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The Rubiaceae
Rubiaceae
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. The family contains about 13,500 species in 611 genera, which makes it the fourth-largest angiosperm family. Rubiaceae
Rubiaceae
has a cosmopolitan distribution; however, the largest species diversity is concentrated in the (sub)tropics.[1] Economic importance includes Coffea, the source of coffee, Cinchona, the source of the antimalarial alkaloid quinine, some dye plants (e.g. Rubia), and ornamental cultivars (e.g. Gardenia, Ixora, Pentas).

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
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 links

Description[edit] The Rubiaceae
Rubiaceae
are morphologically easily recognizable as a coherent group by a combination of characters: opposite leaves that are simple and entire (with the exception of several newly discovered compound-leaved species of Pentagonia)[2], 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
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; 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 alternate resulting from the suppression of one leaf at each node (e.g. Sabicea
Sabicea
sthenula). Characteristic for the Rubiaceae
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 Rubieae
Rubieae
tribe 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 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 has four or five sepals with basally fused lobes. The corolla is sympetalous with four or five lobes, mostly actinomorphic, usually tubular, mostly white or creamy but also yellow (e.g. Gardenia
Gardenia
spp., Mycelia basiflora), and rarely blue (e.g. Faramea
Faramea
calyptrata) or red (e.g. Alberta magna, Ixora
Ixora
coccinea). They have four or five stamens, which are alternipetalous and epipetalous. Anthers are longitudinal in dehiscence, but some genera are poricidal (e.g. Rustia). The gynoecium is syncarpous with an inferior ovary (rarely secondarily superior, e.g. Gaertnera, Pagamea[3]). 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. Psychotria), or schizocarp (e.g. Cremocarpon). Red fruits are fairly dominant (e.g. Coffea arabica); yellow (e.g. Rosenbergiodendron formosum), orange (e.g. Vangueria
Vangueria
infausta), or blackish fruits (e.g. Pavetta
Pavetta
gardeniifolia) are equally common; blue fruits are rather exceptional save in the Psychotrieae
Psychotrieae
and associated tribes. Most fruits are about 1.0 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.[4][5] Distribution and habitat[edit] Rubiaceae
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 Rubieae
Rubieae
tribe, 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
Colombia
and Cuba.[6] The Rubiaceae
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
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[edit] Flower biology[edit] Most Rubiaceae
Rubiaceae
are zoophilous. Entomophilous species produce nectar from an epigynous disk at the base of the corolla tube to attract insects. Ornithophily
Ornithophily
is rare and is found in red-flowered species of Alberta, Bouvardia, and Burchellia. Anemophylous species are found in the tribes Anthospermeae and Theligoneae
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.[4] Although most Rubiaceae
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
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
Posoqueria
(tribe Posoquerieae) that projects a spherical pollen mass onto visiting sphingidae.[7] Heterodistyly is another mechanism to avoid inbreeding and is widely present in the Rubiaceae
Rubiaceae
family.[8] The tribes containing the largest number of heterostylous species are Spermacoceae
Spermacoceae
and Psychotrieae. Heterostyly
Heterostyly
is absent in groups that have secondary pollen presentation (e.g. Vanguerieae). Unisexual flowers also occur in Rubiaceae
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).[4] Flowers that are morphologically hermaphrodite, but functionally dioecious are for example found in Pyrostria.[9] Fruit biology[edit] The dispersal units in Rubiaceae
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
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
Guettarda
speciosa). Other dispersal mechanisms are absent or at least very rare. Some Spermacoceae
Spermacoceae
having seeds with elaiosomes are probably myrmecochorous (e.g. Spermacoce
