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Rubiaceae

January 29, 2023

Not to be confused with Rubiácea, a town in Brazil.

The Rubiaceae /ruːbiˈeɪʃiː/ are a family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs that are recognizable by simple, opposite leaves with interpetiolar stipules and sympetalous actinomorphic flowers. The family contains about 13,500 species in about 620 genera, which makes it the fourth-largest angiosperm family. Rubiaceae has a cosmopolitan distribution; however, the largest species diversity is concentrated in the tropics and subtropics.[1] Economically important genera include Coffea, the source of coffee, Cinchona, the source of the antimalarial alkaloid quinine, ornamental cultivars (e.g., Gardenia, Ixora, Pentas), and historically some dye plants (e.g., Rubia).

Description

The Rubiaceae are morphologically easily recognizable as a coherent group by a combination of characters: opposite or whorled leaves that are simple and entire, interpetiolar stipules, tubular sympetalous actinomorphic corollas and an inferior ovary.

A wide variety of growth forms are present: shrubs are most common (e.g. Coffea, Psychotria), but members of the family can also be trees (e.g. Cinchona, Nauclea), lianas (e.g. Psychotria samoritourei), or herbs (e.g. Galium, Spermacoce). Some epiphytes are also present (e.g. Myrmecodia). The plants usually contain iridoids, various alkaloids, and raphide crystals are common. The leaves are simple, undivided, and entire; there is only one case of pinnately compound leaves (Pentagonia osapinnata[2]). Leaf blades are usually elliptical, with a cuneate base and an acute tip. In three genera (Pavetta, Psychotria, Sericanthe), bacterial leaf nodules can be observed as dark spots or lines on the leaves. The phyllotaxis is usually decussate, rarely whorled (e.g. Fadogia), or rarely seemingly alternate resulting from the reduction of one leaf at each node (e.g. Sabicea sthenula). Characteristic for the Rubiaceae is the presence of stipules that are mostly fused to an interpetiolar structure on either side of the stem between the opposite leaves. Their inside surface often bears glands called "colleters", which produce mucilaginous compounds protecting the young shoot. The "whorled" leaves of the herbaceous tribe Rubieae have classically been interpreted as true leaves plus interpetiolar leaf-like stipules. The inflorescence is a cyme, rarely of solitary flowers (e.g. Rothmannia), and is either terminal or axillary and paired at the nodes. The 4-5-merous (rarely pleiomerous; e.g. six in Richardia[3]) flowers are usually bisexual and usually epigynous. The perianth is usually biseriate, although the calyx is absent in some taxa (e.g. Theligonum). The calyx mostly has the lobes fused at the base; unequal calyx lobes are not uncommon, and sometimes (e.g. Mussaenda) one lobe is enlarged and coloured (a so-called “semaphyl”). The corolla is sympetalous, mostly actinomorphic, usually tubular, mostly white or creamy but also yellow (e.g. Gardenia spp., Mycelia basiflora), and rarely blue (e.g. Faramea calyptrata) or red (e.g. Alberta magna, Ixora coccinea). The stamens are alternipetalous and epipetalous. Anthers are longitudinal in dehiscence, but are poricidal in some genera (e.g. Rustia, Tresanthera). The gynoecium is syncarpous with an inferior ovary (rarely secondarily superior, e.g. Gaertnera, Pagamea[4]). Placentation is axial, rarely parietal (e.g. Gardenia); ovules are anatropous to hemitropous, unitegmic, with a funicular obturator, one to many per carpel. Nectaries are often present as a nectariferous disk atop the ovary. The fruit is a berry, capsule (e.g. Oldenlandia), drupe (e.g. Coffea, Psychotria), or schizocarp (e.g. Cremocarpon). Red fruits are fairly dominant (e.g. Coffea arabica); yellow (e.g. Rosenbergiodendron formosum), orange (e.g. Vangueria infausta), or blackish fruits (e.g. Pavetta gardeniifolia) are equally common; blue fruits are rather exceptional save in the Psychotrieae and associated tribes. Most fruits are about 1 cm in diameter; very small fruits are relatively rare and occur in herbaceous tribes; very large fruits are rare and confined to the Gardenieae. The seeds are endospermous.[5][6]

Distribution and habitat

Rubiaceae have a cosmopolitan distribution and are found in nearly every region of the world, except for extreme environments such as the polar regions and deserts. The distribution pattern of the family is very similar to the global distribution of plant diversity overall. However, the largest diversity is distinctly concentrated in the humid tropics and subtropics. An exception is the tribe Rubieae, which is cosmopolitan but centered in temperate regions. Only a few genera are pantropical (e.g. Ixora, Psychotria), many are paleotropical, while Afro-American distributions are rare (e.g. Sabicea). Endemic rubiaceous genera are found in most tropical and subtropical floristic regions of the world. The highest number of species is found in Colombia, Venezuela, and New Guinea. When adjusted for area, Venezuela is the most diverse, followed by Colombia and Cuba.[7]

The Rubiaceae consist of terrestrial and predominantly woody plants. Woody rubiaceous shrubs constitute an important part of the understorey of low- and mid-altitude rainforests. 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).

Ecology

Flower biology

Most Rubiaceae are zoophilous. Entomophilous species produce nectar from an epigynous disk at the base of the corolla tube to attract insects. Ornithophily is rare and is found in red-flowered species of Alberta, Bouvardia, and Burchellia. Anemophilous species are found in the tribes Anthospermeae and Theligoneae and are characterized by hermaphroditic or unisexual flowers that exhibit a set of specialized features, such as striking sexual dimorphism, increased receptive surface of the stigmas and pendulous anthers.[5]

Although most Rubiaceae species are hermaphroditic, outbreeding is promoted through proterandry and spatial isolation of the reproductive organs. More complex reproductive strategies include secondary pollen presentation, heterodistyly, and unisexual flowers.

