Difference between revisions of "Rubiaceae"

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{{taxobox
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#redirect [[:Category:Rubiaceae]]
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| name = Rubiaceae
 
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| image = Luculia gratissima.jpg
 
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| image_caption = ''[[Luculia gratissima]]''
 
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| regnum = [[Plantae]]
 
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| unranked_divisio = [[Angiosperms]]
 
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| unranked_classis = [[Eudicots]]
 
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| unranked_ordo = [[Asterids]]
 
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| ordo = [[Gentianales]]
 
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| familia = '''Rubiaceae'''
 
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| familia_authority = [[Antoine Laurent de Jussieu|Juss.]]
 
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| type_genus = ''[[Rubia]]''
 
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| type_genus_authority = [[Carl Linnaeus|L.]]
 
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| subdivision_ranks = [[Subfamilies]]
 
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| subdivision =
 
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* [[Cinchonoideae]]
 
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* [[Ixoroideae]]
 
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* [[Rubioideae]]
 
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| synonyms = see text
 
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}}
 
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The '''Rubiaceae''' are a [[family (biology)|family]] of [[flowering plant]]s, 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 [[Petiole (botany)|interpetiolar]] [[stipules]]. The family contains about 13,500 [[species]] in 611 [[genera]], which makes it the fourth-largest angiosperm family. Rubiaceae has a [[cosmopolitan distribution]], however, the largest species diversity is concentrated in the (sub)tropics.<ref name="Stevens"/> 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]]'').
 
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==Description==
 
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The Rubiaceae are morphologically easily recognizable as a coherent group by a combination of characters: opposite leaves that are simple and entire, [[Petiole (botany)|interpetiolar]] stipules, tubular sympetalous actinomorphic corollas and an inferior ovary.
 
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A wide variety of growth forms are present: [[shrub]]s are most common (e.g. ''[[Coffea]]'', ''[[Psychotria]]''), but members of the family can also be [[tree]]s (e.g. ''[[Cinchona]]'', ''[[Nauclea]]''), [[liana]]s (e.g. ''[[Psychotria samoritourei]]''), or [[herb]]s (e.g. ''[[Galium]]'', ''[[Spermacoce]]''). Some epiphytes are also present (e.g. ''[[Myrmecodia]]''). The plants usually contain [[iridoid]]s, various [[alkaloid]]s, and [[raphide|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 sthenula]]''). Characteristic for the Rubiaceae is the presence of [[stipule]]s 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]] tribe have classically been interpreted as true leaves plus interpetiolar leaf-like stipules. The inflorescence is a [[Inflorescence#Organization|cyme]], rarely of solitary flowers (e.g. ''[[Rothmannia]]''), and is either terminal or axillary and paired at the nodes. The flowers are usually [[Plant reproductive morphology#Flowering plants|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 [[petal#Corolla|corolla]] is sympetalous with four or five lobes, 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]]''). They have four or five [[stamen]]s, which are alternipetalous and epipetalous. [[Anther]]s are longitudinal in dehiscence, but some genera are poricidal (e.g. ''[[Rustia (plant)|Rustia]]''). The [[gynoecium]] is syncarpous with an inferior [[Ovary (botany)|ovary]] (rarely secondarily superior, e.g. ''[[Gaertnera]]'', ''[[Pagamea]]''<ref name="Igersheim94"/>). [[Placentation#Placentation in plants|Placentation]] is axial, rarely parietal (e.g. ''[[Gardenia]]''); [[ovule]]s are anatropous to hemitropous, unitegmic, with a funicular [[Ovary (botany)#Parts of the ovary|obturator]], one to many per carpel. [[Nectar#Floral nectaries|Nectaries]] are often present as a nectariferous disk atop the ovary. The fruit is a [[berry (botany)|berry]], [[Capsule (botany)|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 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 1.0&nbsp;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 [[endosperm]]ous.<ref name="Robbrecht1988"/><ref name ="Takhtajan2009"/>
 
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==Distribution and habitat==
 
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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]] 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]] and [[Cuba]].<ref name="Davis2009"/>
 
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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).
 