Spermacoce
hepperiana). Epizoochorous taxa are limited to herbaceous Rubiaceae
Rubiaceae
(e.g. Galium
Galium
aparine fruits are densely covered with hooked bristly hairs). Associations with other organisms[edit] The genera Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum, and Squamellaria
Squamellaria
are succulent epiphytes that have evolved a mutualistic relationship with ants. Their hypocotyl grows out into an ant-inhabited tuber.[10] Some shrubs or trees have ant holes in their stems (e.g. Globulostylis).[11] Some Rubiaceae
Rubiaceae
species have domatia that are inhabited by mites (viz. acarodomatia; e.g. Plectroniella armata).[12] An intimate association between bacteria and plants is found in three rubiaceous genera (viz. Pavetta, Psychotria, and Sericanthe).[13] 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
Burkholderia
bacteria. A second type of bacterial leaf symbiosis is found in the genera Fadogia, Fadogiella, Globulostylis, Rytigynia, Vangueria
Vangueria
(all belonging to the Vanguerieae tribe), where Burkholderia
Burkholderia
bacteria are found freely distributed among the mesophyll cells and no leaf nodules are formed.[14][15][16] The hypothesis regarding the function of the symbiosis is that the endophytes provide chemical protection against herbivory by producing certain toxic secondary metabolites.[17] Systematics[edit] The Rubiaceae
Rubiaceae
family is named after Rubia, a name used by Pliny the Elder in his Naturalis Historia for madder ( Rubia
Rubia
tinctorum).[18] 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
Rubus
(blackberries and raspberries) is unrelated and belongs to Rosaceae, the rose family. Taxonomy[edit] The name Rubiaceae
Rubiaceae
(nomen conservandum) was published in 1789 by Antoine Laurent de Jussieu,[19] but the name was already mentioned in 1782.[20] 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,[21] Henriqueziaceae,[22] and Theligonaceae[23][24][25] were reduced to synonymy of Rubiaceae. Subfamilies
Subfamilies
and tribes[edit] The classical classification system of Rubiaceae
Rubiaceae
distinguished only two subfamilies: Cinchonoideae, characterized by more than one ovule in each locule, and Coffeoideae, having one ovule in each locule.[26][27] This distinction, however, was criticized because of the distant position of two obviously related tribes, viz. Gardenieae with many ovules in Cinchonoideae
Cinchonoideae
and Ixoreae
Ixoreae
with one ovule in Coffeoideae, and because in species of Tarenna
Tarenna
the number of ovules varies from one to several in each locule.[28][29] During the twentieth century other characters were used to delineate subfamilies, e.g. stylar pollen presentation, raphides, endosperm, heterostyly, etc. On this basis, three[30] or eight[31] subfamilies were recognised. The last subfamilial classification solely based on morphological characters divided Rubiaceae
Rubiaceae
into four subfamilies: Cinchonoideae, Ixoroideae, Antirheoideae, and Rubioideae.[4] In general, problems of subfamilies delimitation in Rubiaceae
Rubiaceae
based on morphological characters are linked with the extreme naturalness of the family, hence a relatively low divergence of its members.[4] The introduction of molecular phylogenetics in Rubiaceae
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.[32] The tribe Coptosapelteae
Coptosapelteae
including the genera Acranthera and Coptosapelta, and the monogeneric tribe Luculieae
Luculieae
have not been placed within a subfamily and are sister to the rest of Rubiaceae.[33] Currently, in most molecular research concerning the Rubiaceae
Rubiaceae
family, the classification with three subfamilies (Cinchonoideae, Ixoroideae, and Rubioideae) is followed.[34] However, an alternative view is proposed where only two subfamilies are recognized, an expanded Cinchonoideae (that includes Ixoroideae, Coptosapeltaeae and Luculieae) and Rubioideae.[25] 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
Ixoroideae
should be replaced by Dialypetalanthoideae.[35] However, Dialypetalanthus is morphologically quite aberrant in Rubiaceae
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.[36][37] Also here, according to the Melbourne Code, the tribe Condamineeae
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,[38] however, several genera and species are not yet placed in a tribe.