Secondary pollen presentation (also known as stylar pollen presentation or ixoroid pollen mechanism) is especially known from the Gardenieae and related tribes. The flowers are proterandrous and the pollen is shed early onto the outside of the stigmas or the upper part of the style, which serve as a 'receptaculum pollinis'. Increased surface area and irregularity of the pollen receptacle, caused by swellings, hairs, grooves or ridges often ensure a more efficient pollen deposition. After elongation of the style, animals transport the pollen to flowers in the female or receptive stage with exposed stigmatic surfaces. A pollen catapult mechanism is present in the genera Molopanthera and Posoqueria (tribe Posoquerieae) that projects a spherical pollen mass onto visiting sphingidae.[8]

Heterodistyly is another mechanism to avoid inbreeding and is widely present in the family Rubiaceae.[9] The tribes containing the largest number of heterostylous species are Spermacoceae and Psychotrieae. Heterostyly is absent in groups that have secondary pollen presentation (e.g. Vanguerieae).

Unisexual flowers also occur in Rubiaceae and most taxa that have this characteristic are dioecious. The two flower morphs are however difficult to observe as they are rather morphologically similar; male flowers have a pistillode with the ovaries empty and female flowers have empty, smaller anthers (staminodes).[5] Flowers that are morphologically hermaphrodite, but functionally dioecious are for example found in Pyrostria.[10]

Fruit biology

The dispersal units in Rubiaceae can be entire fruits, syncarps, mericarps, pyrenes or seeds. Fleshy fruit taxa are probably all (endo)zoochorous (e.g. tribes Pavetteae, Psychotrieae), while the dispersal of dry fruits is often unspecialized (e.g. tribes Knoxieae, Spermacoceae). When seeds function as diaspores, the dispersal is either anemochorous or hydrochorous. The three types of wind-dispersed diaspores in Rubiaceae are dust seeds (rare, e.g. Lerchea), plumed seeds (e.g. Hillia), and winged seeds (e.g. Coutarea). Long-distance dispersal by ocean currents is very rare (e.g. the seashore tree Guettarda speciosa). Other dispersal mechanisms are absent or at least very rare. Some Spermacoceae having seeds with elaiosomes are probably myrmecochorous (e.g. Spermacoce hepperiana). Epizoochorous taxa are limited to herbaceous Rubiaceae (e.g. Galium aparine fruits are densely covered with hooked bristly hairs).

Associations with other organisms

The genera Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum, and Squamellaria are succulent epiphytes that have evolved a mutualistic relationship with ants. Their hypocotyl grows out into an ant-inhabited tuber.[11] Some shrubs or trees have ant holes in their stems (e.g. Globulostylis).[12] Some Rubiaceae species have domatia that are inhabited by mites (viz. acarodomatia; e.g. Plectroniella armata).[13]

An intimate association between bacteria and plants is found in three rubiaceous genera (viz. Pavetta, Psychotria, and Sericanthe).[14] The presence of endophytic bacteria is visible by eye because of the formation of dark spots or nodules in the leaf blades. The endophytes have been identified as Burkholderia bacteria. A second type of bacterial leaf symbiosis is found in the genera Fadogia, Fadogiella, Globulostylis, Rytigynia, Vangueria (all belonging to the tribe Vanguerieae), where Burkholderia bacteria are found freely distributed among the mesophyll cells and no leaf nodules are formed.[15][16][17] The hypothesis regarding the function of the symbiosis is that the endophytes provide chemical protection against herbivory by producing certain toxic secondary metabolites.[18]

Systematics

The family Rubiaceae is named after Rubia, a name used by Pliny the Elder in his Naturalis Historia for madder (Rubia tinctorum).[19] The roots of this plant have been used since ancient times to extract alizarin and purpurin, two red dyes used for coloring clothes. The name rubia is therefore derived from the Latin word ruber, meaning red. The well-known genus Rubus (blackberries and raspberries) is unrelated and belongs to Rosaceae, the rose family.

Taxonomy

The name Rubiaceae (nomen conservandum) was published in 1789 by Antoine Laurent de Jussieu,[20] but the name was already mentioned in 1782.[21]

Several historically accepted families are since long included in Rubiaceae: Aparinaceae, Asperulaceae, Catesbaeaceae, Cephalanthaceae, Cinchonaceae, Coffeaceae, Coutariaceae, Galiaceae, Gardeniaceae, Guettardaceae, Hameliaceae, Hedyotidaceae, Houstoniaceae, Hydrophylacaceae, Lippayaceae, Lygodisodeaceae, Naucleaceae, Nonateliaceae, Operculariaceae, Pagamaeaceae, Psychotriaceae, Randiaceae, Sabiceaceae, Spermacoceaceae.[1]

More recently, the morphologically quite different families Dialypetalanthaceae,[22] Henriqueziaceae,[23] and Theligonaceae[24][25][26] were reduced to synonymy of Rubiaceae.

Subfamilies and tribes

The classical classification system of Rubiaceae distinguished only two subfamilies: Cinchonoideae, characterized by more than one ovule in each locule, and Coffeoideae, having one ovule in each locule.[27][28] This distinction, however, was criticized because of the distant position of two obviously related tribes, viz. Gardenieae with many ovules in Cinchonoideae and Ixoreae with one ovule in Coffeoideae, and because in species of Tarenna the number of ovules varies from one to several in each locule.[29][30] During the twentieth century other characters were used to delineate subfamilies, e.g. stylar pollen presentation, raphides, endosperm, heterostyly, etc. On this basis, three[31] or eight[32] subfamilies were recognised. The last subfamilial classification solely based on morphological characters divided Rubiaceae into four subfamilies: Cinchonoideae, Ixoroideae, Antirheoideae, and Rubioideae.[5] In general, problems of subfamilies delimitation in Rubiaceae based on morphological characters are linked with the extreme naturalness of the family, hence a relatively low divergence of its members.[5]