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==Ecology==
 
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===Flower biology===
 
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Most Rubiaceae are [[zoophily|zoophilous]]. [[Entomophily|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 magna|Alberta]]'', ''[[Bouvardia]]'', and ''[[Burchellia]]''. [[Anemophily|Anemophylous]] species are found in the tribes [[Anthospermeae]] and [[Theligoneae]] and are characterized by hermaphroditic and/or unisexual flowers that exhibit a set of specialized features, such as striking sexual dimorphism, increased receptive surface of the stigmas and pendulous anthers.<ref name="Robbrecht1988"/>
 
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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.<br />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 and/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]].<ref name="Delprete2009"/><br />Heterodistyly is another mechanism to avoid inbreeding and is widely present in the Rubiaceae family.<ref name="Anderson1973"/> 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]]).<br />Unisexual flowers also occur in Rubiaceae and most taxa that have this characteristic are [[dioecy|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).<ref name="Robbrecht1988"/> Flowers that are morphologically hermaphrodite, but functionally dioecious are for example found in ''[[Pyrostria]]''.<ref name="Bridson1987"/>
 
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=== Fruit biology ===
 
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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 [[Diaspore (botany)|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 (plant)|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 [[elaiosome]]s 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).
 
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=== Associations with other organisms ===
 
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The genera ''[[Anthorrhiza]]'', ''[[Hydnophytum]]'', ''[[Myrmecodia]]'', ''[[Myrmephytum]]'', and ''[[Squamellaria]]'' are succulent [[epiphyte]]s that have evolved a [[mutualism (biology)|mutualistic]] relationship with ants. Their [[hypocotyl]] grows out into an ant-inhabited tuber.<ref name="Kapitany2007"/> Some shrubs or trees have ant holes in their stems (e.g. ''[[Globulostylis]]'').<ref name="Verstraete2013b"/> Some Rubiaceae species have domatia that are inhabited by mites (viz. [[acarodomatia]]; e.g. ''[[Plectroniella armata]]'').<ref name="Tilney2012"/><br />An intimate association between bacteria and plants is found in three rubiaceous genera (viz. ''[[Pavetta]]'', ''[[Psychotria]]'', and ''[[Sericanthe]]'').<ref name="Lemaire2011"/> 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 [[Vanguerieae]] tribe), where ''[[Burkholderia]]'' bacteria are found freely distributed among the mesophyll cells and no leaf nodules are formed.<ref name="Verstraete2011"/><ref name="Verstraete2013a"/><ref name="Verstraete2013c"/> The hypothesis regarding the function of the symbiosis is that the endophytes provide chemical protection against herbivory by producing certain toxic secondary metabolites.<ref name="Sieber2015"/>
 
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==Systematics==
 
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The Rubiaceae family is named after ''[[Rubia]]'', a name used by [[Pliny the Elder]] in his [[Natural History (Pliny)|Naturalis Historia]] for madder (''[[Rubia tinctorum]]'').<ref name="Simpson2006"/> 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.
 
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===Taxonomy===
 
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The name Rubiaceae ([[nomen conservandum]]) was published in 1789 by [[Antoine Laurent de Jussieu]],<ref name=Jussieu1789/> but the name was already mentioned in 1782.<ref name=Durande1782/>
 
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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.<ref name="Stevens"/><br />More recently, the morphologically quite different families Dialypetalanthaceae,<ref name="Fay2000"/> Henriqueziaceae,<ref name="Rogers1981"/> and Theligonaceae<ref name="Wunderlich1971"/><ref name="Rutishauser1998"/><ref name="Robbrecht2006"/> were reduced to synonymy of Rubiaceae.
 
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====Subfamilies and tribes====
 
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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.<ref name ="Hooker1873"/><ref name ="Schumann1891"/> 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.<ref name ="Baillon1878"/><ref name ="Solereder1893"/> During the twentieth century other characters were used to delineate subfamilies, e.g. stylar pollen presentation, [[raphide]]s, [[endosperm]], [[heterostyly]], etc.  On this basis, three<ref name ="Verdcourt1958"/> or eight<ref name ="Bremekamp1966"/> subfamilies were recognised. The last subfamilial classification solely based on morphological characters divided Rubiaceae into four subfamilies: Cinchonoideae, Ixoroideae, Antirheoideae, and Rubioideae.<ref name="Robbrecht1988"/> 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.<ref name="Robbrecht1988"/>
 
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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]].<ref name="Bremer1995"/> 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.<ref name ="Rydin2009"/> Currently, in most molecular research concerning the Rubiaceae family, the classification with three subfamilies (Cinchonoideae, Ixoroideae, and Rubioideae) is followed.<ref name ="Bremer2009"/> However, an alternative view is proposed where only two subfamilies are recognized, an expanded Cinchonoideae (that includes Ixoroideae, Coptosapeltaeae and Luculieae) and Rubioideae.<ref name="Robbrecht2006"/> The adoption of the [[International Code of Nomenclature for algae, fungi, and plants|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.<ref name="Reveal2012"/> 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.<ref name="Kainulainen2013"/><ref name="Mouly2014"/> 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,<ref name="WCSP"/> however, several genera and species are not yet placed in a tribe.
 
