Vanguerieae is a

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

s in the

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Rubiaceae The 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 a ...

and contains about 655

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

in 30

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

. It is one of the most species-rich groups within the family and it is distributed across the

Paleotropics The Paleotropical Kingdom (Paleotropis) is a floristic kingdom comprising tropical areas of Africa, Asia and Oceania (excluding Australia and New Zealand), as proposed by Ronald Good and Armen Takhtajan. Part of its flora, inherited from the anc ...

Description

Several different life forms are present in the tribe: most species are

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from tree ...

s, but geofructices (plants with woody

rhizome In botany and dendrology, a rhizome (; , ) is a modified subterranean plant stem that sends out roots and shoots from its nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from axillary buds and grow ...

s) (e.g. '' Fadogia homblei'', '' Pygmaeothamnus zeyheri''), small

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

s (e.g. '' Vangueria infausta''), and climbers (e.g. '' Keetia gueinzii'') also occur. As all Rubiaceae species, the leaves are opposite, simple and entire, and they have interpetiolar stipules. The

phyllotaxis In botany, phyllotaxis () or phyllotaxy is the arrangement of leaves on a plant stem. Phyllotactic spirals form a distinctive class of patterns in nature. Leaf arrangement The basic arrangements of leaves on a stem are opposite and alterna ...

is decussate, sometimes conspicuously so (e.g. '' Canthium inerme''), and rarely whorled (e.g. '' Fadogia''). Some species have spines (e.g. '' Canthium''). Secondary pollen presentation is characteristic for the tribe and the species develop a conspicuous “stylar head”-complex, which is a structural unit consisting of a pollen presenting organ combined with stigmatic surfaces.

Distribution and habitat

The species of this tribe are found in the

. Most genera and over 70% of the species occur in

Sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...

, including

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Afric ...

and the islands in the Western

Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by ...

. The remainder of the species is mostly found in

Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...

. A few species are restricted to islands in the Pacific Ocean, especially

New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...

, and another fraction of the species occurs in the

Eastern states of Australia The eastern states of Australia are the states adjoining the east continental coastline of Australia. These are the mainland states of Victoria, New South Wales and Queensland, and the island state of Tasmania. The Australian Capital Territor ...

, especially in coastal

Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , establishe ...

. The countries with the largest diversity with over a hundred species each are

Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...

and D.R.Congo. The tribe is characterized by a high degree of

endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...

. Several monotypic genera have a restricted distribution range: '' Eriosemopsis'' is found in the

KwaZulu-Natal KwaZulu-Natal (, also referred to as KZN and known as "the garden province") is a province of South Africa that was created in 1994 when the Zulu bantustan of KwaZulu ("Place of the Zulu" in Zulu) and Natal Province were merged. It is loca ...

province of

South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...

, '' Everistia'' occurs in the

, '' Perakanthus'' occurs in

Peninsular Malaysia Peninsular Malaysia ( ms, Semenanjung Malaysia; Jawi: سمننجڠ مليسيا), or the States of Malaya ( ms, Negeri-negeri Tanah Melayu; Jawi: نڬري-نڬري تانه ملايو), also known as West Malaysia or the Malaysian Peninsula, ...

, and '' Temnocalyx'' is only found in southwestern

. Other larger genera with a restricted range are '' Cyclophyllum'' that is mainly found in

, '' Peponidium'' that is only found on

or the

Comoros The Comoros,, ' officially the Union of the Comoros,; ar, الاتحاد القمري ' is an independent country made up of three islands in southeastern Africa, located at the northern end of the Mozambique Channel in the Indian Ocean. It ...