basal Rubiaceae
Rubiaceae
(59 sp.)

Coptosapelteae
Coptosapelteae
Bremek. ex S.P.Darwin (55 sp.) Luculieae
Luculieae
Rydin & B.Bremer (4 sp.)

Cinchonoideae
Cinchonoideae
Raf. (1708 sp.)

Chiococceae
Chiococceae
Benth. & Hook.f. (224 sp.) Cinchoneae
Cinchoneae
DC. (125 sp.) Guettardeae
Guettardeae
DC. (747 sp.) Hamelieae
Hamelieae
A.Rich. ex DC. (171 sp.) Hymenodictyeae
Hymenodictyeae
Razafim. & B.Bremer (25 sp.) Hillieae Bremek. ex S.P.Darwin (29 sp.) Isertieae
Isertieae
A.Rich. ex DC. (16 sp.) Naucleeae
Naucleeae
DC. ex Miq. (192 sp.) Rondeletieae
Rondeletieae
DC. ex Miq. (178 sp.) Strumpfieae
Strumpfieae
Delprete & T.J.Motley (1 sp.)

Ixoroideae
Ixoroideae
Raf. (3960 sp.)

Airospermeae Kainul. & B.Bremer (7 sp.) Alberteae
Alberteae
Hook.f. (7 sp.) Aleisanthieae Mouly, J.Florence & B.Bremer (10 sp.) Augusteae Kainul. & B.Bremer (86 sp.) Bertiereae
Bertiereae
Bridson (57 sp.) Coffeeae
Coffeeae
DC. (303 sp.) Condamineeae
Condamineeae
Hook.f. (305 sp.) Cordiereae A.Rich. ex DC. emend. Mouly (124 sp.) Cremasporeae Bremek. ex S.P.Darwin (2 sp.) Crossopterygeae F.White ex Bridson (1 sp.) Gardenieae
Gardenieae
A.Rich. ex DC. (587 sp.) Greeneeae Mouly, J.Florence & B.Bremer (9 sp.) Henriquezieae Benth. & Hook.f. (20 sp.) Ixoreae
Ixoreae
Benth. & Hook.f. (545 sp.) Jackieae Korth. (1 sp.) Mussaendeae
Mussaendeae
Hook.f. (221 sp.) Octotropideae
Octotropideae
Bedd. (96 sp.) Pavetteae
Pavetteae
A.Rich. ex Dumort. (624 sp.) Posoquerieae Delprete (23 sp.) Retiniphylleae Hook.f. (20 sp.) Sabiceeae Bremek. (164 sp.) Scyphiphoreae
Scyphiphoreae
Kainul. & B.Bremer (1 sp.) Sherbournieae Mouly & B.Bremer (54 sp.) Sipaneeae Bremek. (43 sp.) Steenisieae Kainul. & B.Bremer (5 sp.) Trailliaedoxeae Kainul. & B.Bremer (1 sp.) Vanguerieae
Vanguerieae
A.Rich. ex Dumort. (644 sp.)

Rubioideae
Rubioideae
Verdc. (7571 sp.)

Anthospermeae Cham. & Schltdl. ex DC. (208 sp.) Argostemmateae Bremek. ex Verdc. (215 sp.) Clarkelleae Deb (1 sp.) Colletoecemateae Rydin & B.Bremer (3 sp.) Coussareeae Hook.f. (402 sp.) Craterispermeae Verdc. (16 sp.) Cyanoneuroneae Razafim. & B.Bremer (5 sp.) Danaideae B.Bremer & Manen (50 sp.) Dunnieae Rydin & B.Bremer (1 sp.) Gaertnereae Bremek. ex S.P.Darwin (95 sp.) Knoxieae
Knoxieae
Hook.f. (131 sp.) Lasiantheae B.Bremer & Manen (239 sp.) Mitchelleae Razafim. & B.Bremer & Manen (14 sp.) Morindeae
Morindeae
Miq. (165 sp.) Ophiorrhizeae
Ophiorrhizeae
Bremek. ex Verdc. (364 sp.) Paederieae DC. (81 sp.) Palicoureeae Robbr. & Manen (817 sp.) Perameae Bremek. ex S.P.Darwin (14 sp.) Prismatomerideae Y.Z.Ruan (23 sp.) Psychotrieae
Psychotrieae
Cham. & Schltdl. (2114 sp.) Putorieae
Putorieae
(34 sp.) Rubieae
Rubieae
Baill. (938 sp.) Schizocoleeae Rydin & B.Bremer (2 sp.) Schradereae Bremek. (55 sp.) Spermacoceae
Spermacoceae
Cham. & Schltdl. ex DC. (1344 sp.) Theligoneae
Theligoneae
Wunderlich ex S.P.Darwin (4 sp.) Urophylleae Bremek. ex Verdc. (236 sp.)

Genera[edit] For a comprehensive list, see List of Rubiaceae
Rubiaceae
genera. The Rubiaceae
Rubiaceae
family 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 1850 species, is the largest genus within the family and the third-largest genus of the angiosperms, after the legume Astragalus
Astragalus
and the orchid Bulbophyllum. However, the delimitation of Psychotria
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.[6] Phylogeny[edit] Molecular studies have demonstrated the phylogenetic placement of Rubiaceae
Rubiaceae
within the order Gentianales
Gentianales
and the monophyly of the family is confirmed.[39][40] The relationships of the three subfamilies of Rubiaceae
Rubiaceae
together with the tribes Coptosapelteae
Coptosapelteae
and Luculieae
Luculieae
are shown in the phylogenetic tree below. The placement of these two groups relative to the three subfamilies has not been fully resolved.[40]

