Rubiaceae

The Rubiaceae are a family of owering plants, commonly known as the coee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs
that are recognizable by simple, opposite leaves with
interpetiolar stipules. The family contains about 13,500
species in 611 genera, which makes it the fourth-largest
angiosperm family. Rubiaceae has a cosmopolitan distribution, however, the largest species diversity is concentrated in the (sub)tropics.[1] Economic importance includes Coea, the source of coee, Cinchona, the source
of the antimalarial alkaloid quinine, some dye plants (e.g.
Rubia), and ornamental cultivars (e.g. Gardenia, Ixora,
Pentas).

taxa (e.g. Theligonum). The calyx has four or ve sepals

with basally fused lobes. The corolla is sympetalous
with four or ve lobes, mostly actinomorphic, usually
tubular, mostly white or creamy but also yellow (e.g.
Gardenia spp., Mycelia basiora), and rarely blue (e.g.
Faramea calyptrata) or red (e.g. Alberta magna, Ixora
coccinea). They have four or ve 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[2] ).
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. Coea 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 cm in diameter; very small fruits are relatively rare and occur in herbaceous tribes; very large fruits
are rare and conned to the Gardenieae. The seeds are
endospermous.[3][4]

Description

The Rubiaceae are morphologically easily recognizable

as a coherent group by a combination of characters: opposite leaves that are simple and entire, interpetiolar stipules, tubular sympetalous actinomorphic corollas and an
inferior ovary.
A wide variety of growth forms are present: shrubs
are most common (e.g. Coea, 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 inorescence is a cyme, rarely
of solitary owers (e.g. Rothmannia), and is either terminal or axillary and paired at the nodes. The owers
are usually bisexual and usually epigynous. The perianth
is usually biseriate, although the calyx is absent in some

2 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 AfroAmerican distributions are rare (e.g. Sabicea). Endemic
rubiaceous genera are found in most tropical and subtropical oristic 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.[5]
The Rubiaceae consist of terrestrial and predominantly
woody plants. Woody rubiaceous shrubs constitute an
important part of the understorey of low- and mid1

4 SYSTEMATICS

altitude rainforests. Rubiaceae are tolerant of a broad

array of environmental conditions (soil types, altitudes,
community structures, etc.) and do not specialize in one
specic habitat type (although genera within the family
often specialize).

3
3.1

Ecology
Flower biology

3.2 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 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), 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 myrmecochorous
(e.g. Spermacoce hepperiana). Epizoochorous taxa are
limited to herbaceous Rubiaceae (e.g. Galium aparine
fruits are densely covered with hooked bristly hairs).

