Editing Rubiaceae

{{Short description|Family of flowering plants}}
{{Distinguish|text=[[Rubiácea]], a town in Brazil}}
{{Technical|date=March 2025}}
{{Automatic taxobox
| image = Luculia gratissima.jpg
| image_caption = ''[[Luculia gratissima]]''
| taxon = Rubiaceae
| authority = [[Antoine Laurent k de Jussieu|Juss.]]
| type_genus = ''[[Rubia]]''
| type_genus_authority = [[Carl Linnaeus|L.]]
| subdivision_ranks = Subfamilies
| subdivision =
* [[Dialypetalanthoideae]]
* [[Rubioideae]]
| synonyms = See text
}}

'''Rubiaceae''' ({{IPAc-en|r|uː|b|i|ˈ|eɪ|s|iː|ˌ|iː|,_|-|s|i|ˌ|aɪ}}) is a [[family (biology)|family]] of [[flowering plant]]s, commonly known as the '''coffee''', '''madder''', or '''bedstraw family'''. It consists of terrestrial trees, shrubs, [[liana]]s, or herbs that are recognizable by simple, opposite leaves with [[Petiole (botany)|interpetiolar]] [[stipules]] and [[sympetalous]] [[actinomorphic]] flowers. The family contains about 14,100 [[species]] in about 580 [[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]].<ref name="Stevens"/> 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, [[Petiole (botany)|interpetiolar]] stipules, tubular [[Sympetalae|sympetalous]] [[actinomorphic]] [[Corolla (botany)|corollas]] and an [[inferior ovary]].

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 [[herbaceous plant|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; there is only one case of pinnately compound leaves (''[[Pentagonia osapinnata]]''<ref name ="Hammel2015"/>). 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 [[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 tribe [[Rubieae]] 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 4-5-merous (rarely pleiomerous; e.g. six in ''[[Richardia]]''<ref name ="Hall1991"/>) 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 [[Sepal|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 [[petal#Corolla|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 [[stamen]]s are alternipetalous and epipetalous. [[Anther]]s are longitudinal in dehiscence, but are poricidal in some genera (e.g. ''[[Rustia (plant)|Rustia]]'', ''[[Tresanthera]]''). 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. ''[[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&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"/>

==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]].<ref name="Davis2009"/>

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 members of the Rubiaceae are [[zoophily|zoophilous]], pollinated mainly by insects. [[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|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 [[stigma (botany)|stigmas]] and [[pendulous]] [[anthers]].<ref name="Robbrecht1988"/>

Although most Rubiaceae species are hermaphroditic, [[outbreeding]] is promoted through [[proterandry|sequential hermaphroditism]] and spatial isolation of the reproductive organs. More complex reproductive strategies include secondary pollen presentation, [[heterostyly]], 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 pollen receptacle. 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|hawk moths]].<ref name="Delprete2009"/>

Heterostyly is another mechanism to avoid inbreeding and is widely present in the family Rubiaceae.<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]]).

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|rudimentary pistil]] with the ovaries empty and female flowers [[staminode|sterile or rudimentary stamens]] with empty anthers.<ref name="Robbrecht1988"/> Flowers that are morphologically hermaphrodite, but functionally dioecious occur in ''[[Pyrostria]]''.<ref name="Bridson1987"/>

=== 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 [[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).

=== Associations with other organisms ===
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"/>

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]]'', and ''[[Vangueria]]'' (all belonging to the tribe [[Vanguerieae]]),<ref name="Verstraete2011"/><ref name="Verstraete2013a"/><ref name="Verstraete2013c"/> and in some species of ''[[Empogona]]'' and ''[[Tricalysia]]'' (both belonging to the tribe [[Coffeeae]]),<ref name="Verstraete2023"/> where ''[[Burkholderia]]'' bacteria are found freely distributed among the mesophyll cells and no leaf nodules are formed. 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"/>

==Systematics==
The family Rubiaceae 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.