The introduction of molecular phylogenetics in Rubiaceae research has corroborated or rejected several of the conclusions made in the pre-molecular era. There is support for the subfamilies Cinchonoideae, Ixoroideae, and Rubioideae, although differently circumscribed, and Antirheoideae is shown to be polyphyletic.[33] The tribe Coptosapelteae including the genera Acranthera and Coptosapelta, and the monogeneric tribe Luculieae have not been placed within a subfamily and are sister to the rest of Rubiaceae.[34] Currently, in most molecular research concerning the family Rubiaceae, the classification with three subfamilies (Cinchonoideae, Ixoroideae, and Rubioideae) is followed.[35] However, an alternative view is proposed where only two subfamilies are recognized, an expanded Cinchonoideae (that includes Ixoroideae, Coptosapeltaeae and Luculieae) and Rubioideae.[26] The adoption of the Melbourne Code for botanical nomenclature had an unexpected impact on many names that have been long in use and are well-established in literature. According to the Melbourne Code, the subfamilial name Ixoroideae should be replaced by Dialypetalanthoideae.[36] However, Dialypetalanthus is morphologically quite aberrant in Rubiaceae and if it should be excluded from Rubiaceae, the subfamilial name remains Ixoroideae. Molecular studies also have substantial impact on tribal delimitations and taxonomic changes are still being made.[37][38] Also here, according to the Melbourne Code, the tribe Condamineeae should be renamed to Dialypetalantheae. The following list contains the validly published tribe names, however, some tribes might be disputed. The approximate number of species is indicated between brackets,[39] however, several genera and species are not yet placed in a tribe.

Genera

For a comprehensive list, see List of Rubiaceae genera.

The family Rubiaceae contains about 13,500 species in 619 genera. This makes it the fourth-largest family of flowering plants by number of species and fifth-largest by number of genera. Although taxonomic adjustments are still being made, the total number of accepted genera remains stable. In total, around 1338 genus names have been published, indicating that more than half of the published names are synonyms. Psychotria, with around 1530 species, is the largest genus within the family and the third-largest genus of the angiosperms, after the legume Astragalus and the orchid Bulbophyllum. However, the delimitation of Psychotria remains problematic and its adjustment might reduce the number of species. In total, 30 genera have more than 100 species. However, 138 genera are monotypic, which account for 22% of all genera, but only for 1.1% of all species.[7]

Phylogeny

Molecular studies have demonstrated the phylogenetic placement of Rubiaceae within the order Gentianales and the monophyly of the family is confirmed.[40][41] The relationships of the three subfamilies of Rubiaceae together with the tribes Coptosapelteae and Luculieae are shown in the phylogenetic tree below. The placement of these two groups relative to the three subfamilies has not been fully resolved.[41]

Evolution

The fossil history of the 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 has been confirmed as belonging to the Rubiaceae.[42]

The oldest confirmed fossils, which are fruits that strongly resemble those of the genus Emmenopterys, were found in the Washington and are 48–49 million years old. A fossil infructescence and fruit found in 44 million-year-old strata in Oregon was assigned to Emmenopterys dilcheri, an extinct species. The next-oldest fossils 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.[42]

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.[42]

Uses

Food

No staple foods are found in the Rubiaceae, but some species are consumed locally and fruits may be used as famine food. Examples are African medlar fruits (e.g. V. infausta, V. madagascariensis), African peach (Nauclea latifolia), and noni (Morinda citrifolia).

Beverage

The most economically important member of the family is the genus Coffea used in the production of coffee. Coffea includes 124 species, but only three species are cultivated for coffee production: C. arabica, C. canephora, and C. liberica.[7]

Medicinal

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 Carapichea 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.[43] The bark of the species Breonadia salicina have been used in traditional African medicine for many years.[44] The leaves of the Kratom plant (Mitragyna speciosa) contain a variety of alkaloids, including several psychoactive alkaloids and is traditionally prepared and consumed in Southeast Asia, where it has been known to exhibit both painkilling and stimulant qualities, behaving as a μ-opioid receptor agonist, and often being used in traditional Thai medicine in a similar way to and often as a replacement for opioid painkillers like morphine.

Ornamentals

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 contains plants cultivated in warmer-climate gardens; the most commonly grown species, Ixora coccinea, is frequently used for pretty red-flowering hedges. Mussaenda cultivars with enlarged, colored calyx lobes are shrubs with the aspect of Hydrangea; they are mainly cultivated in tropical Asia. 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. Nertera granadensis is a well-known house plant cultivated for its conspicuous orange berries. Other ornamental plants include Mitchella, Morinda, Pentas, and Rubia.

Dyes

Rose madder, the crushed root of Rubia tinctorum, yields a red dye, and the tropical Morinda citrifolia yields a yellow dye.

Culture

Image gallery

References

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External links

 
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  • "Rubiaceae" . Encyclopædia Britannica. Vol. 23 (11th ed.). 1911. p. 808.
  • Rubiaceae at The Plant List
  • Rubiaceae at Encyclopedia of Life
  • Rubiaceae at Angiosperm Phylogeny Website
  • Rubiaceae at Flora of China
  • Rubiaceae at Flora of Pakistan
  • Rubiaceae at Flora of Zimbabwe
  • Rubiaceae[permanent dead link] at Flora of Western Australia 2013-07-04 at the Wayback Machine
  • Rubiaceae at Flora of New Zealand
  • Rubiaceae at Integrated Taxonomic Information System
  • Rubiaceae at USDA NRCS Plants Database
  • Rubiaceae 2011-10-01 at the Wayback Machine at Botanic Garden Meise 2011-02-20 at the Wayback Machine
  • World Checklist of Rubiaceae at Royal Botanic Gardens, Kew