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{|
 
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|- valign=top
 
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* '''basal Rubiaceae''' (59 sp.)
 
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** [[Coptosapelteae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (55 sp.)
 
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** [[Luculieae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (4 sp.)
 
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|
 
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* '''[[Cinchonoideae]]''' <small>[[Constantine Samuel Rafinesque|Raf.]]</small> (1708 sp.)
 
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** [[Chiococceae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (224 sp.)
 
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** [[Cinchoneae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (125 sp.)
 
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** [[Guettardeae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (747 sp.)
 
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** [[Hamelieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (171 sp.)
 
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** [[Hymenodictyeae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]]</small> (25 sp.)
 
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** [[Hillieae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (29 sp.)
 
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** [[Isertieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (16 sp.)
 
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** [[Naucleeae]] <small>[[Augustin Pyramus de Candolle|DC.]] ex [[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (192 sp.)
 
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** [[Rondeletieae]] <small>[[Augustin Pyramus de Candolle|DC.]] ex [[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (178 sp.)
 
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** [[Strumpfieae]] <small>Delprete & T.J.Motley</small> (1 sp.)
 
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* '''[[Ixoroideae]]''' <small>[[Constantine Samuel Rafinesque|Raf.]]</small> (3960 sp.)
 
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** [[Airospermeae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (7 sp.)
 
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** [[Alberteae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (7 sp.)
 
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** [[Aleisanthieae]] <small>Mouly, J.Florence & [[Birgitta Bremer|B.Bremer]]</small> (10 sp.)
 
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** [[Augusteae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (86 sp.)
 
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** [[Bertiereae]] <small>[[Diane Mary Bridson|Bridson]]</small> (57 sp.)
 
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** [[Coffeeae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (303 sp.)
 
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** [[Condamineeae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (305 sp.)
 
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** [[Cordiereae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]] emend. Mouly</small> (124 sp.)
 
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** [[Cremasporeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (2 sp.)
 
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** [[Crossopterygeae]] <small>F.White ex [[Diane Mary Bridson|Bridson]]</small> (1 sp.)
 
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** [[Gardenieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (587 sp.)
 
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** [[Greeneeae]] <small>Mouly, J.Florence & [[Birgitta Bremer|B.Bremer]]</small> (9 sp.)
 
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** [[Henriquezieae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (20 sp.)
 
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** [[Ixoreae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (545 sp.)
 
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** [[Jackieae]] <small>[[Pieter Willem Korthals|Korth.]]</small> (1 sp.)
 
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** [[Mussaendeae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (221 sp.)
 
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** [[Octotropideae]] <small>[[Richard Henry Beddome|Bedd.]]</small> (96 sp.)
 
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** [[Pavetteae]] <small>[[Achille Richard|A.Rich.]] ex [[Barthélemy Charles Joseph Dumortier|Dumort.]]</small> (624 sp.)
 
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** [[Posoquerieae]] <small>Delprete</small> (23 sp.)
 
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** [[Retiniphylleae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (20 sp.)
 
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** [[Sabiceeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (164 sp.)
 
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** [[Scyphiphoreae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (1 sp.)
 
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** [[Sherbournieae]] <small>Mouly & [[Birgitta Bremer|B.Bremer]]</small> (54 sp.)
 
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** [[Sipaneeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (43 sp.)
 
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** [[Steenisieae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (5 sp.)
 
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** [[Trailliaedoxeae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (1 sp.)
 
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** [[Vanguerieae]] <small>[[Achille Richard|A.Rich.]] ex [[Barthélemy Charles Joseph Dumortier|Dumort.]]</small> (644 sp.)
 
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|
 
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* '''[[Rubioideae]]''' <small>[[Bernard Verdcourt|Verdc.]]</small> (7571 sp.)
 
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** [[Anthospermeae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (208 sp.)
 
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** [[Argostemmateae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (215 sp.)
 
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** [[Clarkelleae]] <small>Deb</small> (1 sp.)
 
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** [[Colletoecemateae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (3 sp.)
 
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** [[Coussareeae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (402 sp.)
 
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** [[Craterispermeae]] <small>[[Bernard Verdcourt|Verdc.]]</small> (16 sp.)
 
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** [[Cyanoneuroneae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]]</small> (5 sp.)
 
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** [[Danaideae]] <small>[[Birgitta Bremer|B.Bremer]] & Manen</small> (50 sp.)
 
