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Vanguerieae
	Canthium coromandelicum
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Asterids
Order:	Gentianales
Family:	Rubiaceae
Subfamily:	Dialypetalanthoideae
Tribe:	Vanguerieae; A.Rich. ex Dumort.
Type genus
	Vangueria; Juss.

Vanguerieae is a tribe of flowering plants in the family Rubiaceae and contains 671 species in 29 genera.^[1] It is one of the most species-rich groups within the family and it is distributed across the Paleotropics.

Description

Several different life forms are present in the tribe: most species are shrubs, but geofructices (plants with woody rhizomes) (e.g. Fadogia homblei, Pygmaeothamnus zeyheri), small trees (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 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.^[2]

Distribution and habitat

The species of this tribe are found in the Paleotropics. Most genera and over 70% of the species occur in Sub-Saharan Africa, including Madagascar and the islands in the Western Indian Ocean. The remainder of the species is mostly found in Southeast Asia. A few species are restricted to islands in the Pacific Ocean, especially New Caledonia, and another fraction of the species occurs in the Eastern states of Australia, especially in coastal Queensland.^[1] The countries with the largest diversity with over a hundred species each are Tanzania, Madagascar and D.R.Congo.

The tribe is characterized by a high degree of endemism. Several monotypic genera have a restricted distribution range: Eriosemopsis is found in the KwaZulu-Natal province of South Africa, Everistia occurs in the Eastern states of Australia, Perakanthus occurs in Peninsular Malaysia, and Temnocalyx is only found in southwestern Tanzania. Other larger genera with a restricted range are Cyclophyllum that is mainly found in New Caledonia, Peponidium that is only found on Madagascar or the Comoros, and Pyrostria of which most species are restricted to Madagascar.

The tribe is a common and important constituent of many different kinds of habitats. 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 and of the southernmost part of Madagascar. The tribe is found at elevations of over 2,000 m (6,600 ft) in the mountains of Malawi and Tanzania, but also at sea-level along the coast of South Africa.

Ecology

There is a wide range of flower sizes and fruit morphologies, suggesting different adaptations to pollination 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 mostly found intercellularly within the leaves.^[3]^[4]^[5] The presence of such bacteria is consistent on a genus level and all species belong to the following 5 genera: Fadogia, Fadogiella, Globulostylis, Rytigynia, and Vangueria. The bacteria are identified as Burkholderia, which is a genus that is also found in the leaves of other Rubiaceae species.^[6] Bacterial leaf nodules were observed in Keetia nodulosa, although without microscopic and molecular confirmation for now.^[7] The hypothesis is that these endophytic bacteria provide chemical protection against insect herbivory.^[8]

Uses

Some species are consumed locally and fruits may be used as famine food. Examples are African medlar (Vangueria infausta) and Spanish tamarind (Vangueria madagascariensis), but the fruits of other Vangueria and Fadogia species are edible too.

In West Africa, Vangueria agrestis (known under its synonym Fadogia agrestis) is traditionally used as an aphrodisiac. It is known as bakin gagai (Hausa) or black aphrodisiac.^[9]

Systematics

Taxonomy

The first genus of the tribe to be described was Canthium, containing the species C. coronatum (now Catunaregam spinosa in the Gardenieae tribe) and C. parviflorum (now C. coromandelicum).^[10] The second genus to be described was Psydrax, containing the species Psydrax dicoccos.^[11] 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 Vangueria, based on material collected on Western Indian Ocean islands.^[12] The first genus from continental Africa was Cuviera, with the species Cuviera acutiflora from Sierra Leone.^[13] Dumortier first mentioned a tribe Vanguerieae (as “Vaugnerieae”) attributed to Achille Richard and put the genera Hamelia, Evosmia, Mitchella, “Vaugneria” (Vangueria), and Nonatelia in it.^[14] 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 Vangueria; the other genera belong elsewhere. At that time, authors often used the number of locules as a character for delimiting groups, but this is problematic for Vanguerieae as both 2-locular (e.g. Canthium) and 5-locular (e.g. Vangueria) 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.^[15] 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^[1]

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. (18 sp)
Cyclophyllum Hook.f. (42 sp)
Dibridsonia K.M.Wong (3 sp)
Eriosemopsis Robyns (1 sp)
Everistia S.T.Reynolds & R.J.F.Hend. (1 sp)
Fadogia Schweinf. (39 sp)
Fadogiella Robyns (3 sp)
Globulostylis Wernham (7 sp)
Hutchinsonia Robyns (2 sp)
Kanapia Arriola & Alejandro (2 sp)
Keetia E.Philipps (39 sp)
Meyna Roxb. ex Link (9 sp)
Multidentia Gilli (10 sp)
Peponidium (Baill.) Arènes (48 sp)
Perakanthus Robyns ex Ridl. (2 sp)
Psydrax Gaertn. (97 sp)
Pygmaeothamnus Robyns (1 sp)
Pyrostria Comm. ex A.Juss. (76 sp)
Robynsia Hutch. (1 sp)
Rytigynia Blume (81 sp)
Temnocalyx Robyns (1 sp)
Vangueria Juss. (58 sp)
Vangueriella Verdc. (17 sp)
Vangueriopsis Robyns (4 sp)