===Taxonomy===
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/>

Several historically accepted families are included in Rubiaceae: Aparinaceae, Asperulaceae, Catesbaeaceae, Cephalanthaceae, Cinchonaceae, Coffeaceae, Coutariaceae, Dialypetalanthaceae,<ref name="Fay2000"/> Galiaceae, Gardeniaceae, Guettardaceae, Hameliaceae, Hedyotidaceae, Henriqueziaceae,<ref name="Rogers1981"/> Houstoniaceae, Hydrophylacaceae, Lippayaceae, Lygodisodeaceae, Naucleaceae, Nonateliaceae, Operculariaceae, Pagamaeaceae, Psychotriaceae, Randiaceae, Sabiceaceae, Spermacoceaceae, Theligonaceae.<ref name="Wunderlich1971"/><ref name="Rutishauser1998"/><ref name="Robbrecht2006"/><ref name="Stevens"/><ref name="RazaRydin2024"/>

====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.<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 20th century, other morphological 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"/>

The introduction of molecular phylogenetics in Rubiaceae research has corroborated or rejected several of the conclusions made in the pre-molecular era. There was support for the subfamilies Cinchonoideae, Ixoroideae, and Rubioideae, although differently circumscribed, and Antirheoideae was shown to be [[polyphyletic]].<ref name="Bremer1995"/> For a long time, the classification with three subfamilies (Cinchonoideae, Ixoroideae, and Rubioideae) was followed.<ref name ="Bremer2009"/> However, an alternative opinion existed with only two subfamilies: an expanded Cinchonoideae (that includes Ixoroideae, Coptosapelteae, and Luculieae) and Rubioideae.<ref name="Robbrecht2006"/> Finally, more and more evidence pointed towards a two-family classification.<ref name ="RazaRydin2024"/> 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 had to be replaced by Dialypetalanthoideae.<ref name="Reveal2012"/> This means that the two subfamilies in Rubiaceae now are: Dialypetalanthoideae and Rubioideae.<ref name ="RazaRydin2024"/> The monogeneric tribes [[Coptosapelteae]], [[Acranthereae]], and [[Luculieae]] are not placed within a subfamily and are sister to the rest of Rubiaceae.<ref name ="Rydin2009"/><ref name ="RazaRydin2024"/> The following overview shows the latest classification of the family, with two subfamilies and 71 tribes.<ref name ="RazaRydin2024"/> The approximate number of species and genera are indicated between brackets (species/genera).

;Unplaced tribes (76/3)
* [[Acranthereae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (16/1)
* [[Coptosapelteae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (56/1)
* [[Luculieae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (4/1)