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** [[Dunnieae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (1 sp.)
 
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** [[Gaertnereae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (95 sp.)
 
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** [[Knoxieae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (131 sp.)
 
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** [[Lasiantheae]] <small>[[Birgitta Bremer|B.Bremer]] & Manen</small> (239 sp.)
 
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** [[Mitchelleae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]] & Manen</small> (14 sp.)
 
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** [[Morindeae]] <small>[[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (165 sp.)
 
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** [[Ophiorrhizeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (364 sp.)
 
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** [[Paederieae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (81 sp.)
 
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** [[Palicoureeae]] <small>Robbr. & Manen</small> (817 sp.)
 
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** [[Perameae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (14 sp.)
 
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** [[Prismatomerideae]] <small>Y.Z.Ruan</small> (23 sp.)
 
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** [[Psychotrieae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]]</small> (2114 sp.)
 
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** [[Putorieae]] (34 sp.)
 
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** [[Rubieae]] <small>[[Henri Ernest Baillon|Baill.]]</small> (938 sp.)
 
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** [[Schizocoleeae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (2 sp.)
 
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** [[Schradereae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (55 sp.)
 
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** [[Spermacoceae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (1344 sp.)
 
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** [[Theligoneae]] <small>Wunderlich ex S.P.Darwin</small> (4 sp.)
 
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** [[Urophylleae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (236 sp.)
 
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|}
 
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====Genera====
 
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{{For|a comprehensive list|List of Rubiaceae genera}}
 
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The Rubiaceae family contains about 13,500 species in 611 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 1300 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]]'' 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, 136 genera are [[monotypic]], which account for 22% of all genera, but only for 1.1% of all species.<ref name ="Davis2009"/>
 
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===Phylogeny===
 
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Molecular studies have demonstrated the phylogenetic placement of Rubiaceae within the order [[Gentianales]] and the [[monophyly]] of the family is confirmed.<ref name="Bremer2002"/><ref name="BremerEriksson2009"/> 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.<ref name="BremerEriksson2009"/>
 
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{{clade
 
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|label1='''Rubiaceae'''
 
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|1={{clade
 
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|1=[[Rubioideae]]
 
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|2={{clade
 
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  |1=[[Ixoroideae]]
 
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  |2=[[Cinchonoideae]]
 
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  }}
 
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|3={{clade
 
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  |1=[[Coptosapelteae]]
 
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  |2=[[Luculieae]]
 
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  }}
 
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}}
 
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}}
 
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===Evolution===
 
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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.<ref name="Graham2009"/>
 
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The oldest confirmed fossils, which are fruits that strongly resemble those of the genus ''[[Emmenopterys]]'', were found in the [[Washington (state)|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]].<ref name ="Graham2009"/>
 
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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.<ref name ="Graham2009"/>
 
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==Uses==
 
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===Food===
 
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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. ''[[Vangueria infausta|V. infausta]]'', ''[[Vangueria madagascariensis|V. madagascariensis]]''), African peach (''[[Nauclea latifolia]]''), and noni (''[[Morinda citrifolia]]'').
 
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===Beverage===
 
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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 124 species, but only three species are cultivated for coffee production: ''[[Coffea arabica|C. arabica]]'', ''[[Coffea canephora|C. canephora]]'', and ''[[Coffea liberica|C. liberica]]''.<ref name ="Davis2009"/>
 
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===Medicinal===
 
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The bark of trees in the genus ''[[Cinchona]]'' is the source of a variety of [[alkaloid]]s, 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]] [[Syrup of ipecac|ipecac]]. ''[[Psychotria viridis]]'' is frequently used as a source of [[dimethyltryptamine]] in the preparation of [[ayahuasca]], a psychoactive decoction.<ref name="Riba2003"/> The bark of the species ''[[Breonadia]] salicina'' have been used in traditional African medicine for many years.<ref name=Neuwinger94>{{cite book|last1=Neuwinger|first1=Hans Dieter|title=African Ethnobotany: Poisons and Drugs: Chemistry, Pharmacology, Toxicology|date=1994|publisher=Chapman & Hall|location=Stuttgart, Germany|accessdate=10 December 2015}}</ref> 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]].
 
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===Ornamentals===
 
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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 [[hedge]]s. 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]]''.
 
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===Dyes===
 
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[[Rose madder]], the crushed root of ''[[Rubia tinctorum]]'', yields a red dye, and the tropical ''[[Morinda citrifolia]]'' yields a yellow dye.
 
