Rubiaceae are a family of flowering plants, commonly known as the
coffee, madder, or bedstraw family. It consists of terrestrial trees,
shrubs, lianas, or herbs that are recognizable by simple, opposite
leaves with interpetiolar stipules. The family contains about 13,500
species in 611 genera, which makes it the fourth-largest angiosperm
Rubiaceae has a cosmopolitan distribution; however, the
largest species diversity is concentrated in the (sub)tropics.
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).
2 Distribution and habitat
3.1 Flower biology
3.2 Fruit biology
3.3 Associations with other organisms
Subfamilies and tribes
7 Image gallery
9 External links
Rubiaceae are morphologically easily recognizable as a coherent
group by a combination of characters: opposite leaves that are simple
and entire (with the exception of several newly discovered
compound-leaved species of Pentagonia), interpetiolar stipules,
tubular sympetalous actinomorphic corollas and an inferior ovary.
A wide variety of growth forms are present: shrubs are most common
(e.g. Coffea, Psychotria), but members of the family can also be trees
(e.g. Cinchona, Nauclea), lianas (e.g.
Psychotria samoritourei), or
herbs (e.g. Galium, Spermacoce). Some epiphytes are also present (e.g.
Myrmecodia). The plants usually contain iridoids, various alkaloids,
and raphide crystals are common. The leaves are simple, undivided, and
entire; 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 stipules that are mostly fused to an
interpetiolar structure on either side of the stem between the
opposite leaves. Their inside surface often bears glands called
"colleters", which produce mucilaginous compounds protecting the young
shoot. The "whorled" leaves of the herbaceous
Rubieae tribe have
classically been interpreted as true leaves plus interpetiolar
leaf-like stipules. The inflorescence is a cyme, rarely of solitary
flowers (e.g. Rothmannia), and is either terminal or axillary and
paired at the nodes. The flowers are usually bisexual and usually
epigynous. The perianth is usually biseriate, although the calyx is
absent in some taxa (e.g. Theligonum). The calyx has four or five
sepals with basally fused lobes. The corolla is sympetalous with four
or five lobes, mostly actinomorphic, usually tubular, mostly white or
creamy but also yellow (e.g.
Gardenia spp., Mycelia basiflora), and
rarely blue (e.g.
Faramea calyptrata) or red (e.g. Alberta magna,
Ixora coccinea). They have four or five stamens, which are
alternipetalous and epipetalous. Anthers are longitudinal in
dehiscence, but some genera are poricidal (e.g. Rustia). The gynoecium
is syncarpous with an inferior ovary (rarely secondarily superior,
e.g. Gaertnera, Pagamea). Placentation is axial, rarely parietal
(e.g. Gardenia); ovules are anatropous to hemitropous, unitegmic, with
a funicular obturator, one to many per carpel. Nectaries are often
present as a nectariferous disk atop the ovary. The fruit is a berry,
capsule (e.g. Oldenlandia), drupe (e.g. Psychotria), or schizocarp
(e.g. Cremocarpon). Red fruits are fairly dominant (e.g. Coffea
arabica); yellow (e.g. Rosenbergiodendron formosum), orange (e.g.
Vangueria infausta), or blackish fruits (e.g.
are equally common; blue fruits are rather exceptional save in the
Psychotrieae and associated tribes. Most fruits are about 1.0 cm
in diameter; very small fruits are relatively rare and occur in
herbaceous tribes; very large fruits are rare and confined to the
Gardenieae. The seeds are endospermous.
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
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.
Rubiaceae consist of terrestrial and predominantly woody plants.
Woody rubiaceous shrubs constitute an important part of the
understorey of low- and mid-altitude rainforests.
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
Rubiaceae are zoophilous. Entomophilous species produce nectar
from an epigynous disk at the base of the corolla tube to attract
Ornithophily is rare and is found in red-flowered species of
Alberta, Bouvardia, and Burchellia. Anemophylous species are found in
Theligoneae and are characterized by
hermaphroditic or unisexual flowers that exhibit a set of specialized
features, such as striking sexual dimorphism, increased receptive
surface of the stigmas and pendulous anthers.