Most Rubiaceae are zoophilous. 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-owered species of Alberta, Bouvardia,
and Burchellia. Anemophylous species are found in the
tribes Anthospermeae and Theligoneae and are characterized by hermaphroditic and/or unisexual owers that
exhibit a set of specialized features, such as striking sex- 3.3 Associations with other organisms
ual dimorphism, increased receptive surface of the stigmas and pendulous anthers.[3]
The genera Anthorrhiza, Hydnophytum, Myrmecodia,
Although most Rubiaceae species are hermaphroditic, Myrmephytum, and Squamellaria are succulent epiphytes
outbreeding is promoted through proterandry and spatial that have evolved a mutualistic relationship with ants.
isolation of the reproductive organs. More complex re- Their hypocotyl grows out into an ant-inhabited tuber.[9]
productive strategies include secondary pollen presenta- Some shrubs or trees have ant holes in their stems (e.g.
Globulostylis).[10]
tion, heterodistyly, and unisexual owers.
Secondary pollen presentation (also known as stylar Some Rubiaceae species have domatia that are inhabpollen presentation or ixoroid pollen mechanism) is es- ited by mites (viz. acarodomatia; e.g. Plectroniella arpecially known from the Gardenieae and related tribes. mata).[11]
The owers are proterandrous and the pollen is shed An intimate association between bacteria and plants
early onto the outside of the stigmas and/or the up- is found in three rubiaceous genera (viz. Pavetta,
per part of the style, which serve as a 'receptaculum Psychotria, and Sericanthe).[12] The presence of endopollinis. Increased surface area and irregularity of the phytic bacteria is visible by eye because of the formapollen receptacle, caused by swellings, hairs, grooves tion of dark spots or nodules in the leaf blades. The enor ridges often ensure a more ecient pollen deposi- dophytes have been identied as Burkholderia bacteria.
tion. After elongation of the style, animals transport the Their function, however, remains enigmatic. A dierent
pollen to owers in the female or receptive stage with ex- type of plantbacteria association is found in the genera
posed stigmatic surfaces. A pollen catapult mechanism is Fadogia, Fadogiella, Globulostylis, Rytigynia, Vangueria
present in the genera Molopanthera and Posoqueria (tribe (all belonging to the Vanguerieae tribe), where bacteria
Posoquerieae) that projects a spherical pollen mass onto are found freely distributed among the mesophyll cells
and no leaf nodules are formed.[13][14][15]
visiting sphingidae.[6]
Heterodistyly is another mechanism to avoid inbreeding and is widely present in the Rubiaceae family.[7]
The tribes containing the largest number of heterostylous
species are Spermacoceae and Psychotrieae. Heterostyly 4 Systematics
is absent in groups that have secondary pollen presentaThe Rubiaceae family is named after Rubia, a name used
tion (e.g. Vanguerieae).
Unisexual owers also occur in Rubiaceae and most taxa by Pliny the Elder in his Naturalis Historia for madder
that have this characteristic are dioecious. The two ower (Rubia tinctorum).[16] The roots of this plant have been
morphs are however dicult to observe as they are rather used since ancient times to extract alizarin and purpurin,
morphologically similar; male owers have a pistillode two red dyes used for coloring clothes. The name rubia
with the ovaries empty and female owers have empty, is therefore derived from the Latin word ruber, meaning
smaller anthers (staminodes).[3] Flowers that are morpho- red. The well-known genus Rubus (blackberries and rasplogically hermaphrodite, but functionally dioecious are berries) is unrelated and belongs to Rosaceae, the rose
family.
for example found in Pyrostria.[8]

4.2

4.1

Phylogeny

Taxonomy

The name Rubiaceae (nomen conservandum) was published in 1789 by Antoine Laurent de Jussieu,[17] but the
name was already mentioned in 1782.[18]
Several historically accepted families are since long
included in Rubiaceae: Aparinaceae, Asperulaceae,
Catesbaeaceae, Cephalanthaceae, Cinchonaceae, Coffeaceae, Coutariaceae, Galiaceae, Gardeniaceae, Guettardaceae, Hameliaceae, Hedyotidaceae, Houstoniaceae,
Hydrophylacaceae, Lippayaceae, Lygodisodeaceae,
Naucleaceae, Nonateliaceae, Operculariaceae, Pagamaeaceae, Psychotriaceae, Randiaceae, Sabiceaceae,
Spermacoceaceae.[1]
More recently, the morphologically quite dierent
families Dialypetalanthaceae,[19] Henriqueziaceae,[20]
and Theligonaceae[21][22][23] were reduced to synonymy
of Rubiaceae.

3
Rubioideae.[23] The adoption of the Melbourne Code for
botanical nomenclature had an unexpected impact on
many names that have been long in use and are wellestablished in literature. According to the Melbourne
Code, the subfamilial name Ixoroideae should be replaced
by Dialypetalanthoideae.[33] 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.[34][35] 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,[36] however, several genera
and species are not yet placed in a tribe.
4.1.2 Genera

4.1.1

Subfamilies and tribes

The classical classication 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.[24][25] 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 Coeoideae, and because in species of Tarenna
the number of ovules varies from one to several in each
locule.[26][27] During the twentieth century other characters were used to delineate subfamilies, e.g. stylar pollen
presentation, raphides, endosperm, heterostyly, etc. On
this basis, three[28] or eight[29] subfamilies were recognised. The last subfamilial classication solely based
on morphological characters divided Rubiaceae into four
subfamilies: Cinchonoideae, Ixoroideae, Antirheoideae,
and Rubioideae.[3] 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.[3]
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 dierently circumscribed, and
Antirheoideae is shown to be polyphyletic.[30] 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.[31] Currently, in most molecular research concerning the Rubiaceae family, the classication with three subfamilies (Cinchonoideae, Ixoroideae,
and Rubioideae) is followed.[32] However, an alternative view is proposed where only two subfamilies
are recognized, an expanded Cinchonoideae (that includes Ixoroideae, Coptosapeltaeae and Luculieae) and

For a comprehensive list, see List of Rubiaceae genera.