January 29, 2023
rubiaceae, confused, with, rubiácea, town, brazil, family, flowering, plants, commonly, known, coffee, madder, bedstraw, family, consists, terrestrial, trees, shrubs, lianas, herbs, that, recognizable, simple, opposite, leaves, with, interpetiolar, stipules, s. Not to be confused with Rubiacea a town in Brazil The Rubiaceae r uː b i ˈ eɪ ʃ iː are a family of flowering plants commonly known as the coffee madder or bedstraw family It consists of terrestrial trees shrubs lianas or herbs that are recognizable by simple opposite leaves with interpetiolar stipules and sympetalous actinomorphic flowers The family contains about 13 500 species in about 620 genera which makes it the fourth largest angiosperm family Rubiaceae has a cosmopolitan distribution however the largest species diversity is concentrated in the tropics and subtropics 1 Economically important genera include Coffea the source of coffee Cinchona the source of the antimalarial alkaloid quinine ornamental cultivars e g Gardenia Ixora Pentas and historically some dye plants e g Rubia RubiaceaeLuculia gratissimaScientific classificationKingdom PlantaeClade TracheophytesClade AngiospermsClade EudicotsClade AsteridsOrder GentianalesFamily RubiaceaeJuss Type genusRubiaL SubfamiliesCinchonoideae Ixoroideae RubioideaeSynonymsSee text Contents 1 Description 2 Distribution and habitat 3 Ecology 3 1 Flower biology 3 2 Fruit biology 3 3 Associations with other organisms 4 Systematics 4 1 Taxonomy 4 1 1 Subfamilies and tribes 4 1 2 Genera 4 2 Phylogeny 4 3 Evolution 5 Uses 5 1 Food 5 2 Beverage 5 3 Medicinal 5 4 Ornamentals 5 5 Dyes 6 Culture 7 Image gallery 8 References 9 External linksDescription EditThe Rubiaceae are morphologically easily recognizable as a coherent group by a combination of characters opposite or whorled leaves that are simple and entire interpetiolar stipules tubular sympetalous actinomorphic corollas and an inferior ovary A wide variety of growth forms are present shrubs are most common e g Coffea Psychotria but members of the family can also be trees e g Cinchona Nauclea lianas e g Psychotria samoritourei or herbs e g Galium Spermacoce Some epiphytes are also present e g Myrmecodia The plants usually contain iridoids various alkaloids and raphide crystals are common The leaves are simple undivided and entire there is only one case of pinnately compound leaves Pentagonia osapinnata 2 Leaf blades are usually elliptical with a cuneate base and an acute tip In three genera Pavetta Psychotria Sericanthe bacterial leaf nodules can be observed as dark spots or lines on the leaves The phyllotaxis is usually decussate rarely whorled e g Fadogia or rarely seemingly alternate resulting from the reduction of one leaf at each node e g Sabicea sthenula Characteristic for the Rubiaceae is the presence of stipules that are mostly fused to an interpetiolar structure on either side of the stem between the opposite leaves Their inside surface often bears glands called colleters which produce mucilaginous compounds protecting the young shoot The whorled leaves of the herbaceous tribe Rubieae have classically been interpreted as true leaves plus interpetiolar leaf like stipules The inflorescence is a cyme rarely of solitary flowers e g Rothmannia and is either terminal or axillary and paired at the nodes The 4 5 merous rarely pleiomerous e g six in Richardia 3 flowers are usually bisexual and usually epigynous The perianth is usually biseriate although the calyx is absent in some taxa e g Theligonum The calyx mostly has the lobes fused at the base unequal calyx lobes are not uncommon and sometimes e g Mussaenda one lobe is enlarged and coloured a so called semaphyl The corolla is sympetalous mostly actinomorphic usually tubular mostly white or creamy but also yellow e g Gardenia spp Mycelia basiflora and rarely blue e g Faramea calyptrata or red e g Alberta magna Ixora coccinea The stamens are alternipetalous and epipetalous Anthers are longitudinal in dehiscence but are poricidal in some genera e g Rustia Tresanthera The gynoecium is syncarpous with an inferior ovary rarely secondarily superior e g Gaertnera Pagamea 4 Placentation is axial rarely parietal e g Gardenia ovules are anatropous to hemitropous unitegmic with a funicular obturator one to many per carpel Nectaries are often present as a nectariferous disk atop the ovary The fruit is a berry capsule e g Oldenlandia drupe e g Coffea Psychotria or schizocarp e g Cremocarpon Red fruits are fairly dominant e g Coffea arabica yellow e g Rosenbergiodendron formosum orange e g Vangueria infausta or blackish fruits e g Pavetta gardeniifolia are equally common blue fruits are rather exceptional save in the Psychotrieae and associated tribes Most fruits are about 1 cm in diameter very small fruits are relatively rare and occur in herbaceous tribes very large fruits are rare and confined to the Gardenieae The seeds are endospermous 5 6 Distribution and habitat EditRubiaceae have a cosmopolitan distribution and are found in nearly every region of the world except for extreme environments such as the polar regions and deserts The distribution pattern of the family is very similar to the global distribution of plant diversity overall However the largest diversity is distinctly concentrated in the humid tropics and subtropics An exception is the tribe Rubieae which is cosmopolitan but centered in temperate regions Only a few genera are pantropical e g Ixora Psychotria many are paleotropical while Afro American distributions are rare e g Sabicea Endemic rubiaceous genera are found in most tropical and subtropical floristic regions of the world The highest number of species is found in Colombia Venezuela and New Guinea When adjusted for area Venezuela is the most diverse followed by Colombia and Cuba 7 The Rubiaceae consist of terrestrial and predominantly woody plants Woody rubiaceous shrubs constitute an important part of the understorey of low and mid altitude rainforests 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 