Rubiaceae species are hermaphroditic, outbreeding is
promoted through proterandry and spatial isolation of the reproductive
organs. More complex reproductive strategies include secondary pollen
presentation, heterodistyly, and unisexual flowers.
Secondary pollen presentation (also known as stylar pollen
presentation or ixoroid pollen mechanism) is especially known from the
Gardenieae and related tribes. The flowers are proterandrous and the
pollen is shed early onto the outside of the stigmas or the upper part
of the style, which serve as a 'receptaculum pollinis'. Increased
surface area and irregularity of the pollen receptacle, caused by
swellings, hairs, grooves or ridges often ensure a more efficient
pollen deposition. After elongation of the style, animals transport
the pollen to flowers in the female or receptive stage with exposed
stigmatic surfaces. A pollen catapult mechanism is present in the
genera Molopanthera and
Posoqueria (tribe Posoquerieae) that projects
a spherical pollen mass onto visiting sphingidae.
Heterodistyly is another mechanism to avoid inbreeding and is widely
present in the
Rubiaceae family. The tribes containing the largest
number of heterostylous species are
Spermacoceae and Psychotrieae.
Heterostyly is absent in groups that have secondary pollen
presentation (e.g. Vanguerieae).
Unisexual flowers also occur in
Rubiaceae and most taxa that have this
characteristic are dioecious. The two flower morphs are however
difficult to observe as they are rather morphologically similar; male
flowers have a pistillode with the ovaries empty and female flowers
have empty, smaller anthers (staminodes). Flowers that are
morphologically hermaphrodite, but functionally dioecious are for
example found in Pyrostria.
The dispersal units in
Rubiaceae can be entire fruits, syncarps,
mericarps, pyrenes or seeds. Fleshy fruit taxa are probably all
(endo)zoochorous (e.g. tribes Pavetteae, Psychotrieae), while the
dispersal of dry fruits is often unspecialized (e.g. tribes Knoxieae,
Spermacoceae). When seeds function as diaspores, the dispersal is
either anemochorous or hydrochorous. The three types of wind-dispersed
Rubiaceae are dust seeds (rare, e.g. Lerchea), plumed
seeds (e.g. Hillia), and winged seeds (e.g. Coutarea). Long-distance
dispersal by ocean currents is very rare (e.g. the seashore tree
Guettarda speciosa). Other dispersal mechanisms are absent or at least
very rare. Some
Spermacoceae having seeds with elaiosomes are probably
Spermacoce hepperiana). Epizoochorous taxa are
limited to herbaceous
Galium aparine fruits are
densely covered with hooked bristly hairs).
Associations with other organisms
The genera Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum, and
Squamellaria are succulent epiphytes that have evolved a mutualistic
relationship with ants. Their hypocotyl grows out into an
ant-inhabited tuber. Some shrubs or trees have ant holes in their
stems (e.g. Globulostylis). Some
Rubiaceae species have domatia
that are inhabited by mites (viz. acarodomatia; e.g. Plectroniella
An intimate association between bacteria and plants is found in three
rubiaceous genera (viz. Pavetta, Psychotria, and Sericanthe). 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,
Vangueria (all belonging to the Vanguerieae
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.
Rubiaceae family is named after Rubia, a name used by Pliny the
Elder in his Naturalis Historia for madder (
Rubia tinctorum). 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.
Rubiaceae (nomen conservandum) was published in 1789 by
Antoine Laurent de Jussieu, but the name was already mentioned in
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.
More recently, the morphologically quite different families
Dialypetalanthaceae, Henriqueziaceae, and
Theligonaceae were reduced to synonymy of Rubiaceae.