, and '' Pyrostria'' of which most species are restricted to

. The tribe is a common and important constituent of many different kinds of

habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...

s. Species of Vanguerieae are found in both extreme wet habitats, such as the rainforests of tropical Africa, and in the very dry desert-like areas of the

horn of Africa The Horn of Africa (HoA), also known as the Somali Peninsula, is a large peninsula and geopolitical region in East Africa.Robert Stock, ''Africa South of the Sahara, Second Edition: A Geographical Interpretation'', (The Guilford Press; 2004 ...

and of the southernmost part of

. The tribe is found at elevations of over in the mountains of

Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...

and

, but also at sea-level along the coast of

Ecology

There is a wide range of flower sizes and fruit morphologies, suggesting different adaptations to

pollination Pollination is the transfer of pollen from an Stamen, anther of a plant to the stigma (botany), stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by Anemophily, wind. Pollinating agents can ...

and dispersal. Most of the species are probably pollinated by insects, but it is suspected that at least some of the large-flowered species of '' Fadogia'' are bird-pollinated. The edibility of many fruits suggests that they are dispersed by animals. Almost 30% of the species within the tribe have a symbiosis with endophytic bacteria, which are found intercellularly within the leaves. The presence of these bacteria is consistent on a genus level and all species belong to the following 5 genera: '' Fadogia'', '' Fadogiella'', '' Globulostylis'', '' Rytigynia'', and ''

Vangueria ''Vangueria'' is a genus of flowering plants in the family Rubiaceae. The genus is named for , as '' V. madagascariensis'' is known in Malagasy. Distribution The genus contains over 50 species distributed in Africa south of the Sahara with one ...

''. The bacteria are identified as ''

Burkholderia ''Burkholderia'' is a genus of Pseudomonadota whose pathogenic members include the ''Burkholderia cepacia'' complex, which attacks humans and ''Burkholderia mallei'', responsible for glanders, a disease that occurs mostly in horses and related ...

'', which is a genus that is also found in the leaves of other Rubiaceae species. The hypothesis is that these endophytic bacteria provide chemical protection against insect herbivory.

Uses

Some species are consumed locally and fruits may be used as

famine food A famine food or poverty food is any inexpensive or readily available food used to nourish people in times of hunger and starvation, whether caused by extreme poverty, such as during economic depression or war, or by natural disasters such a ...

. Examples are African medlar ('' Vangueria infausta'') and Spanish tamarind ('' Vangueria madagascariensis''), but the fruits of other ''

'' and '' Fadogia'' species are edible too. In

West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Mali ...

, '' Vangueria agrestis'' (known under its old name ''Fadogia agrestis'') is traditionally used as an

aphrodisiac An aphrodisiac is a substance that increases sexual desire, sexual attraction, sexual pleasure, or sexual behavior. Substances range from a variety of plants, spices, foods, and synthetic chemicals. Natural aphrodisiacs like cannabis or cocai ...

. It is known as ''bakin gagai'' ( Hausa) or black aphrodisiac.

Systematics

Taxonomy

The first genus of the tribe to be described was '' Canthium'', containing the species ''C. coronatum'' (now ''

Catunaregam spinosa ''Catunaregam spinosa'', the mountain pomegranate, is a flowering plant in the family Rubiaceae, found in South Asia and other Asian countries. Almost all parts of the plant are used as a traditional medicine in Ayurveda and fruits have been repo ...

'' in the

Gardenieae Gardenieae is a tribe of flowering plants in the family Rubiaceae and contains about 586 species in 53 genera. Genera Currently accepted names * '' Adenorandia'' Vermoesen (1 sp) * '' Agouticarpa'' C.H.Press. (7 sp) * '' Aidia'' Lour. (55 sp ...

tribe) and ''C. parviflorum'' (now '' C. coromandelicum''). The second genus to be described was '' Psydrax'', containing the species '' Psydrax dicoccos''. Although the tribe is predominantly found in Africa, these first described species are Asiatic taxa. The first non-Asiatic taxa to be described were the genera '' Pyrostria'' and ''

'', based on material collected on Western Indian Ocean islands. The first genus from continental Africa was '' Cuviera'', with the species '' Cuviera acutiflora'' from

Sierra Leone Sierra Leone,)]. officially the Republic of Sierra Leone, is a country on the southwest coast of West Africa. It is bordered by Liberia to the southeast and Guinea surrounds the northern half of the nation. Covering a total area of , Sierr ...