The Rubiaceae family contains about 13,500 species in
611 genera. This makes it the fourth-largest family of
owering plants by number of species and fth-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.[5]

4.2 Phylogeny
Molecular studies have demonstrated the phylogenetic
placement of Rubiaceae within the order Gentianales and
the monophyly of the family is conrmed.[37][38] 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.[38]

4.3 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

6 CULTURE

medicine for many years.[41] 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
The oldest conrmed fossils, which are fruits that strongly qualities, behaving as a -opioid receptor agonist, and ofresemble those of the genus Emmenopterys, were found in ten being used in traditional Thai medicine in a similar
the Washington and are 4849 million years old. A fos- way to and often as a replacement for opioid painkillers
like morphine.
sil 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 Pale- 5.4 Ornamentals
orubiaceophyllum, an extinct genus from the southeastern
United States.[39]
Originally from China, the common gardenia (Gardenia
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 conrmed as belonging to the Rubiaceae.[39]

jasminoides) is a widely grown garden plant and ower 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-owering 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
5 Uses
repens is a commonly used plant for hedges. The South
African Rothmannia globosa is seen as a specimen tree in
5.1 Food
horticulture. Nertera granadensis is a well-known house
plant cultivated for its conspicuous orange berries. Other
Staple foods are not found in the Rubiaceae; instead, ornamental plants include Mitchella, Morinda, Pentas,
some species are consumed locally and fruits may be and Rubia.
used as famine food. Examples are African medlar fruits
(e.g. V. infausta, V. madagascariensis), African peach
(Nauclea latifolia), and noni (Morinda citrifolia).
Fossil Rubiaceae are known from three regions in the
Eocene (North America north of Mexico, MexicoCentral America-Caribbean, and Southeast PacicAsia). 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.[39]

5.5 Dyes
5.2

Beverage

Rose madder, the crushed root of Rubia tinctorum, yields

The most economically important member of the family a red dye, and the tropical Morinda citrifolia yields a yeland the worlds second-most important commodity (af- low dye.
ter petroleum) is the genus Coea used in the production of coee. Coea includes 124 species, but only three
species are cultivated for coee production: C. arabica,
C. canephora, and C. liberica.[5]

6 Culture

5.3

Medicinal

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 rst agents eective in treating malaria. Woodru (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.[40] The bark of the species
Breonadia salicina have been used in traditional African

Warszewiczia coccinea is the national ower of

Trinidad and Tobago.

Coea arabica is the national ower of Yemen.

Cinchona is the national tree of Ecuador and Peru.
The International Coee Day is held each year on
September 29.

Image gallery

Galium uliginosum

Gardenia thunbergia

Ixora coccinea

Ixora javanica

Mitragyna speciosa

Morinda pubescens

Nertera granadensis

Alberta magna

Arachnothryx leucophylla

Asperula tinctoria

Bikkia philippinensis

Chiococca alba

Coea arabica

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Psychotria poeppigiana

Sherardia arvensis

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9 External links

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[34] Kainulainen K, Razamandimbison SG, Bremer B
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doi:10.1111/boj.12038.
[35] Mouly A, Kainulainen K, Persson C, Davis AP, Wong
KM, Razamandimbison SG, Bremer B (2014). Phylogenetic structure and clade circumscriptions in the Gardenieae complex (Rubiaceae)". Taxon. 63 (4): 801818.
doi:10.12705/634.4.
[36] the World Checklist of Rubiaceae. Retrieved March
2016. Check date values in: |access-date= (help)
[37] Bremer B, Bremer K, Heidari N, Erixon P, Olmstead RG, Anderberg AA, Kllersj 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: 274301.
doi:10.1016/s1055-7903(02)00240-3.

Rubiaceae at The Plant List

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

10

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10.1

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

Text and image sources, contributors, and licenses

Text

Rubiaceae Source: https://en.wikipedia.org/wiki/Rubiaceae?oldid=751078115 Contributors: Eclecticology, Josh Grosse, PierreAbbat,

Ben-Zin~enwiki, Henriette~enwiki, Twilsonb, Looxix~enwiki, Stan Shebs, Glenn, KayEss, Marshman, Topbanana, Wiwaxia, Moriori,
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