Subfamilies and tribes
The classical classification system of
Rubiaceae distinguished only
two subfamilies: Cinchonoideae, characterized by more than one ovule
in each locule, and Coffeoideae, having one ovule in each
locule. This distinction, however, was criticized because of
the distant position of two obviously related tribes, viz. Gardenieae
with many ovules in
Ixoreae with one ovule in
Coffeoideae, and because in species of
Tarenna the number of ovules
varies from one to several in each locule. During the
twentieth century other characters were used to delineate subfamilies,
e.g. stylar pollen presentation, raphides, endosperm, heterostyly,
etc. On this basis, three or eight subfamilies were
recognised. The last subfamilial classification solely based on
morphological characters divided
Rubiaceae into four subfamilies:
Cinchonoideae, Ixoroideae, Antirheoideae, and Rubioideae. 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.
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. 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. Currently, in
most molecular research concerning the
Rubiaceae family, the
classification with three subfamilies (Cinchonoideae, Ixoroideae, and
Rubioideae) is followed. However, an alternative view is proposed
where only two subfamilies are recognized, an expanded Cinchonoideae
(that includes Ixoroideae, Coptosapeltaeae and Luculieae) and
Rubioideae. The adoption of the Melbourne Code for botanical
nomenclature had an unexpected impact on many names that have been
long in use and are well-established in literature. According to the
Melbourne Code, the subfamilial name
Ixoroideae should be replaced by
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. 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, however, several
genera and species are not yet placed in a tribe.
Rubiaceae (59 sp.)
Coptosapelteae Bremek. ex S.P.Darwin (55 sp.)
Luculieae Rydin & B.Bremer (4 sp.)
Cinchonoideae Raf. (1708 sp.)
Chiococceae Benth. & Hook.f. (224 sp.)
Cinchoneae DC. (125 sp.)
Guettardeae DC. (747 sp.)
Hamelieae A.Rich. ex DC. (171 sp.)
Hymenodictyeae Razafim. & B.Bremer (25 sp.)
Hillieae Bremek. ex S.P.Darwin (29 sp.)
Isertieae A.Rich. ex DC. (16 sp.)
Naucleeae DC. ex Miq. (192 sp.)
Rondeletieae DC. ex Miq. (178 sp.)
Strumpfieae Delprete & T.J.Motley (1 sp.)
Ixoroideae Raf. (3960 sp.)
Airospermeae Kainul. & B.Bremer (7 sp.)
Alberteae Hook.f. (7 sp.)
Aleisanthieae Mouly, J.Florence & B.Bremer (10 sp.)
Augusteae Kainul. & B.Bremer (86 sp.)
Bertiereae Bridson (57 sp.)
Coffeeae DC. (303 sp.)
Condamineeae Hook.f. (305 sp.)
Cordiereae A.Rich. ex DC. emend. Mouly (124 sp.)
Cremasporeae Bremek. ex S.P.Darwin (2 sp.)
Crossopterygeae F.White ex Bridson (1 sp.)
Gardenieae A.Rich. ex DC. (587 sp.)
Greeneeae Mouly, J.Florence & B.Bremer (9 sp.)
Henriquezieae Benth. & Hook.f. (20 sp.)
Ixoreae Benth. & Hook.f. (545 sp.)
Jackieae Korth. (1 sp.)
Mussaendeae Hook.f. (221 sp.)
Octotropideae Bedd. (96 sp.)
Pavetteae A.Rich. ex Dumort. (624 sp.)
Posoquerieae Delprete (23 sp.)
Retiniphylleae Hook.f. (20 sp.)
Sabiceeae Bremek. (164 sp.)
Scyphiphoreae Kainul. & B.Bremer (1 sp.)
Sherbournieae Mouly & B.Bremer (54 sp.)
Sipaneeae Bremek. (43 sp.)
Steenisieae Kainul. & B.Bremer (5 sp.)
Trailliaedoxeae Kainul. & B.Bremer (1 sp.)
Vanguerieae A.Rich. ex Dumort. (644 sp.)