Ecology EditFlower biology Edit Most Rubiaceae are zoophilous Entomophilous species produce nectar from an epigynous disk at the base of the corolla tube to attract insects Ornithophily is rare and is found in red flowered species of Alberta Bouvardia and Burchellia Anemophilous species are found in the tribes Anthospermeae and Theligoneae and are characterized by hermaphroditic or unisexual flowers that exhibit a set of specialized features such as striking sexual dimorphism increased receptive surface of the stigmas and pendulous anthers 5 Although most Rubiaceae species are hermaphroditic outbreeding is promoted through proterandry and spatial isolation of the reproductive organs More complex reproductive strategies include secondary pollen presentation heterodistyly and unisexual flowers Secondary pollen presentation also known as stylar pollen presentation or ixoroid pollen mechanism is especially known from the Gardenieae and related tribes The flowers are proterandrous and the pollen is shed early onto the outside of the stigmas or the upper part of the style which serve as a receptaculum pollinis Increased surface area and irregularity of the pollen receptacle caused by swellings hairs grooves or ridges often ensure a more efficient pollen deposition After elongation of the style animals transport the pollen to flowers in the female or receptive stage with exposed stigmatic surfaces A pollen catapult mechanism is present in the genera Molopanthera and Posoqueria tribe Posoquerieae that projects a spherical pollen mass onto visiting sphingidae 8 Heterodistyly is another mechanism to avoid inbreeding and is widely present in the family Rubiaceae 9 The tribes containing the largest number of heterostylous species are Spermacoceae and Psychotrieae Heterostyly is absent in groups that have secondary pollen presentation e g Vanguerieae Unisexual flowers also occur in Rubiaceae and most taxa that have this characteristic are dioecious The two flower morphs are however difficult to observe as they are rather morphologically similar male flowers have a pistillode with the ovaries empty and female flowers have empty smaller anthers staminodes 5 Flowers that are morphologically hermaphrodite but functionally dioecious are for example found in Pyrostria 10 Fruit biology Edit The dispersal units in Rubiaceae can be entire fruits syncarps mericarps pyrenes or seeds Fleshy fruit taxa are probably all endo zoochorous e g tribes Pavetteae Psychotrieae while the dispersal of dry fruits is often unspecialized e g tribes Knoxieae Spermacoceae When seeds function as diaspores the dispersal is either anemochorous or hydrochorous The three types of wind dispersed diaspores in Rubiaceae are dust seeds rare e g Lerchea plumed seeds e g Hillia and winged seeds e g Coutarea Long distance dispersal by ocean currents is very rare e g the seashore tree Guettarda speciosa Other dispersal mechanisms are absent or at least very rare Some Spermacoceae having seeds with elaiosomes are probably myrmecochorous e g Spermacoce hepperiana Epizoochorous taxa are limited to herbaceous Rubiaceae e g Galium aparine fruits are densely covered with hooked bristly hairs Associations with other organisms Edit The genera Anthorrhiza Hydnophytum Myrmecodia Myrmephytum and Squamellaria are succulent epiphytes that have evolved a mutualistic relationship with ants Their hypocotyl grows out into an ant inhabited tuber 11 Some shrubs or trees have ant holes in their stems e g Globulostylis 12 Some Rubiaceae species have domatia that are inhabited by mites viz acarodomatia e g Plectroniella armata 13 An intimate association between bacteria and plants is found in three rubiaceous genera viz Pavetta Psychotria and Sericanthe 14 The presence of endophytic bacteria is visible by eye because of the formation of dark spots or nodules in the leaf blades The endophytes have been identified as Burkholderia bacteria A second type of bacterial leaf symbiosis is found in the genera Fadogia Fadogiella Globulostylis Rytigynia Vangueria all belonging to the tribe Vanguerieae where Burkholderia bacteria are found freely distributed among the mesophyll cells and no leaf nodules are formed 15 16 17 The hypothesis regarding the function of the symbiosis is that the endophytes provide chemical protection against herbivory by producing certain toxic secondary metabolites 18 Systematics EditThe family Rubiaceae is named after Rubia a name used by Pliny the Elder in his Naturalis Historia for madder Rubia tinctorum 19 The roots of this plant have been used since ancient times to extract alizarin and purpurin two red dyes used for coloring clothes The name rubia is therefore derived from the Latin word ruber meaning red The well known genus Rubus blackberries and raspberries is unrelated and belongs to Rosaceae the rose family Taxonomy Edit The name Rubiaceae nomen conservandum was published in 1789 by Antoine Laurent de Jussieu 20 but the name was already mentioned in 1782 21 Several historically accepted families are since long included in Rubiaceae Aparinaceae Asperulaceae Catesbaeaceae Cephalanthaceae Cinchonaceae Coffeaceae Coutariaceae Galiaceae Gardeniaceae Guettardaceae Hameliaceae Hedyotidaceae Houstoniaceae Hydrophylacaceae Lippayaceae Lygodisodeaceae Naucleaceae Nonateliaceae Operculariaceae Pagamaeaceae Psychotriaceae Randiaceae Sabiceaceae Spermacoceaceae 1 More recently the morphologically quite different families Dialypetalanthaceae 22 Henriqueziaceae 23 and Theligonaceae 24 25 26 were reduced to synonymy of Rubiaceae Subfamilies and tribes Edit The classical classification system of Rubiaceae distinguished only two subfamilies Cinchonoideae characterized by more than one ovule in each locule and Coffeoideae having one ovule in each locule 27 28 This distinction however was criticized because of the distant position of two obviously related tribes viz Gardenieae with many ovules in Cinchonoideae and Ixoreae with one ovule in Coffeoideae and because in species of