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==Culture==
 
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* ''[[Warszewiczia coccinea]]'' is the national flower of [[Trinidad and Tobago]].
 
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* ''Coffea arabica'' is the national flower of [[Yemen]].
 
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* ''Cinchona'' is the national tree of [[Ecuador]] and [[Peru]].
 
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* The [[International Coffee Day]] is held each year on September 29.
 
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==Image gallery==
 
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<gallery>
 
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File:Alberta magna.jpg|''[[Alberta magna]]''
 
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File:IMG 7387-Arachnothryx leucophylla.jpg|''[[Arachnothryx leucophylla]]''
 
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File:Marzanka barwierska - Asperula tinctoria 01.jpg|''[[Asperula tinctoria]]''
 
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File:Bikkia philippinensis.jpg|''[[Bikkia philippinensis]]''
 
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File:Chiococca alba.jpg|''[[Chiococca alba]]''
 
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File:Coffee Flowers.JPG|''[[Coffea arabica]]''
 
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File:Galium uliginosum W.jpg|''[[Galium uliginosum]]''
 
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File:Gardenia thunbergia.JPG|''[[Gardenia thunbergia]]''
 
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File:Ixora coccinea- jungle geranium.JPG|''[[Ixora coccinea]]''
 
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File:Soka-Ixora javanica1.jpg|''[[Ixora javanica]]''
 
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File:Mitragyna speciosa111.JPG|''[[Mitragyna speciosa]]''
 
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File:Morinda pubescens in Ananthagiri forest, AP W IMG 9225.jpg|''[[Morinda pubescens]]''
 
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File:Nertera depressa 1.jpg|''[[Nertera granadensis]]''
 
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File:Psychotria poeppigiana (bracts).jpg|''[[Psychotria poeppigiana]]''
 
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File:Sherardia arvensis Kaldari 01.jpg|''[[Sherardia arvensis]]''
 
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</gallery>
 
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==References==
 
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{{reflist|refs=
 
−
<ref name="Stevens">{{cite web|url=http://www.mobot.org/MOBOT/research/APweb|title=Angiosperm Phylogeny Website|accessdate=1 June 2014}}</ref>
 
−
<ref name="Igersheim94">{{cite journal|vauthors=Igersheim A, Puff C, Leins P, Erbar C |year=1994|title=Gynoecial development of ''Gaertnera'' Lam. and of presumably allied taxa of the Psychotrieae (Rubiaceae): secondary 'superior' vs. inferior ovaries|journal=Botanische Jahrbücher fur Systematik|volume=116|pages=401–414}}</ref>
 
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<ref name="Robbrecht1988">{{cite journal|author=Robbrecht E|year=1988|title=Tropical woody Rubiaceae|journal=Opera Botanica Belgica|volume=1|pages=1–271}}</ref>
 
−
<ref name ="Takhtajan2009">{{cite book|author=Takhtajan A|year=2009|title=Flowering Plants|edition=2|publisher=Springer|ISBN=978-1-4020-9608-2}}</ref>
 
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<ref name="Davis2009">{{cite journal|vauthors=Davis AP, Govaerts R, Bridson DM, Ruhsam M, Moat J, Brummitt NA |year=2009|title=A global assessment of distribution, diversity, endemism, and taxonomic effort in the Rubiaceae|journal=Annals of the Missouri Botanical Garden|volume=96|issue=1|pages=68–78| doi=10.3417/2006205}}</ref>
 
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<ref name="Delprete2009">{{cite journal|author=Delprete PG|year=2009|title=Taxonomic history, morphology, and reproductive biology of the tribe Posoquerieae (Rubiaceae, Ixoroideae)|journal=Annals of the Missouri Botanical Garden|volume=96|issue=1|pages=79–89| doi=10.3417/2006192}}</ref>
 
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<ref name="Anderson1973">{{cite journal|author=Anderson WR|year=1973|title=A morphological hypothesis for the origin of heterostyly in the Rubiaceae|journal=Taxon|volume=22|issue=5/6|pages=537–542|doi=10.2307/1218628}}</ref>
 
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<ref name="Bridson1987">{{cite journal|author=Bridson DM|year=1987|title=Studies in African Rubiaceae-Vanguerieae: a new circumscription of ''Pyrostria'' and a new subgenus, ''Canthium'' subgen. ''Bullockia''|journal=Kew Bulletin|volume=42|pages=611–639|doi=10.2307/4110068}}</ref>
 
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<ref name="Kapitany2007">{{cite book|author=Kapitany A|title=Australian succulent plants: an introduction|publisher=Kapitany Concepts|location=Boronia, Victoria|year=2007|pages=144–155|isbn=0-646-46381-0}}</ref>
 