Rubioideae Verdc. (7571 sp.)
Anthospermeae Cham. & Schltdl. ex DC. (208 sp.)
Argostemmateae Bremek. ex Verdc. (215 sp.)
Clarkelleae Deb (1 sp.)
Colletoecemateae Rydin & B.Bremer (3 sp.)
Coussareeae Hook.f. (402 sp.)
Craterispermeae Verdc. (16 sp.)
Cyanoneuroneae Razafim. & B.Bremer (5 sp.)
Danaideae B.Bremer & Manen (50 sp.)
Dunnieae Rydin & B.Bremer (1 sp.)
Gaertnereae Bremek. ex S.P.Darwin (95 sp.)
Knoxieae Hook.f. (131 sp.)
Lasiantheae B.Bremer & Manen (239 sp.)
Mitchelleae Razafim. & B.Bremer & Manen (14 sp.)
Morindeae Miq. (165 sp.)
Ophiorrhizeae Bremek. ex Verdc. (364 sp.)
Paederieae DC. (81 sp.)
Palicoureeae Robbr. & Manen (817 sp.)
Perameae Bremek. ex S.P.Darwin (14 sp.)
Prismatomerideae Y.Z.Ruan (23 sp.)
Psychotrieae Cham. & Schltdl. (2114 sp.)
Putorieae (34 sp.)
Rubieae Baill. (938 sp.)
Schizocoleeae Rydin & B.Bremer (2 sp.)
Schradereae Bremek. (55 sp.)
Spermacoceae Cham. & Schltdl. ex DC. (1344 sp.)
Theligoneae Wunderlich ex S.P.Darwin (4 sp.)
Urophylleae Bremek. ex Verdc. (236 sp.)
For a comprehensive list, see List of
Rubiaceae family contains about 13,500 species in 619 genera. This
makes it the fourth-largest family of flowering plants by number of
species and fifth-largest by number of genera. Although taxonomic
adjustments are still being made, the total number of accepted genera
remains stable. In total, around 1338 genus names have been published,
indicating that more than half of the published names are synonyms.
Psychotria, with around 1850 species, is the largest genus within the
family and the third-largest genus of the angiosperms, after the
Astragalus and the orchid Bulbophyllum. However, the
Psychotria remains problematic and its adjustment
might reduce the number of species. In total, 30 genera have more than
100 species. However, 138 genera are monotypic, which account for 22%
of all genera, but only for 1.1% of all species.
Molecular studies have demonstrated the phylogenetic placement of
Rubiaceae within the order
Gentianales and the monophyly of the family
is confirmed. The relationships of the three subfamilies of
Rubiaceae together with the tribes
shown in the phylogenetic tree below. The placement of these two
groups relative to the three subfamilies has not been fully
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
Palaeocene have been referred to the family by various authors, none
of these fossils has been confirmed as belonging to the Rubiaceae.
The oldest confirmed fossils, which are fruits that strongly resemble
those of the genus Emmenopterys, were found in the Washington and are
48–49 million years old. A fossil infructescence and fruit found in
44 million-year-old strata in
Oregon was assigned to Emmenopterys
dilcheri, an extinct species. The next-oldest fossils date to the Late
Eocene and include
Canthium from Australia,
Faramea from Panama,
Guettarda from New Caledonia, and Paleorubiaceophyllum, an extinct
genus from the southeastern United States.
Rubiaceae are known from three regions in the
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.
Staple foods are not found in the Rubiaceae; instead, some species are
consumed locally and fruits may be used as famine food. Examples are
African medlar fruits (e.g. V. infausta, V. madagascariensis), African
Nauclea latifolia), and noni (
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: C. arabica,
C. canephora, and C. liberica.
The bark of trees in the genus
Cinchona is the source of a variety of
alkaloids, the most familiar of which is quinine, one of the first
agents effective in treating malaria. Woodruff (
Galium odoratum) is a
small herbaceous perennial that contains coumarin, a natural precursor
of warfarin, and the South American plant
Carapichea ipecacuanha is
the source of the emetic ipecac.