. Dumortier first mentioned a tribe Vanguerieae (as “Vaugnerieae”) attributed to

Achille Richard Achille Richard was a French botanist, botanical illustrator and physician (27 April 1794 in Paris – 5 October 1852). Biography Achille was the son of the botanist Louis-Claude Marie Richard (1754–1821). He was a pharmacist in the Frenc ...

and put the genera ''

Hamelia ''Hamelia'' is a genus of flowering plants in the coffee family, Rubiaceae. The name honors French botanist Henri-Louis Duhamel du Monceau (1700–1782). Selected species * '' Hamelia axillaris'' Sw. – Guayabo Negro * '' Hamelia cupr ...

'', '' Evosmia'', '' Mitchella'', ''“Vaugneria”'' (''

''), and '' Nonatelia'' in it. However, this circumscription of the tribe was far from correct, as '' Canthium'' and '' Cuviera'' were put in different tribes and '' Psydrax'' and '' Pyrostria'' were not even mentioned. Also, the only genus that actually belongs in the tribe is ''

''; the other genera belong elsewhere. At that time, authors often used the number of

locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...

s as a character for delimiting groups, but this is problematic for Vanguerieae as both 2-locular (e.g. '' Canthium'') and 5-locular (e.g. ''

'') ovaries occur in the tribe. The first monograph of the tribe was performed by Walter Robyns in 1928 and a total of 17 genera were allocated to the tribe. Although 9 new genera were described, this work was not entirely complete, e.g. '' Canthium'', '' Cuviera'', and '' Pyrostria'' were not treated. Since then, several new genera have been described or made synonym and more taxonomic adjustments on genus level will probably still be necessary.

Genera

Currently accepted names * '' Afrocanthium'' ( Bridson) Lantz & B.Bremer (17 sp) * '' Bridsonia'' Verstraete & A.E.van Wyk (1 sp) * '' Bullockia'' ( Bridson) Razafim., Lantz & B.Bremer (6 sp) * '' Canthium'' Lam. (81 sp) * '' Canthiumera'' K.M.Wong & Mahyuni (4 sp) * '' Cuviera''

DC. Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...

(10+6 sp) * '' Cyclophyllum''

Hook.f. Sir Joseph Dalton Hooker (30 June 1817 – 10 December 1911) was a British botanist and explorer in the 19th century. He was a founder of geographical botany and Charles Darwin's closest friend. For twenty years he served as director of t ...

(44 sp) * '' Dibridsonia'' K.M.Wong (3 sp) * '' Eriosemopsis'' Robyns (1 sp) * '' Everistia'' S.T.Reynolds & R.J.F.Hend. (1 sp) * '' Fadogia'' Schweinf. (40 sp) * '' Fadogiella'' Robyns (3 sp) * '' Globulostylis'' Wernham (8 sp) * ''

Hutchinsonia ''Hutchinsonia'' is a genus of flowering plants in the family Rubiaceae The 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 ...

'' Robyns (2 sp) * '' Kanapia'' Arriola & Alejandro (2 sp) * '' Keetia'' E.Philipps (36 sp) * '' Meyna'' Roxb. ex Link (9 sp) * '' Multidentia'' Gilli (10 sp) * '' Peponidium'' (

Baill. Henri Ernest Baillon was a French botanist and physician. He was born in Calais on 30 November 1827 and died in Paris on 19 July 1895. Baillon spent his professional life as a professor of natural history, and he published numerous works on b ...

) Arènes (48 sp) * '' Perakanthus'' Robyns ex

Ridl. Henry Nicholas Ridley CMG (1911), MA (Oxon), FRS, FLS, F.R.H.S. (10 December 1855 – 24 October 1956) was an English botanist, geologist and naturalist who lived much of his life in Singapore. He was instrumental in promoting rubber trees i ...