Tarenna the number of ovules varies from one to several in each locule 29 30 During the twentieth century other characters were used to delineate subfamilies e g stylar pollen presentation raphides endosperm heterostyly etc On this basis three 31 or eight 32 subfamilies were recognised The last subfamilial classification solely based on morphological characters divided Rubiaceae into four subfamilies Cinchonoideae Ixoroideae Antirheoideae and Rubioideae 5 In general problems of subfamilies delimitation in Rubiaceae based on morphological characters are linked with the extreme naturalness of the family hence a relatively low divergence of its members 5 The introduction of molecular phylogenetics in Rubiaceae research has corroborated or rejected several of the conclusions made in the pre molecular era There is support for the subfamilies Cinchonoideae Ixoroideae and Rubioideae although differently circumscribed and Antirheoideae is shown to be polyphyletic 33 The tribe Coptosapelteae including the genera Acranthera and Coptosapelta and the monogeneric tribe Luculieae have not been placed within a subfamily and are sister to the rest of Rubiaceae 34 Currently in most molecular research concerning the family Rubiaceae the classification with three subfamilies Cinchonoideae Ixoroideae and Rubioideae is followed 35 However an alternative view is proposed where only two subfamilies are recognized an expanded Cinchonoideae that includes Ixoroideae Coptosapeltaeae and Luculieae and Rubioideae 26 The adoption of the Melbourne Code for botanical nomenclature had an unexpected impact on many names that have been long in use and are well established in literature According to the Melbourne Code the subfamilial name Ixoroideae should be replaced by Dialypetalanthoideae 36 However Dialypetalanthus is morphologically quite aberrant in Rubiaceae and if it should be excluded from Rubiaceae the subfamilial name remains Ixoroideae Molecular studies also have substantial impact on tribal delimitations and taxonomic changes are still being made 37 38 Also here according to the Melbourne Code the tribe Condamineeae should be renamed to Dialypetalantheae The following list contains the validly published tribe names however some tribes might be disputed The approximate number of species is indicated between brackets 39 however several genera and species are not yet placed in a tribe basal Rubiaceae 60 sp Coptosapelteae Bremek ex S P Darwin 56 sp Luculieae Rydin amp B Bremer 4 sp Cinchonoideae Raf 1704 sp Chiococceae Benth amp Hook f 226 sp Cinchoneae DC 130 sp Guettardeae DC 737 sp Hamelieae A Rich ex DC 172 sp Hillieae Bremek ex S P Darwin 30 sp Hymenodictyeae Razafim amp B Bremer 26 sp Isertieae A Rich ex DC 17 sp Naucleeae DC ex Miq 190 sp Rondeletieae DC ex Miq 175 sp Strumpfieae Delprete amp T J Motley 1 sp Ixoroideae Raf 4052 sp Airospermeae Kainul amp B Bremer 7 sp Alberteae Hook f 7 sp Aleisanthieae Mouly J Florence amp B Bremer 10 sp Augusteae Kainul amp B Bremer 87 sp Bertiereae Bridson 57 sp Coffeeae DC 335 sp Condamineeae Hook f 310 sp Cordiereae A Rich ex DC emend Mouly 128 sp Cremasporeae Bremek ex S P Darwin 2 sp Crossopterygeae F White ex Bridson 1 sp Gardenieae A Rich ex DC 581 sp Greeneeae Mouly J Florence amp B Bremer 9 sp Henriquezieae Benth amp Hook f 22 sp Ixoreae Benth amp Hook f 565 sp Jackieae Korth 1 sp Mussaendeae Hook f 216 sp Octotropideae Bedd 106 sp Pavetteae A Rich ex Dumort 640 sp Posoquerieae Delprete 23 sp Retiniphylleae Hook f 21 sp Sabiceeae Bremek 157 sp Scyphiphoreae Kainul amp B Bremer 1 sp Sherbournieae Mouly amp B Bremer 58 sp Sipaneeae Bremek 43 sp Steenisieae Kainul amp B Bremer 5 sp Trailliaedoxeae Kainul amp B Bremer 1 sp Vanguerieae A Rich ex Dumort 651 sp Virectarieae Verdc 8 sp Rubioideae Verdc 7681 sp Anthospermeae Cham amp Schltdl ex DC 210 sp Argostemmateae Bremek ex Verdc 233 sp Clarkelleae Deb 1 sp Colletoecemateae Rydin amp B Bremer 3 sp Coussareeae Hook f 412 sp Craterispermeae Verdc 26 sp Cyanoneuroneae Razafim amp B Bremer 5 sp Danaideae B Bremer amp Manen 67 sp Dunnieae Rydin amp B Bremer 1 sp Gaertnereae Bremek ex S P Darwin 113 sp Knoxieae Hook f 130 sp Lasiantheae B Bremer amp Manen 288 sp Mitchelleae Razafim amp B Bremer amp Manen 14 sp Morindeae Miq 165 sp Ophiorrhizeae Bremek ex Verdc 380 sp Paederieae DC 82 sp Palicoureeae Robbr amp Manen 1161 sp Perameae Bremek ex S P Darwin 14 sp Prismatomerideae Y Z Ruan 24 sp Psychotrieae Cham amp Schltdl 1696 sp Putorieae Sweet 35 sp Rubieae Baill 974 sp Schizocoleeae Rydin amp B Bremer 2 sp Schradereae Bremek 54 sp Seychelleeae Razafim Kainul amp Rydin 1 sp Spermacoceae Cham amp Schltdl ex DC 1346 sp Theligoneae Wunderlich ex S P Darwin 4 sp Urophylleae Bremek ex Verdc 241 sp Genera Edit For a comprehensive list see List of Rubiaceae genera The family Rubiaceae contains about 13 500 species in 619 genera This makes it the fourth largest family of flowering plants by number of species and fifth largest by number of genera Although taxonomic adjustments are still being made the total number of accepted genera remains stable In total around 1338 genus names have been published indicating that more than half of the published names are synonyms Psychotria with around 1530 species is the largest genus within the family and the third largest genus of the angiosperms after the legume Astragalus and the orchid Bulbophyllum However the delimitation of Psychotria remains problematic and its adjustment might reduce the number of species In total 30 genera have more than 100 species However 138 genera are monotypic which account for 22 of all genera but only for 1 1 of all species 7 Phylogeny Edit Molecular studies have demonstrated the phylogenetic placement of Rubiaceae within the order Gentianales and the monophyly of the family is confirmed 40 41 The relationships of the three subfamilies of Rubiaceae together with the tribes Coptosapelteae and Luculieae are shown in the phylogenetic tree below The placement of these two groups relative to the three