=====[[Rubioideae]]=====
Authority: [[Bernard Verdcourt|Verdc.]]; over 8000 spp.
{{div col|colwidth=20em}}
* [[Aitchisonieae]] <small>Bordbar, Razafim. & Mirtadz.</small> (1/1)
* [[Anthospermeae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (210/12)
* [[Argostemmateae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (251/6)
* [[Colletoecemateae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (3/1)
* [[Coussareeae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (414/10)
* [[Craterispermeae]] <small>[[Bernard Verdcourt|Verdc.]]</small> (32/1)
* [[Cyanoneuroneae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]]</small> (5/1)
* [[Danaideae]] <small>[[Birgitta Bremer|B.Bremer]] & Manen</small> (67/3)
* [[Dunnieae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (1/1)
* [[Foonchewieae]] <small>R.J.Wang</small> (2/1)
* [[Gaertnereae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (116/2)
* [[Knoxieae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (130/16)
* [[Lasiantheae]] <small>[[Birgitta Bremer|B.Bremer]] & Manen</small> (291/4)
* [[Mitchelleae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]] & Manen</small> (16/2)
* [[Morindeae]] <small>[[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (163/5)
* [[Seychelleeae]] <small>Razafim., Kainul. & Rydin</small> (1/1)
* [[Ophiorrhizeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (420/6)
* [[Paederieae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (98/6)
* [[Palicoureeae]] <small>Robbr. & Manen</small> (1112/9)
* [[Perameae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (14/1)
* [[Prismatomerideae]] <small>Y.Z.Ruan</small> (27/2)
* [[Psychotrieae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]]</small> (1819/1)
* [[Putorieae]] <small>[[Robert Sweet (botanist)|Sweet]]</small> (36/1)
* [[Rubieae]] <small>[[Henri Ernest Baillon|Baill.]]</small> (969/14)
* [[Schizocoleeae]] <small>Rydin & [[Birgitta Bremer|B.Bremer]]</small> (2/1)
* [[Schradereae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (62/1)
* [[Spermacoceae]] <small>[[Adelbert von Chamisso|Cham.]] & [[Diederich Franz Leonhard von Schlechtendal|Schltdl.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (1553/86)
* [[Temnopterygeae]] <small>Razafim. & Rydin</small> (1/1)
* [[Theligoneae]] <small>Wunderlich ex S.P.Darwin</small> (4/1)
* [[Urophylleae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex [[Bernard Verdcourt|Verdc.]]</small> (251/5)
{{div col end}}
=====Dialypetalanthoideae=====
Authority: [[James L. Reveal|Reveal]]; almost 6000 spp.
{{div col|colwidth=20em}}
* [[Airospermeae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (7/2)
* [[Alberteae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (9/3)
* [[Aleisanthieae]] <small>Mouly, J.Florence & [[Birgitta Bremer|B.Bremer]]</small> (10/3)
* [[Augusteae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (104/2)
* [[Bertiereae]] <small>[[Diane Mary Bridson|Bridson]]</small> (57/1)
* [[Chiococceae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (239/32)
* [[Chioneae]] <small>Razafim. & Rydin</small> (27/2)
* [[Cinchoneae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (125/8)
* [[Clavistigmateae]] <small>T.Y.Tu & P.W.Xie</small> (1/1)
* [[Coffeeae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (311/12)
* [[Cordiereae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]] emend. Mouly</small> (123/12)
* [[Crossopterygeae]] <small>F.White ex [[Diane Mary Bridson|Bridson]]</small> (1/1)
* [[Dialypetalantheae]] <small>Reveal</small> (310/35)
* [[Gardenieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (655/64)
* [[Glionnetieae]] <small>Razafim. & Rydin</small> (1/1)
* [[Greeneeae]] <small>Mouly, J.Florence & [[Birgitta Bremer|B.Bremer]]</small> (9/1)
* [[Guettardeae]] <small>[[Augustin Pyramus de Candolle|DC.]]</small> (769/20)
* [[Hamelieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (181/13)
* [[Henriquezieae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (36/3)
* [[Hillieae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]] ex S.P.Darwin</small> (29/3)
* [[Hymenodictyeae]] <small>Razafim. & [[Birgitta Bremer|B.Bremer]]</small> (32/2)
* [[Isertieae]] <small>[[Achille Richard|A.Rich.]] ex [[Augustin Pyramus de Candolle|DC.]]</small> (17/2)
* [[Ixoreae]] <small>[[George Bentham|Benth.]] & [[Joseph Dalton Hooker|Hook.f.]]</small> (564/1)
* [[Jackieae]] <small>[[Pieter Willem Korthals|Korth.]]</small> (1/1)
* [[Mussaendeae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (223/7)
* [[Naucleeae]] <small>[[Augustin Pyramus de Candolle|DC.]] ex [[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (194/19)
* [[Octotropideae]] <small>[[Richard Henry Beddome|Bedd.]]</small> (139/33)
* [[Pavetteae]] <small>[[Achille Richard|A.Rich.]] ex [[Barthélemy Charles Joseph Dumortier|Dumort.]]</small> (652/20)
* [[Posoquerieae]] <small>Delprete</small> (23/2)
* [[Retiniphylleae]] <small>[[Joseph Dalton Hooker|Hook.f.]]</small> (21/1)
* [[Rondeletieae]] <small>[[Augustin Pyramus de Candolle|DC.]] ex [[Friedrich Anton Wilhelm Miquel|Miq.]]</small> (192/18)
* [[Sabiceeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (169/4)
* [[Scyphiphoreae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (1/1)
* [[Sherbournieae]] <small>Mouly & [[Birgitta Bremer|B.Bremer]]</small> (57/4)
* [[Sipaneeae]] <small>[[Cornelis Eliza Bertus Bremekamp|Bremek.]]</small> (43/10)
* [[Steenisieae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (5/1)
* [[Strumpfieae]] <small>Delprete & T.J.Motley</small> (1/1)
* [[Trailliaedoxeae]] <small>Kainul. & [[Birgitta Bremer|B.Bremer]]</small> (1/1)
* [[Vanguerieae]] <small>[[Achille Richard|A.Rich.]] ex [[Barthélemy Charles Joseph Dumortier|Dumort.]]</small> (650/29)
{{div col end}}

====Genera====
{{For|a comprehensive list|List of Rubiaceae genera}}
The family Rubiaceae contains about 14,100 species in 580 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 1630 species, is the largest genus within the family and the third-largest genus of the angiosperms, after the legume ''[[Astragalus (plant)|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, 197 genera are [[monotypic]], which account for a third of all genera, but only for 1.4% of all species.<ref name ="Davis2009"/>