(1 sp) * '' Plectroniella'' Robyns (1 sp) * '' Psydrax'' Gaertn. (90 sp) * '' Pygmaeothamnus'' Robyns (1 sp) * '' Pyrostria'' Comm. ex A.Juss. (67 sp) * '' Robynsia'' Hutch. (1 sp) * '' Rytigynia'' Blume (82 sp) * '' Temnocalyx'' Robyns (1 sp) * ''

'' Juss. (58 sp) * '' Vangueriella'' Verdc. (18 sp) * '' Vangueriopsis'' Robyns (4 sp) Synonyms * '' Ancylanthos'' Desf. = ''

'' * ''Clusiophyllea''

= '' Canthium'' * ''Dinocanthium'' Bremek. = '' Pyrostria'' * '' Dondisia''

= '' Canthium'' * '' Lagynias'' E.Mey. ex Robyns = ''

'' * '' Leroya'' Cavaco = '' Pyrostria'' * ''Lycioserissa'' Roem. &

Schult. Josef (Joseph) August Schultes (15 April 1773 in Vienna – 21 April 1831 in Landshut) was an Austrian botanist and professor from Vienna. Together with Johann Jacob Roemer (1763–1819), he published the 16th edition of Linnaeus' ''Systema ...

= '' Canthium'' * ''Mesoptera''

= '' Psydrax'' * '' Mitrastigma'' Harv. = '' Psydrax'' * '' Neoleroya'' Cavaco = '' Pyrostria'' * '' Pachystigma'' Hochst. = ''

'' * ''Phallaria'' Schumach. & Thonn. = '' Psydrax'' * ''Plectronia'' Lour. = '' Canthium'', '' Peponidium'', '' Psydrax'' * '' Pseudopeponidium'' Homolle ex Arènes = '' Pyrostria'' * ''Psilostoma''

Klotzsch Johann Friedrich Klotzsch (9 June 1805 – 5 November 1860) was a German pharmacist and botanist. His principal work was in the field of mycology, with the study and description of many species of mushroom. Klotzsch was born in Wittenberg. Orig ...

ex Eckl. & Zeyh. = '' Canthium'' * '' Rhopalobrachium''

Schltr. Friedrich Richard Rudolf Schlechter (16 October 1872 – 16 November 1925) was a German taxonomist, botanist, and author of several works on orchids. He went on botanical expeditions in Africa, Indonesia, New Guinea, South and Central America ...

& K.Krause = '' Cyclophyllum'' * ''Scyphochlamys'' Balf.f. = '' Pyrostria'' * '' Tapiphyllum'' Robyns = ''

'' * '' Vavanga'' Rohr = ''

Phylogenetic relationships

The following

phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

is based on

molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

studies Study or studies may refer to: General * Education ** Higher education * Clinical trial * Experiment * Observational study * Research * Study skills, abilities and approaches applied to learning Other * Study (art), a drawing or series of d ...

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. T ...

Image gallery

File:Canthium angustifolium Roxb. 7.JPG, '' Canthium angustifolium'' File:Canthium coromandelicum in Keesaraguda, AP W IMG 9056.jpg, '' Canthium coromandelicum'' File:Cyclophyllum coprosmoides flowers and foliage.jpg, '' Cyclophyllum coprosmoides'' File:Everistia vacciniifolia var. vacciniifolia flowers.jpg, '' Everistia vacciniifolia'' File:Fadogia homblei01.jpg, '' Fadogia homblei'' File:False Turkeyberry (Canthium armatum) (16462475912).jpg, '' Plectroniella armata'' File:Psydrax banksii (8467185479).jpg, '' Psydrax banksii'' File:Psydrax foliage.jpg, '' Psydrax latifolia'' File:Psydrax livida, bloeiwyses, b, Seringveld.jpg, '' Psydrax livida'' File:Psydrax odorata (5816464018).jpg, '' Psydrax odorata'' File:Vangueria infausta, habitus, a, Pretoria NBT.jpg, '' Vangueria infausta'' File:Vangueria infausta, blomme, Schanskop, b.jpg, '' Vangueria infausta'' File:Vangueria infausta, groen vrugte, Faerie Glen NR.jpg, '' Vangueria infausta'' File:Vangueria parvifolia, vrug, Walter Sisulu NBT.jpg, ''

Vangueria parvifolia ''Vangueria'' is a genus of flowering plants in the family Rubiaceae. The genus is named for , as '' V. madagascariensis'' is known in Malagasy. Distribution The genus contains over 50 species distributed in Africa south of the Sahara with one ...

'' File:Vangueria macrocalyx, b, Skeerpoort.jpg, '' Vangueria macrocalyx''

References

{{Taxonbar, from=Q3595854 Ixoroideae tribes