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<ref name="Verstraete2013b">{{cite journal|vauthors=Verstraete B, Lachenaud O, Smets E, Dessein S, Sonké B |year=2013|title=Taxonomy and phylogeny of ''Cuviera'' (Rubiaceae-Vanguerieae) and reinstatement of the genus ''Globulostylis'' with the description of three new species|journal=Botanical Journal of the Linnean Society|volume=173|issue=3|pages=407–441|doi=10.1111/boj.12062}}</ref>
 
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<ref name="Tilney2012">{{cite journal|vauthors=Tilney PM, van Wyk AE, van deer Merwe CF |year=2012|title=Structural evidence in ''Plectroniella armada'' (Rubiaeae) for possible material exchange between domatia and mites|journal=PLoS ONE|volume=7|issue=7|pages=e39984|doi=10.1371/journal.pone.0039984|pmid=22792206|pmc=3390328}}</ref>
 
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<ref name="Lemaire2011">{{cite journal|vauthors=Lemaire B, Vandamme P, Merckx V, Smets E, Dessein S |year=2011|title=Bacterial leaf symbiosis in angiosperms: host specificity without co-speciation|journal=PLoS ONE|volume=6|issue=9|pages=e24430|doi=10.1371/journal.pone.0024430|pmid=21915326|pmc=3168474}}</ref>
 
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<ref name="Verstraete2011">{{cite journal|vauthors=Verstraete B, Van Elst D, Steyn H, Van Wyk B, Lemaire B, Smets E, Dessein S |year=2011|title=Endophytic bacteria in toxic South African plants: identification, phylogeny and possible involvement in gousiekte|journal=PLoS ONE|volume=6|issue=4|pages=e19265|doi=10.1371/journal.pone.0019265|pmid=21541284|pmc=3082559}}</ref>
 
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<ref name="Verstraete2013a">{{cite journal|vauthors=Verstraete B, Janssens S, Smets E, Dessein S |year=2013|title=Symbiotic beta-proteobacteria beyond legumes: ''Burkholderia'' in Rubiaceae|journal=PLoS ONE|volume=8|issue=1|pages=e55260|doi=10.1371/journal.pone.0055260|pmid=23372845|pmc=3555867}}</ref>
 
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<ref name="Verstraete2013c">{{cite journal|vauthors=Verstraete B, Janssens S, Lemaire B, Smets E, Dessein S |year=2013|title=Phylogenetic lineages in Vanguerieae (Rubiaceae) associated with ''Burkholderia'' bacteria in sub-Saharan Africa|journal=American Journal of Botany|volume=100|issue=|pages=2380–2387|doi=10.3732/ajb.1300303|pmid=24275705}}</ref>
 
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<ref name="Sieber2015">{{cite journal|author=Sieber S, Carlier AL, Neuburger M, Grabenweger G, Eberl L, Gademann K|year=2015|title=Isolation and total synthesis of kirkamide, an aminocyclitol from an obligate leaf nodule symbiont|journal=Angewandte Chemie - International Edition|volume=54|pages=7968–7970|doi=10.1002/anie.201502696}}</ref>
 
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<ref name="Simpson2006">{{cite book|author=Simpson MG|year=2006|title=Plant Systematics|edition=1|publisher=Elsevier Academic Press|ISBN=978-0-12-644460-5}}</ref>
 
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<ref name=Jussieu1789>{{cite book|author=Jussieu A L de|year=1789|title=Genera Plantarum|page=206|publisher=Herissant & Barrois|location=Paris|url= http://biodiversitylibrary.org/page/5437428}}</ref>
 
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<ref name=Durande1782>{{cite book|author=Durand JF|year=1782|title=Notions Élémentaires de Botanique |page=274|publisher=LN Frantin|location=Dijon}}</ref>
 
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<ref name="Fay2000">{{cite journal|vauthors=Fay MF, Bremer B, Prance GT, van der Bank M, Bridson D, Chase MW |year=2000|title=Plastid ''rbcL'' sequence data show Dialypetalanthus to be a member of Rubiaceae|journal=Kew Bulletin|volume=55|issue=4|pages=853–864|doi=10.2307/4113630}}</ref>
 
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<ref name="Rogers1981">{{cite journal|author=Rogers GK|year=1981|title=The wood of ''Gleasonia'', ''Henriquezia'', and ''Platycarpum'' (Rubiaceae) and its bearing on their classification: some new considerations|journal=Brittonia|volume=33|issue=3|pages=461–465|doi=10.2307/2806441}}</ref>
 
