Editing Rubiaceae (section)

==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"/>