===Phylogeny===
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 two subfamilies of Rubiaceae together with the tribes Acranthereae, Coptosapelteae, and Luculieae are shown in the phylogenetic tree below. The placement of these three tribes relative to the two subfamilies has not been fully resolved.<ref name="BremerEriksson2009"/><ref name="RazaRydin2024"/>

{{clade
|label1='''Rubiaceae'''
|1={{clade
  |1={{clade
    |1=[[Luculieae]]
    }}
  |2={{clade
      |1=[[Acranthereae]]
      |2=[[Coptosapelteae]]
      }}
  |3={{clade
   |1=[[Rubioideae]]
   |2=[[Dialypetalanthoideae]]
   }}
 }}
}}

===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.<ref name="Graham2009"/>

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"/>

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"/>

==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. ''[[Vangueria infausta|V. infausta]]'', ''[[Vangueria madagascariensis|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: ''[[Coffea arabica|C. arabica]]'', ''[[Coffea canephora|C. canephora]]'', and ''[[Coffea liberica|C. liberica]]''.<ref name ="Davis2009"/>

===Medicinal===
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}}</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]].

===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 [[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]]''.

===Dyes===
[[Rose madder]], the crushed root of ''[[Rubia tinctorum]]'', yields a red dye, and the tropical ''[[Morinda citrifolia]]'' yields a yellow dye.

==Culture==
* ''[[Cinchona 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]].

==Image gallery==
<gallery>
File:Alberta magna.jpg|''[[Alberta magna]]''
File:IMG 7387-Arachnothryx leucophylla.jpg|''[[Arachnothryx leucophylla]]''
File:Asperula tinctoria Marzanka barwierska 01.jpg|''[[Asperula tinctoria]]''
File:Bikkia philippinensis.jpg|''[[Bikkia philippinensis]]''
File:Chiococca alba.jpg|''[[Chiococca alba]]''
File:Coffee Flowers.JPG|''[[Coffea arabica]]''
File:Galium uliginosum W.jpg|''[[Galium uliginosum]]''
File:Gardenia thunbergia.JPG|''[[Gardenia thunbergia]]''
File:Ixora coccinea- jungle geranium.JPG|''[[Ixora coccinea]]''
File:Soka-Ixora javanica1.jpg|''[[Ixora javanica]]''
File:Mitragyna speciosa111.JPG|''[[Mitragyna speciosa]]''
File:Morinda pubescens in Ananthagiri forest, AP W IMG 9225.jpg|''[[Morinda pubescens]]''
File:Nertera depressa 1.jpg|''[[Nertera granadensis]]''
File:Psychotria poeppigiana (bracts).jpg|''[[Psychotria poeppigiana]]''
File:Sherardia arvensis Kaldari 01.jpg|''[[Sherardia arvensis]]''
</gallery>

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}}

==External links==
{{Commons category|Rubiaceae}}
{{Wikispecies|Rubiaceae}}
* {{cite EB1911 |wstitle=Rubiaceae |volume=23 |page=808 |short=x}}
* [http://www.theplantlist.org/browse/A/Rubiaceae Rubiaceae] at [http://www.theplantlist.org ''The Plant List'']
* [http://eol.org/pages/4202/overview Rubiaceae] at [http://eol.org ''Encyclopedia of Life'']
* [http://www.mobot.org/mobot/research/APweb/orders/gentianalesweb.htm#Rubiaceae Rubiaceae] at [http://www.mobot.org/mobot/research/apweb ''Angiosperm Phylogeny Website'']
* [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'']
* [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'']
* [http://www.zimbabweflora.co.zw/speciesdata/family.php?family_id=198 Rubiaceae] at [http://www.zimbabweflora.co.zw ''Flora of Zimbabwe'']
* [https://florabase.dbca.wa.gov.au/browse/profile.php/22916 Rubiaceae] at [https://florabase.dbca.wa.gov.au ''Flora of Western Australia'']
* [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'']
* [https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=34784 Rubiaceae] at [https://www.itis.gov ''Integrated Taxonomic Information System'']
* [https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:30032120-2 Rubiaceae] at [https://powo.science.kew.org/ Plants of the World Online]

{{Angiosperm families}}
{{Taxonbar|from=Q156569}}
{{Authority control}}

[[Category:Rubiaceae| ]]
[[Category:Asterid families]]
[[Category:Extant Eocene first appearances]]
[[Category:Taxa named by Antoine Laurent de Jussieu]]