Psychotria viridis is frequently used
as a source of dimethyltryptamine in the preparation of ayahuasca, a
psychoactive decoction. The bark of the species
have been used in traditional African medicine for many years. 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.
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.
Ixora contains plants cultivated in warmer-climate gardens;
the most commonly grown species,
Ixora coccinea, is frequently used
for pretty red-flowering hedges.
Mussaenda cultivars with enlarged,
colored calyx lobes are shrubs with the aspect of Hydrangea; they are
mainly cultivated in tropical Asia. The New Zealand native Coprosma
repens is a commonly used plant for hedges. The South African
Rothmannia globosa is seen as a specimen tree in horticulture. Nertera
granadensis is a well-known house plant cultivated for its conspicuous
orange berries. Other ornamental plants include Mitchella, Morinda,
Pentas, and Rubia.
Rose madder, the crushed root of
Rubia tinctorum, yields a red dye,
and the tropical
Morinda citrifolia yields a yellow dye.
Warszewiczia coccinea is the national flower of Trinidad and Tobago.
Coffea arabica is the national flower of Yemen.
Cinchona is the national tree of
Ecuador and Peru.
Coffee Day is held each year on September 29.
^ a b "Angiosperm Phylogeny Website". Retrieved 1 June 2014.
^ Hammel BE (2015). "Three new species of
Pentagonia (Rubiaceae) from
southern Central America, one foreseen, two surprising" (PDF).
Phyotneuron. 46: 1–13.
^ Igersheim A, Puff C, Leins P, Erbar C (1994). "Gynoecial development
Gaertnera Lam. and of presumably allied taxa of the Psychotrieae
(Rubiaceae): secondary 'superior' vs. inferior ovaries". Botanische
Jahrbücher fur Systematik. 116: 401–414.
^ a b c d e Robbrecht E (1988). "Tropical woody Rubiaceae". Opera
Botanica Belgica. 1: 1–271.
^ Takhtajan A (2009). Flowering Plants (2 ed.). Springer.
^ a b c Davis AP, Govaerts R, Bridson DM, Ruhsam M, Moat J, Brummitt
NA (2009). "A global assessment of distribution, diversity, endemism,
and taxonomic effort in the Rubiaceae". Annals of the Missouri
Botanical Garden. 96 (1): 68–78. doi:10.3417/2006205.
^ Delprete PG (2009). "Taxonomic history, morphology, and reproductive
biology of the tribe
Posoquerieae (Rubiaceae, Ixoroideae)". Annals of
the Missouri Botanical Garden. 96 (1): 79–89.
^ Anderson WR (1973). "A morphological hypothesis for the origin of
heterostyly in the Rubiaceae". Taxon. 22 (5/6): 537–542.
^ Bridson DM (1987). "Studies in African Rubiaceae-Vanguerieae: a new
Pyrostria and a new subgenus,
Bullockia". Kew Bulletin. 42: 611–639. doi:10.2307/4110068.
^ Kapitany A (2007). Australian succulent plants: an introduction.
Boronia, Victoria: Kapitany Concepts. pp. 144–155.
^ Verstraete B, Lachenaud O, Smets E, Dessein S, Sonké B (2013).
"Taxonomy and phylogeny of Cuviera (Rubiaceae-Vanguerieae) and
reinstatement of the genus
Globulostylis with the description of three
new species". Botanical Journal of the Linnean Society. 173 (3):
^ Tilney PM, van Wyk AE, van deer Merwe CF (2012). "Structural
evidence in Plectroniella armada (Rubiaeae) for possible material
exchange between domatia and mites". PLoS ONE. 7 (7): e39984.
doi:10.1371/journal.pone.0039984. PMC 3390328 .
^ Lemaire B, Vandamme P, Merckx V, Smets E, Dessein S (2011).