Synonyms

Ancylanthos Desf. = Vangueria
Caranda Gaertn. = Psydrax
Clusiophyllea Baill. = Peponidium
Dinocanthium Bremek. = Pyrostria
Dondisia DC. = Canthium
Lagynias E.Mey. ex Robyns = Vangueria
Leroyia Cavaco = Pyrostria
Lycioserissa Roem. & Schult. = Canthium
Mesoptera Hook.f. = Psydrax
Mitrastigma Harv. = Psydrax
Neoleroya Cavaco = Pyrostria
Pachystigma Hochst. = Vangueria
Phallaria Schumach. & Thonn. = Psydrax
Plectronia Lour. = Canthium, Peponidium, Psydrax
Plectroniella Robyns = Canthium
Pseudopeponidium Homolle ex Arènes = Pyrostria
Psilostoma Klotzsch ex Eckl. & Zeyh. = Canthium
Rhopalobrachium Schltr. & K.Krause = Cyclophyllum
Rytigyniopsis Capuron = Rytigynia
Scyphochlamys Balf.f. = Pyrostria
Tapiphyllum Robyns = Vangueria
Vavanga Rohr = Vangueria
Webera Cramer = Canthium
Wittmannia Vahl = Vangueria

Phylogenetic relationships

The following phylogenetic tree is based on molecular phylogenetic studies of DNA sequences.^[3]^[4]^[5]^[16]^[17]^[18]^[19]

Vanguerieae

	Kanapia

	Keetia

Psydrax

dioecious group

	Bullockia

	Pyrostria

Canthium group

Bridsonia chamaedendrum

Eriosemopsis

	Canthium

	Rytigynia bugoyensis

	Rytigynia mrimaensis

	Meyna

	Vangueriella sect. Stenosepalae

	Dibridsonia

	Canthiumera

	Fadogia

	Fadogiella

	Rytigynia

	Hutchinsonia

	Cuviera

	Globulostylis

	Vangueriella sect. Vangueriella

Vangueria group

Multidentia

	Pygmaeothamnus zeyheri

	Robynsia glabrata

	Vangueriopsis

Vangueria

Image gallery

References

^ ^a ^b ^c POWO (2024). "Plants of the World Online". Royal Botanic Gardens, Kew. Retrieved 2024-10-08.
^ Igersheim A (1993). "Gynoecium development in Rubiaceae-Vanguerieae, with particular reference to the "stylar head"-complex and secondary pollen presentation". Plant Systematics and Evolution. 187 (1–4): 175–190. doi:10.1007/bf00994098. S2CID 33654368.
^ ^a ^b 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.
^ ^a ^b 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): 407–441. doi:10.1111/boj.12062.
^ ^a ^b 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 (12): 2380–2387. doi:10.3732/ajb.1300303. PMID 24275705.
^ 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. Bibcode:2011PLoSO...624430L. doi:10.1371/journal.pone.0024430. PMC 3168474. PMID 21915326.
^ Cheek M, Onana JM (2024). "Keetia nodulosa sp. nov. (Rubiaceae - Vanguerieae) of West-Central Africa: bacterial leaf nodulation discovered in a fourth genus and tribe of Rubiaceae". Webbia. 79: 31–46. doi:10.36253/jopt-15946.
^ Sieber S, Carlier A, Neuburger M, Grabenweger G, Eberl L, Gademann K (2015). "Isolation and total synthesis of kirkamide, an aminocyclitol from an obligate leaf nodule symbiont" (PDF). Angewandte Chemie International Edition. 54 (27): 7968–7970. doi:10.1002/anie.201502696. PMID 26033226.
^ Yakubu MT, Oladiji AT, Akanji MA (2009). "Mode of cellular toxicity of aqueous extract of Fadogia agrestis (Schweinf. ex Hiern) stem in male rat liver and kidney". Human & Experimental Toxicology. 28 (8): 469–478. doi:10.1177/0960327109106973. PMID 19755438. S2CID 34724459.
^ Lamarck JBAPM de (1783). Encyclopédie Méthodique, Botanique. Vol. 1. Paris & Liège: Panckoucke & Plomteux. p. 602.
^ Gaertner J (1788). De Fructibus et Seminibus Plantarum. Vol. 1. Stuttgart: Typis Academiae Carolinae. p. 125.
^ Jussieu AL de (1789). Genera Plantarum. Paris: Herissant & Barrois. p. 206.
^ de Candolle AP (1807). "Mémoire sur le Cuviera". Annales du Muséum National d'Histoire Naturelle. 9: 216–222.
^ Dumortier BCJ (1829). Analyse des Familles des Plantes. Tournay: J Casterman. p. 32.
^ Robyns W (1928). "Tentamen monographiae Vangueriae generumque affinium". Bulletin du Jardin Botanique de l'état Bruxelles. 11 (1): 1–359. doi:10.2307/3666476. JSTOR 3666476.
^ Arriola AH, Paraguison LD, Alejandro GJD (2016). "Kanapia (Vanguerieae): a new endemic genus of Philippine Rubiaceae". Plant Systematics and Evolution. 302 (7): 911–920. doi:10.1007/s00606-016-1307-5. S2CID 14815565.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ Lantz H, Bremer B (2004). "Phylogeny inferred from morphology and DNA data: characterizing well-supported groups in Vanguerieae (Rubiaceae)". Botanical Journal of the Linnean Society. 146 (3): 257–283. doi:10.1111/j.1095-8339.2004.00338.x.
^ Lantz H, Bremer B (2005). "Phylogeny of the complex Vanguerieae (Rubiaceae) genera Fadogia, Rytigynia, and Vangueria with close relatives and a new circumscription of Vangueria". Plant Systematics and Evolution. 253 (1–4): 159–183. doi:10.1007/s00606-005-0313-9. S2CID 30867982.
^ Razafimandimbison SG, Lantz H, Mouly A, Bremer B (2009). "Evolutionary trends, major lineages, and new generic limits in the dioecious group of the tribe Vanguerieae (Rubiaceae): insights into the evolution of functional dioecy". Annals of the Missouri Botanical Garden. 96 (1): 161–181. doi:10.3417/2006191. S2CID 86065468.