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<ref name="Wunderlich1971">{{cite journal|author=Wunderlich R|year=1971|title=Die systematische Stellung von ''Theligonum''|journal=Österreichische botanische Zeitschrift |volume=119|pages=329–394|doi=10.1007/bf01377490}}</ref>
 
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<ref name="Rutishauser1998">{{cite journal|vauthors=Rutishauser F, Ronse Decraene LP, Smets E, Mendoza-Heuer I |year=1998|title=''Theligonum cynocrambe'': developmental morphology of a peculiar rubiaceous herb|journal=Plant Systematics and Evolution|volume=210|issue=1|pages=1–24|doi=10.1007/BF00984724}}</ref>
 
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<ref name="Robbrecht2006">{{cite journal|vauthors=Robbrecht E, Manen JF |year=2006|title=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|journal=Systematic Geography of Plants|volume=76|pages=85–146}}</ref>
 
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<ref name ="Hooker1873">{{cite book|author=Hooker JD|veditors=Bentham G, Hooker JD |title=Genera planetarium ad exemplaria imprimis in herbaria kewensibus servata defirmata|volume=2|location=London|date=1873|pages=7–151|chapter=Ordo LXXXIV. Rubiaceae}}</ref>
 
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<ref name ="Schumann1891">{{cite book|author=Schumann K|veditors=Engler A, Prantl K |title=Die natürlichen Pflanzenfamilien|volume=4|issue=4|publisher=Engelmann|location=Leipzig|date=1891|pages=1–156|chapter=Rubiaceae}}</ref>
 
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<ref name ="Baillon1878">{{cite journal|author=Baillon H|year=1878|title=Sur les limits du genre ''Ixora''|journal=Adansonia|volume=12|pages=213–219}}</ref>
 
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<ref name ="Solereder1893">{{cite journal|author=Solereder H|year=1893|title=Ein Beitrag zur anatomischen Charakteristik und zur Systematik deer Rubiaceen|journal=Bull. Herb. Boissier|volume=1|pages=167–183}}</ref>
 
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<ref name ="Verdcourt1958">{{cite journal|author=Verdcourt B|year=1958|title=Remarks on the classification of the Rubiaceae|journal=Bulletin du Jardin botanique de l'état, Bruxelles|volume=28|pages=209–281|doi=10.2307/3667090}}</ref>
 
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<ref name ="Bremekamp1966">{{cite journal|author=Bremekamp CEB|year=1966|title=Remarks on the position, the delimitation and the subdivision of the Rubiaceae|journal=Acta Botanica Neerlandica|volume=15|pages=1–33|doi=10.1111/j.1438-8677.1966.tb00207.x}}</ref>
 
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<ref name="Bremer1995">{{cite journal|vauthors=Bremer B, Andreasen K, Olsson D |year=1995|title=Subfamilial and tribal relationships in the Rubiaceae based on rbcL sequence data|journal=Annals of the Missouri Botanical Garden|volume=82|pages=383–397|doi=10.2307/2399889}}</ref>
 
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<ref name ="Rydin2009">{{cite journal|vauthors=Rydin C, Kainulainen K, Razafimandimbison SG, Smedmark JE, Bremer B |year=2009|title=Deep divergences in the coffee family and the systematic position of ''Acranthera''|journal=Plant Systematics and Evolution|volume=278|pages=101–123|doi=10.1007/s00606-008-0138-4}}</ref>
 
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<ref name ="Bremer2009">{{cite journal|author=Bremer B|year=2009|title=A review of molecular phylogenetic studies of Rubiaceae|journal=Annals of the Missouri Botanical Garden|volume=96|issue=1|pages=4–26|doi=10.3417/2006197}}</ref>
 
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<ref name="Kainulainen2013">{{cite journal|vauthors=Kainulainen K, Razafimandimbison SG, Bremer B |year=2013|title=Phylogenetic relationships and new tribal delimitations in subfamily Ixoroideae (Rubiaceae)|journal=Botanical Journal of the Linnean Society|volume=173|issue=3|pages=387–406|doi=10.1111/boj.12038}}</ref>
 
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<ref name="Mouly2014">{{cite journal|vauthors=Mouly A, Kainulainen K, Persson C, Davis AP, Wong KM, Razafimandimbison SG, Bremer B |year=2014|title=Phylogenetic structure and clade circumscriptions in the Gardenieae complex (Rubiaceae)|journal=Taxon|volume=63|issue=4|pages=801–818|doi=10.12705/634.4}}</ref>
 