"Bacterial leaf symbiosis in angiosperms: host specificity without
co-speciation". PLoS ONE. 6 (9): e24430.
doi:10.1371/journal.pone.0024430. PMC 3168474 .
^ Verstraete B, Van Elst D, Steyn H, Van Wyk B, Lemaire B, Smets E,
Dessein S (2011). "Endophytic bacteria in toxic South African plants:
identification, phylogeny and possible involvement in gousiekte". PLoS
ONE. 6 (4): e19265. doi:10.1371/journal.pone.0019265.
PMC 3082559 . PMID 21541284.
^ Verstraete B, Janssens S, Smets E, Dessein S (2013). "Symbiotic
beta-proteobacteria beyond legumes:
Burkholderia in Rubiaceae". PLoS
ONE. 8 (1): e55260. doi:10.1371/journal.pone.0055260.
PMC 3555867 . PMID 23372845.
^ Verstraete B, Janssens S, Lemaire B, Smets E, Dessein S (2013).
"Phylogenetic lineages in
Vanguerieae (Rubiaceae) associated with
Burkholderia bacteria in sub-Saharan Africa". American Journal of
Botany. 100: 2380–2387. doi:10.3732/ajb.1300303.
^ Sieber S, Carlier AL, Neuburger M, Grabenweger G, Eberl L, Gademann
K (2015). "Isolation and total synthesis of kirkamide, an
aminocyclitol from an obligate leaf nodule symbiont". Angewandte
Chemie International Edition. 54: 7968–7970.
doi:10.1002/anie.201502696. CS1 maint: Multiple names: authors
^ Simpson MG (2006).
Plant Systematics (1 ed.). Elsevier Academic
Press. ISBN 978-0-12-644460-5.
^ Jussieu A L de (1789).
Genera Plantarum. Paris: Herissant &
Barrois. p. 206.
^ Durand JF (1782). Notions Élémentaires de Botanique. Dijon: LN
Frantin. p. 274.
^ Fay MF, Bremer B, Prance GT, van der Bank M, Bridson D, Chase MW
(2000). "Plastid rbcL sequence data show
Dialypetalanthus to be a
member of Rubiaceae". Kew Bulletin. 55 (4): 853–864.
^ Rogers GK (1981). "The wood of Gleasonia, Henriquezia, and
Platycarpum (Rubiaceae) and its bearing on their classification: some
new considerations". Brittonia. 33 (3): 461–465.
^ Wunderlich R (1971). "Die systematische Stellung von Theligonum".
Österreichische botanische Zeitschrift. 119: 329–394.
^ Rutishauser F, Ronse Decraene LP, Smets E, Mendoza-Heuer I (1998).
Theligonum cynocrambe: developmental morphology of a peculiar
Plant Systematics and Evolution. 210 (1): 1–24.
^ a b Robbrecht E, Manen JF (2006). "The major evolutionary lineages
of the coffee family (Rubiaceae, angiosperms). Combined analysis (nDNA
and cpDNA) to infer the position of
Coptosapelta and Luculia, and
supertree construction based on rbcL, rps16, trnL-trnF and atpB-rbcL
data. A new classification in two subfamilies,
Rubioideae". Systematic Geography of Plants. 76: 85–146.
^ Hooker JD (1873). "Ordo LXXXIV. Rubiaceae". In Bentham G, Hooker JD.
Genera planetarium ad exemplaria imprimis in herbaria kewensibus
servata defirmata. 2. London. pp. 7–151.
^ Schumann K (1891). "Rubiaceae". In Engler A, Prantl K. Die
natürlichen Pflanzenfamilien. 4. Leipzig: Engelmann.