Rubiaceae

Rubioideae

Ixoroideae

Cinchonoideae

Coptosapelteae

Luculieae

Evolution[edit] The fossil history of the Rubiaceae
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
Cretaceous
and Palaeocene
Palaeocene
have been referred to the family by various authors, none of these fossils has been confirmed as belonging to the Rubiaceae.[41] 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
Oregon
was assigned to Emmenopterys dilcheri, an extinct species. The next-oldest fossils date to the Late Eocene
Eocene
and include Canthium
Canthium
from Australia, Faramea
Faramea
from Panama, Guettarda
Guettarda
from New Caledonia, and Paleorubiaceophyllum, an extinct genus from the southeastern United States.[41] Fossil Rubiaceae
Rubiaceae
are known from three regions in the Eocene
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.[41] Uses[edit] Food[edit] Staple foods are not found in the Rubiaceae; instead, 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
Nauclea
latifolia), and noni ( Morinda
Morinda
citrifolia). Beverage[edit] 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
Coffea
includes 124 species, but only three species are cultivated for coffee production: C. arabica, C. canephora, and C. liberica.[6] Medicinal[edit] The bark of trees in the genus Cinchona
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
Galium
odoratum) is a small herbaceous perennial that contains coumarin, a natural precursor of warfarin, and the South American plant Carapichea ipecacuanha
Carapichea ipecacuanha
is the source of the emetic ipecac. Psychotria
Psychotria
viridis is frequently used as a source of dimethyltryptamine in the preparation of ayahuasca, a psychoactive decoction.[42] The bark of the species Breonadia
Breonadia
salicina have been used in traditional African medicine for many years.[43] 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[edit] Originally from China, the common gardenia ( 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
Ixora
contains plants cultivated in warmer-climate gardens; the most commonly grown species, Ixora
Ixora
coccinea, is frequently used for pretty red-flowering hedges. Mussaenda
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
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
Rubia
tinctorum, yields a red dye, and the tropical Morinda citrifolia
Morinda citrifolia
yields a yellow dye. Culture[edit]

Warszewiczia coccinea
Warszewiczia coccinea
is the national flower of Trinidad and Tobago. Coffea
Coffea
arabica is the national flower of Yemen. Cinchona
Cinchona
is the national tree of Ecuador
Ecuador
and Peru. The International Coffee
Coffee
Day is held each year on September 29.

Image gallery[edit]

Alberta magna

Arachnothryx leucophylla

Asperula tinctoria

Bikkia philippinensis

Chiococca alba

Coffea
Coffea
arabica

Galium
Galium
uliginosum

Gardenia
Gardenia
thunbergia

Ixora
Ixora
coccinea

Ixora
Ixora
javanica

Mitragyna speciosa

Morinda
Morinda
pubescens

Nertera granadensis

Psychotria
Psychotria
poeppigiana

Sherardia arvensis

References[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.  ^ Igersheim A, Puff C, Leins P, Erbar C (1994). "Gynoecial development of Gaertnera
Gaertnera
Lam. and of presumably allied taxa of the Psychotrieae (Rubiaceae): secondary 'superior' vs. inferior ovaries". Botanische Jahrbücher 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.  ^ 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.  ^ Anderson WR (1973). "A morphological hypothesis for the origin of heterostyly in the Rubiaceae". Taxon. 22 (5/6): 537–542. doi:10.2307/1218628.  ^ Bridson DM (1987). "Studies in African Rubiaceae-Vanguerieae: a new circumscription of Pyrostria
Pyrostria
and a new subgenus, Canthium
Canthium
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External links[edit]

Wikimedia Commons has media related to Rubiaceae.

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has information related to Rubiaceae

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at Encyclopedia of Life Rubiaceae
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at Angiosperm Phylogeny Website Rubiaceae
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at Flora of China Rubiaceae
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at Flora of Pakistan Rubiaceae
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at Flora of Zimbabwe Rubiaceae
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at Flora of Western Australia Rubiaceae
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at Flora of New Zealand Rubiaceae
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at Integrated Taxonomic Information System Rubiaceae
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at USDA NRCS Plants Database Rubiaceae
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at Botanic Garden Meise World Checklist of Rubiaceae
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at Royal Botanic Gardens, Kew

Taxon identifiers

Wd: Q156569 EoL: 4202 EPPO: 1RUBF FloraBase: 22916 FoC: 10778 Fossilworks: 55784 GBIF: 8798 GRIN: 975 IPNI: 30032120-2 ITIS: 34784 NCBI: 24966 Tropicos: 42000315 VASCAN: 208 Watson & Dallwitz: rubi

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