subfamilies has not been fully resolved 41 Rubiaceae RubioideaeIxoroideaeCinchonoideaeCoptosapelteaeLuculieaeEvolution Edit The fossil history of the 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 has been confirmed as belonging to the Rubiaceae 42 The oldest confirmed fossils which are fruits that strongly resemble those of the genus Emmenopterys were found in the Washington and are 48 49 million years old A fossil infructescence and fruit found in 44 million year old strata in Oregon was assigned to Emmenopterys dilcheri an extinct species The next oldest fossils 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 42 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 42 Uses EditFood Edit No staple foods are found in the Rubiaceae but some species are consumed locally and fruits may be used as famine food Examples are African medlar fruits e g V infausta V madagascariensis African peach Nauclea latifolia and noni Morinda citrifolia Beverage Edit The most economically important member of the family is the genus Coffea used in the production of coffee Coffea includes 124 species but only three species are cultivated for coffee production C arabica C canephora and C liberica 7 Medicinal Edit 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 Carapichea 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 43 The bark of the species Breonadia salicina have been used in traditional African medicine for many years 44 The leaves of the Kratom plant Mitragyna speciosa contain a variety of alkaloids including several psychoactive alkaloids and is traditionally prepared and consumed in Southeast Asia where it has been known to exhibit both painkilling and stimulant qualities behaving as a m opioid receptor agonist and often being used in traditional Thai medicine in a similar way to and often as a replacement for opioid painkillers like morphine Ornamentals Edit 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 contains plants cultivated in warmer climate gardens the most commonly grown species Ixora coccinea is frequently used for pretty red flowering hedges Mussaenda cultivars with enlarged colored calyx lobes are shrubs with the aspect of Hydrangea they are mainly cultivated in tropical Asia 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 Nertera granadensis is a well known house plant cultivated for its conspicuous orange berries Other ornamental plants include Mitchella Morinda Pentas and Rubia Dyes Edit Rose madder the crushed root of Rubia tinctorum yields a red dye and the tropical Morinda citrifolia yields a yellow dye Culture EditCinchona officinalis is the national tree of Ecuador and Peru Coffea arabica is the national flower of Yemen Ixora coccinea is the national flower of Suriname Warszewiczia coccinea is the national flower of Trinidad and Tobago The International Coffee Day is held each year on September 29 Image gallery Edit Alberta magna Arachnothryx leucophylla Asperula tinctoria Bikkia philippinensis Chiococca alba Coffea arabica Galium uliginosum Gardenia thunbergia Ixora coccinea Ixora javanica Mitragyna speciosa Morinda pubescens Nertera granadensis Psychotria poeppigiana Sherardia arvensisReferences Edit a b Angiosperm Phylogeny Website Retrieved 1 June 2014 Hammel BE 2015 Three new species of Pentagonia Rubiaceae from southern Central America one foreseen two surprising PDF Phyotneuron 46 1 13 Hall D Vandiver V Sellers B 1991 Brazil Pusley Richardia brasiliensis Moq PDF University of Florida Retrieved 9 August 2018 Igersheim A Puff C Leins P Erbar C 1994 Gynoecial development of Gaertnera Lam and of presumably allied taxa of the Psychotrieae Rubiaceae secondary superior vs inferior ovaries Botanische Jahrbucher fur Systematik 116 401 414 a b c d e Robbrecht E 1988 Tropical woody Rubiaceae Opera Botanica Belgica 1 1 271 Takhtajan A 2009 Flowering Plants 2 ed Springer ISBN 978 1 4020 9608 2 a b c Davis AP Govaerts R Bridson DM Ruhsam M Moat J Brummitt NA 2009 A global assessment of distribution diversity endemism and taxonomic effort in the Rubiaceae Annals of the Missouri Botanical Garden 96 1 68 78 doi 10 3417 2006205 S2CID 86558504 Delprete PG 2009 Taxonomic history morphology and reproductive biology of the tribe Posoquerieae Rubiaceae Ixoroideae Annals of the Missouri Botanical Garden 96 1 79 89 doi 10 3417 2006192 S2CID 86495664 Anderson WR 1973 A morphological hypothesis for the origin of heterostyly in the Rubiaceae Taxon 22 5 6 537 542 doi 10 2307 1218628 JSTOR 1218628 Bridson DM 1987 Studies in African Rubiaceae Vanguerieae a new circumscription of Pyrostria and a new subgenus Canthium subgen Bullockia Kew Bulletin 42 3 611 639 doi 10 2307 4110068 JSTOR 4110068 Kapitany A 2007 Australian succulent plants an introduction Boronia Victoria Kapitany Concepts pp 144 155 ISBN 978 0 646 46381 0 Verstraete B Lachenaud O Smets E Dessein S Sonke B 2013 Taxonomy and phylogeny of Cuviera Rubiaceae Vanguerieae and reinstatement of the genus Globulostylis with the description of three new species Botanical Journal of the Linnean Society 173 3 407 441 doi 10 1111 boj 12062 Tilney PM van Wyk AE van deer Merwe CF 2012 Structural evidence in Plectroniella armada Rubiaeae for possible material exchange between domatia and mites PLOS ONE 7 7 e39984 Bibcode 2012PLoSO 739984T doi 10 1371 journal pone 0039984 PMC 3390328 PMID 22792206 Lemaire B Vandamme P Merckx V Smets E Dessein S 2011 Bacterial leaf symbiosis in angiosperms host specificity without co speciation PLOS ONE 6 9 e24430 Bibcode 2011PLoSO 624430L doi 10 1371 journal pone 0024430 PMC 3168474 PMID 21915326 Verstraete B Van Elst D Steyn H Van Wyk B Lemaire B Smets E Dessein S 2011 Endophytic bacteria in toxic South African plants identification phylogeny and possible involvement in gousiekte PLOS ONE 6 4 e19265 Bibcode 2011PLoSO 619265V doi 10 1371 journal pone 0019265 PMC 3082559 PMID 21541284 Verstraete B Janssens S Smets E Dessein S 2013 Symbiotic beta proteobacteria beyond legumes Burkholderia in Rubiaceae PLOS ONE 8 1 e55260 doi 10 1371 