally valuable madder (Rubia), west Indian jasmine (Ixora), partridgeberry (Mitchella), Morinda, Gardenia, and Pentas.
Members of the coffee family tend to be concentrated in warmer and tropical climates around the world. Currently, about 611 genera and more than 13,000 species are placed in Rubiaceae. This makes it the fourth-largest family of flowering plants by number of species, and fifth-largest by number of genera.
The family takes its name from the madder genus Rubia, which derives from the Latin word ruber, meaning "red". Rubia as a name for madder was coined by Pliny. (The well-known genus Rubus (blackberries and raspberries) is unrelated and belongs to Rosaceae, the rose family.)
Even though Rubiaceae are found in nearly every major region of the world (with the exception of continental Antarctica, the high arctic, and portions of central Africa and Asia), diversity is highest in the humid tropics. The pattern of diversity in the family is very similar to the global distribution of plant diversity overall.
The greatest number of species occurs in Colombia, Venezuela and New Guinea. When adjusted for area, Venezuela is the most diverse, followed by Colombia and Cuba. A large number of poorly studied species exist as understorey shrubs in Madagascar and are threatened with habitat destruction.
Rubiaceae are an easily recognizable family characterized by opposite leaves that are simple and entire, with interpetiolar stipules, tubular sympetalous corollas and an inferior ovary. Exceptionally, there are some plants that have only a single leaf at each node, alternating from one side to the other. In these cases, the alternate leaf arrangement is produced through the suppression of one leaf at each node. A wide variety of growth forms are present in the Rubiaceae. While shrubs are most common, members of the family can also be trees, lianas or herbs. The flowers, which are usually bisexual, have a 4–5 lobed calyx and generally a 4–5 lobed corolla, 4 or 5 stamens and two carpels.
The fossil history of Rubiaceae goes back at least as far as the Eocene. The geographic distribution of these fossils, coupled with the fact that they represent all three subfamilies, is indicative of an earlier origin for the family, probably in the Late Cretaceous or Paleocene. Although fossils dating back to the Cretaceous and Palaeocene have been referred to the family by various authors, none of these fossils have been confirmed as belonging to Rubiaceae.
The oldest confirmed fossils, which are of fruit that bear strong resemblance to the genus Emmenopterys were found in Washington State and are 48–49 million years old. A fossil infructescence and fruit found in 44-million-year-old strata in Oregon were assigned to Emmenopterys dilcheri, an extinct species. The next oldest fossils after these date to the Late Eocene and include Canthium from Australia, Faramea from Panama, Guettarda from New Caledonia, and Paleorubiaceophyllum, an extinct genus, from the southeastern United States.
Fossil Rubiaceae are known from three regions in the Eocene (North America north of Mexico, Mexico-Central America-Caribbean, and Southeast Pacific-Asia). In the Oligocene they are found in these three regions plus Africa. In the Miocene they are found in these four regions, plus South America, and Europe.
Rubiaceae are tolerant of a broad array of environmental conditions (soil types, altitudes, community structures, etc.), and do not specialize in one specific habitat type (although genera within the family often specialize). The plants tend not to be eaten by the larvae of butterflies, but some sphingids (Semanophorae) do appear to prefer them. The genera Myrmecodia and Hydnophytum are interesting, as they are epiphytes that have evolved mutualistic relationships with ants.
The most economically important member of the family, and the world's second most important commodity (after petroleum) is the genus Coffea used in the production of coffee. Coffea includes 103 species, but only three species are cultivated for coffee production: C. arabica, C. canephora, and C. liberica.
The bark of trees in the genus Cinchona is the source of a variety of alkaloids, the most familiar of which is quinine, one of the first agents effective in treating malaria. Woodruff (Galium odoratum) is a small herbaceous perennial that contains coumarin—a natural precursor of warfarin—and the South American plant Psychotria ipecacuanha is the source of the emetic ipecac. Psychotria viridis is frequently used as a source of dimethyltryptamine in the preparation of ayahuasca, a psychoactive decoction.
Originally from China, the common gardenia (Gardenia jasminoides) is a widely grown garden plant and flower in frost-free climates worldwide. Several other species from the genus are also seen in horticulture. The genus Ixora also contains plants seen cultivated in warmer climate gardens. The New Zealand native Coprosma repens is a commonly used plant for hedges. The South African Rothmannia globosa is seen as a specimen tree in horticulture.
The family was described for the first time by Antoine Laurent de Jussieu in 1789. Currently, a number of traditionally accepted families (Dialypetalanthaceae, Henriqueziaceae, Naucleaceae, and Theligonaceae) are included in Rubiaceae following molecular phylogenetic research by the Angiosperm Phylogeny Group.
Subfamilies and tribes
Rubiaceae were "classically" divided into two subfamilies, Coffeoideae, which have placentas bearing a single ovule, and Cinchonoideae, which have more than one ovule per placenta. However, as early as 1893 Hans Solereder identified this system as artificial, since individuals with a single species of Tarenna had placentas with one or more ovules. During the twentieth century other characteristics were used to delineate subfamilies including the distribution of raphides, the absence of endosperm or heterostyly. On this basis, three, five or eight subfamilies were recognised. In 1988, Elmar Robbrecht divided the Rubiaceae into four subfamilies: Ixoroideae, Cinchonoideae, Antirheoideae and Rubioideae. While the limits of Rubioideae remained almost unchanged in the face of molecular studies, Antirheoideae was shown to be polyphyletic, while Ixoroideae was broadened and Cinchonoideae narrowed. Currently three subfamilies (Cinchonoideae, Ixoroideae and Rubioideae) and 52 tribes are recognised. Elmar Robbrecht and Jean-François Manen, however, have proposed that only two subfamilies be recognised, an expanded Cinchonoideae (that includes Ixoroideae, Coptosapelta and Luculia) and Rubioideae. One tribe, Coptosapelteae Bremek. ex Darwin, and one genus, Luculia Sweet, have not been placed within a subfamily and are sister to the rest of Rubiaceae.
- For a full list, see List of Rubiaceae genera
According to the World Checklist of Rubiaceae, 611 genera and 13,143 species were accepted as of 2009. Psychotria, with 1834 species, is the largest genus within the family, and the third-largest genus of the angiosperms, after the legume Astragalus and the orchid Bulbophyllum. Twenty-nine other genera also have more than 100 species. On the other hand, 211 genera are monotypic. Together these account for more than a third of all genera in Rubiaceae, but only around 1% of all species.
Relationships of the three subfamilies of Rubiaceae, together with the tribe Coptosapelteae and the genus Luculia. The placement of these two groups relative to the three subfamilies has not been fully resolved.
In comparison to other families, minimal research has focused on understanding the evolutionary history of the Rubiaceae. Rubiaceae is large; the number of species in the Rubiaceae is more than double that of mammals, and its fossil record is incomplete. Apart from Coffea, Rubiaceae includes relatively few genera of economic importance, providing little economic incentive to stimulate research concerning a systematically difficult family.
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