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<ref name="Bremer2002">{{cite journal|vauthors=Bremer B, Bremer K, Heidari N, Erixon P, Olmstead RG, Anderberg AA, Källersjö M, Barkhordarian E |year=2002|title=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|journal=Molecular Phylogenetics and Evolution|volume=24|pages=274–301|doi=10.1016/s1055-7903(02)00240-3|pmid=12144762}}</ref>
 
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<ref name="BremerEriksson2009">{{cite journal|vauthors=Bremer B, Eriksson T |year=2009|title=Time tree of Rubiaceae: phylogeny and dating the family, subfamilies, and tribes|journal=International Journal of Plant Sciences|volume=170|pages=766–793|doi=10.1086/599077}}</ref>
 
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<ref name="Graham2009">{{cite journal|author=Graham A|year=2009|title=Fossil record of the Rubiaceae|journal=Annals of the Missouri Botanical Garden|volume=96|issue=1|pages=90–108|doi=10.3417/2006165}}</ref>
 
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<ref name="Riba2003">{{cite journal|vauthors=Riba J, Valle M, Urbano G, Yritia M, Morte A, Barbanoj MJ |year=2003|title=Human pharmacology of ayahuasca: subjective and cardiovascular effects, monoamine metabolite excretion, and pharmacokinetics|journal=Journal of Pharmacology and Experimental Therapeutics|pmid=12660312|volume=306|issue=1|pages=73–83|doi=10.1124/jpet.103.049882}}</ref>
 
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<ref name="Reveal2012">{{cite journal|author=Reveal JL|year=2012|title=Newly required infrafamilial names mandated by changes in the code of nomenclature for algae, fungi and plants|journal=Phytoneuron|volume=33|pages=1–32}}</ref>
 
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<ref name="WCSP">{{cite web|url=http://apps.kew.org/wcsp/home.do|title=World Checklist of Rubiaceae|accessdate=1 March 2016}}</ref>
 
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}}
 
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==External links==
 
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{{commons category|Rubiaceae}}
 
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{{wikispecies|Rubiaceae}}
 
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* [http://www.theplantlist.org/browse/A/Rubiaceae Rubiaceae] at [http://www.theplantlist.org ''The Plant List'']
 
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* [http://eol.org/pages/4202/overview Rubiaceae] at [http://eol.org ''Encyclopedia of Life'']
 
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* [http://www.mobot.org/mobot/research/APweb/orders/gentianalesweb.htm#Rubiaceae Rubiaceae] at [http://www.mobot.org/mobot/research/apweb ''Angiosperm Phylogeny Website'']
 
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* [http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=10778 Rubiaceae] at [http://www.efloras.org/flora_page.aspx?flora_id=2 ''Flora of China'']
 
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* [http://www.efloras.org/florataxon.aspx?flora_id=5&taxon_id=10778 Rubiaceae] at [http://www.efloras.org/flora_page.aspx?flora_id=5 ''Flora of Pakistan'']
 
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* [http://www.zimbabweflora.co.zw/speciesdata/family.php?family_id=198 Rubiaceae] at [http://www.zimbabweflora.co.zw ''Flora of Zimbabwe'']
 
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* [http://florabase.dec.wa.gov.au/browse/profile.php/22916 Rubiaceae] at [http://florabase.dec.wa.gov.au ''Flora of Western Australia'']
 
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* [http://floraseries.landcareresearch.co.nz/pages/Taxon.aspx?id=_22034db9-2fe7-40b1-8b2d-3633d3d683dc&fileName=Flora%201.xml Rubiaceae] at [http://floraseries.landcareresearch.co.nz ''Flora of New Zealand'']
 
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* [http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=34784 Rubiaceae] at [http://www.itis.gov ''Integrated Taxonomic Information System'']
 
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* [http://plants.usda.gov/java/ClassificationServlet?source=display&classid=Rubiaceae Rubiaceae] at [http://plants.usda.gov ''USDA NRCS Plants Database'']
 
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* [http://www.br.fgov.be/RESEARCH/PROJECTS/rubiaceae.php Rubiaceae] at [http://www.br.fgov.be ''Botanic Garden Meise'']
 
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* [http://apps.kew.org/wcsp/rubiaceae World Checklist of Rubiaceae] at [http://www.kew.org ''Royal Botanic Gardens, Kew'']
 
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{{taxonbar}}
 
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[[Category:Asterid families]]
 
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[[Category:Rubiaceae| ]]
 
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[[Category:Extant Eocene first appearances]]
 

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