journal pone 0055260 PMC 3555867 PMID 23372845 Verstraete B Janssens S Lemaire B Smets E Dessein S 2013 Phylogenetic lineages in Vanguerieae Rubiaceae associated with Burkholderia bacteria in sub Saharan Africa American Journal of Botany 100 12 2380 2387 doi 10 3732 ajb 1300303 PMID 24275705 Sieber S Carlier AL Neuburger M Grabenweger G Eberl L Gademann K 2015 Isolation and total synthesis of kirkamide an aminocyclitol from an obligate leaf nodule symbiont PDF Angewandte Chemie International Edition 54 27 7968 7970 doi 10 1002 anie 201502696 PMID 26033226 Simpson MG 2006 Plant Systematics 1 ed Elsevier Academic Press ISBN 978 0 12 644460 5 Jussieu A L de 1789 Genera Plantarum Paris Herissant amp Barrois p 206 Durand JF 1782 Notions Elementaires de Botanique Dijon LN Frantin p 274 Fay MF Bremer B Prance GT van der Bank M Bridson D Chase MW 2000 Plastid rbcL sequence data show Dialypetalanthus to be a member of Rubiaceae Kew Bulletin 55 4 853 864 doi 10 2307 4113630 JSTOR 4113630 Rogers GK 1981 The wood of Gleasonia Henriquezia and Platycarpum Rubiaceae and its bearing on their classification some new considerations Brittonia 33 3 461 465 doi 10 2307 2806441 JSTOR 2806441 S2CID 85070681 Wunderlich R 1971 Die systematische Stellung von Theligonum Osterreichische Botanische Zeitschrift 119 4 5 329 394 doi 10 1007 bf01377490 S2CID 41742399 Rutishauser F Ronse Decraene LP Smets E Mendoza Heuer I 1998 Theligonum cynocrambe developmental morphology of a peculiar rubiaceous herb PDF Plant Systematics and Evolution 210 1 1 24 doi 10 1007 BF00984724 S2CID 22331639 a b Robbrecht E Manen JF 2006 The major evolutionary lineages of the coffee family Rubiaceae angiosperms Combined analysis nDNA and cpDNA to infer the position of Coptosapelta and Luculia and supertree construction based on rbcL rps16 trnL trnF and atpB rbcL data A new classification in two subfamilies Cinchonoideae and Rubioideae Systematic Geography of Plants 76 85 146 Hooker JD 1873 Ordo LXXXIV Rubiaceae In Bentham G Hooker JD eds Genera planetarium ad exemplaria imprimis in herbaria kewensibus servata defirmata Vol 2 London pp 7 151 Schumann K 1891 Rubiaceae In Engler A Prantl K eds Die naturlichen Pflanzenfamilien Vol 4 Leipzig Engelmann pp 1 156 Baillon H 1878 Sur les limits du genre Ixora Adansonia 12 213 219 Solereder H 1893 Ein Beitrag zur anatomischen Charakteristik und zur Systematik deer Rubiaceen Bull Herb Boissier 1 167 183 Verdcourt B 1958 Remarks on the classification of the Rubiaceae Bulletin du Jardin Botanique de l Etat a Bruxelles 28 3 209 281 doi 10 2307 3667090 JSTOR 3667090 Bremekamp CEB 1966 Remarks on the position the delimitation and the subdivision of the Rubiaceae Acta Botanica Neerlandica 15 1 33 doi 10 1111 j 1438 8677 1966 tb00207 x Bremer B Andreasen K Olsson D 1995 Subfamilial and tribal relationships in the Rubiaceae based on rbcL sequence data Annals of the Missouri Botanical Garden 82 3 383 397 doi 10 2307 2399889 JSTOR 2399889 Rydin C Kainulainen K Razafimandimbison SG Smedmark JE Bremer B 2009 Deep divergences in the coffee family and the systematic position of Acranthera PDF Plant Systematics and Evolution 278 1 2 101 123 doi 10 1007 s00606 008 0138 4 S2CID 26350567 Bremer B 2009 A review of molecular phylogenetic studies of Rubiaceae Annals of the Missouri Botanical Garden 96 1 4 26 doi 10 3417 2006197 S2CID 53378010 Reveal JL 2012 Newly required infrafamilial names mandated by changes in the code of nomenclature for algae fungi and plants Phytoneuron 33 1 32 Kainulainen K Razafimandimbison SG Bremer B 2013 Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae Rubiaceae Botanical Journal of the Linnean Society 173 3 387 406 doi 10 1111 boj 12038 Mouly A Kainulainen K Persson C Davis AP Wong KM Razafimandimbison SG Bremer B 2014 Phylogenetic structure and clade circumscriptions in the Gardenieae complex Rubiaceae Taxon 63 4 801 818 doi 10 12705 634 4 World Checklist of Rubiaceae Retrieved 1 March 2016 Bremer B Bremer K Heidari N Erixon P Olmstead RG Anderberg AA Kallersjo M Barkhordarian E 2002 Phylogenetics of asteroids based on 3 coding and 3 non coding chloroplast DNA markers and the utility of non coding DNA at higher taxonomic levels Molecular Phylogenetics and Evolution 24 2 274 301 doi 10 1016 s1055 7903 02 00240 3 PMID 12144762 a b Bremer B Eriksson T 2009 Time tree of Rubiaceae phylogeny and dating the family subfamilies and tribes International Journal of Plant Sciences 170 6 766 793 doi 10 1086 599077 S2CID 49332892 a b c Graham A 2009 Fossil record of the Rubiaceae Annals of the Missouri Botanical Garden 96 1 90 108 doi 10 3417 2006165 S2CID 84196922 Riba J Valle M Urbano G Yritia M Morte A Barbanoj MJ 2003 Human pharmacology of ayahuasca subjective and cardiovascular effects monoamine metabolite excretion and pharmacokinetics Journal of Pharmacology and Experimental Therapeutics 306 1 73 83 doi 10 1124 jpet 103 049882 PMID 12660312 S2CID 6147566 Neuwinger Hans Dieter 1994 African Ethnobotany Poisons and Drugs Chemistry Pharmacology Toxicology Stuttgart Germany Chapman amp Hall External links Edit Wikimedia Commons has media related to Rubiaceae Wikispecies has information related to Rubiaceae Rubiaceae Encyclopaedia Britannica Vol 23 11th ed 1911 p 808 Rubiaceae at The Plant List Rubiaceae at Encyclopedia of Life Rubiaceae at Angiosperm Phylogeny Website Rubiaceae at Flora of China Rubiaceae at Flora of Pakistan Rubiaceae at Flora of Zimbabwe Rubiaceae permanent dead link at Flora of Western Australia Archived 2013 07 04 at the Wayback Machine Rubiaceae at Flora of New Zealand Rubiaceae at Integrated Taxonomic Information System Rubiaceae at USDA NRCS Plants Database Rubiaceae Archived 2011 10 01 at the Wayback Machine at Botanic Garden Meise Archived 2011 02 20 at the Wayback Machine World Checklist of Rubiaceae at Royal Botanic Gardens Kew Retrieved from https en wikipedia org w index php title Rubiaceae amp oldid 1123462701, wikipedia, wiki, book, books, library,

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