^ Baillon H (1878). "Sur les limits du genre Ixora". Adansonia. 12:
^ Solereder H (1893). "Ein Beitrag zur anatomischen Charakteristik und
zur Systematik deer Rubiaceen". Bull. Herb. Boissier. 1:
^ Verdcourt B (1958). "Remarks on the classification of the
Rubiaceae". Bulletin du Jardin botanique de l'état, Bruxelles. 28:
^ Bremekamp CEB (1966). "Remarks on the position, the delimitation and
the subdivision of the Rubiaceae". Acta Botanica Neerlandica. 15:
^ Bremer B, Andreasen K, Olsson D (1995). "Subfamilial and tribal
relationships in the
Rubiaceae based on rbcL sequence data". Annals of
the Missouri Botanical Garden. 82: 383–397.
^ Rydin C, Kainulainen K, Razafimandimbison SG, Smedmark JE, Bremer B
(2009). "Deep divergences in the coffee family and the systematic
position of Acranthera".
Plant Systematics and Evolution. 278:
^ Bremer B (2009). "A review of molecular phylogenetic studies of
Rubiaceae". Annals of the Missouri Botanical Garden. 96 (1): 4–26.
^ Reveal JL (2012). "Newly required infrafamilial names mandated by
changes in the code of nomenclature for algae, fungi and plants".
Phytoneuron. 33: 1–32.
^ Kainulainen K, Razafimandimbison SG, Bremer B (2013). "Phylogenetic
relationships and new tribal delimitations in subfamily Ixoroideae
(Rubiaceae)". Botanical Journal of the Linnean Society. 173 (3):
^ Mouly A, Kainulainen K, Persson C, Davis AP, Wong KM,
Razafimandimbison SG, Bremer B (2014). "Phylogenetic structure and
clade circumscriptions in the
Gardenieae complex (Rubiaceae)". Taxon.
63 (4): 801–818. doi:10.12705/634.4.
^ "World Checklist of Rubiaceae". Retrieved 1 March 2016.
^ Bremer B, Bremer K, Heidari N, Erixon P, Olmstead RG, Anderberg AA,
Källersjö M, Barkhordarian E (2002). "Phylogenetics of asteroids
based on 3 coding and 3 non-coding chloroplast DNA markers and the
utility of non-coding DNA at higher taxonomic levels". Molecular
Phylogenetics and Evolution. 24: 274–301.
doi:10.1016/s1055-7903(02)00240-3. PMID 12144762.
^ a b Bremer B, Eriksson T (2009). "Time tree of Rubiaceae: phylogeny
and dating the family, subfamilies, and tribes". International Journal
Plant Sciences. 170: 766–793. doi:10.1086/599077.
^ a b c Graham A (2009). "Fossil record of the Rubiaceae". Annals of
the Missouri Botanical Garden. 96 (1): 90–108.
^ Riba J, Valle M, Urbano G, Yritia M, Morte A, Barbanoj MJ (2003).
"Human pharmacology of ayahuasca: subjective and cardiovascular
effects, monoamine metabolite excretion, and pharmacokinetics".
Journal of Pharmacology and Experimental Therapeutics. 306 (1):
73–83. doi:10.1124/jpet.103.049882. PMID 12660312.
^ Neuwinger, Hans Dieter (1994). African Ethnobotany: Poisons and
Drugs: Chemistry, Pharmacology, Toxicology. Stuttgart, Germany:
Chapman & Hall. access-date= requires url= (help)
Wikimedia Commons has media related to Rubiaceae.
Wikispecies has information related to Rubiaceae
Rubiaceae at The
Rubiaceae at Encyclopedia of Life
Rubiaceae at Angiosperm Phylogeny Website
Rubiaceae at Flora of China
Rubiaceae at Flora of Pakistan
Rubiaceae at Flora of Zimbabwe
Rubiaceae at Flora of Western Australia
Rubiaceae at Flora of New Zealand
Rubiaceae at Integrated Taxonomic Information System
Rubiaceae at USDA NRCS Plants Database
Rubiaceae at Botanic Garden Meise
World Checklist of
Rubiaceae at Royal Botanic Gardens, Kew
Watson & Dallwitz: rubi