Combretum Explained
Combretum, the bushwillows or combretums, make up the type genus of the family Combretaceae. The genus comprises about 272 species of trees and shrubs, most of which are native to tropical and southern Africa, about 5 to Madagascar, but there are others that are native to tropical Asia, New Guinea and the Bismarck Archipelago, Australia, and tropical America. Though somewhat reminiscent of willows (Salix) in their habitus, they are not particularly close relatives of these.
Ecology
Bushwillow trees often are important plants in their habitat. Savannahs in Africa, in particular those growing on granitic soils, are often dominated by Combretum and its close relative Terminalia. For example, C. apiculatum is a notable tree in the Angolan mopane woodlands ecoregion in the Kunene River basin in southern Africa.[1]
Other species of this genus are a major component of Southwestern Amazonian moist forests. This genus contains several species that are pollinated by mammals other than bats, which is quite rare indeed.[2] But most species are more conventionally pollinated by insects or birds.
Typhlodromus combretum, a mite of the family Phytoseiidae, was discovered on a bushwillow plant and is named after this genus. Other herbivores that eat Combretum foliage include the caterpillars of the Brown Awl (Badamia exclamationis) which is found on C. albidum, C. latifolium and C. ovalifolium; those of the Orange-tailed Awl (Bibasis sena) are recorded from C. extensum and C. latifolium.
Use by humans
Several species are used in African or Indian herbal medicine. Several species of this genus in Madagascar known collectively as voantamenaka or voatamenaka (from voa 'fruit' and menaka 'oil') are used in traditional Malagasy medicine as deworming remedies.[3] The class of chemical compounds known as combretastatins were first isolated from South African bushwillow (Combretum afrum), from which they get their name. One synthetic derivative, fosbretabulin disodium (combretastatin A4 phosphate), underwent preliminary study for the treatment of anaplastic thyroid cancer,[4] but it was not effective enough to progress to more advanced trials. C. molle is also recorded to contain antioxidants such as punicalagin, which is also found in pomegranates (Punica granatum), a somewhat related plant.
The botanist George Don studied this genus extensively. The Luvuvhu River in southern Africa was named after the river's bushwillows (C. erythrophyllum), locally known as muvuvhu.
Species
As of April 2021, there are 272 accepted species of Combretum:
- Combretum aculeatum
- Combretum acuminatum
- Combretum acutifolium
- Combretum acutum
- Combretum adenogonium
- Combretum adrianii
- Combretum afzelii
- Combretum alatum
- Combretum albiflorum
- Combretum albopunctatum
- Combretum alfredi
- Combretum andongense
- Combretum andradae
- Combretum angolense
- Combretum angustipetalum
- Combretum annulatum
- Combretum apetalum
- Combretum aphanopetalum
- Combretum apiculatum
- Combretum argenteum
- Combretum argyrotrichum
- Combretum assimile
- Combretum atropurpureum
- Combretum aureonitens
- Combretum auriculatum
- Combretum barbatum
- Combretum batesii
- Combretum bauchiense
- Combretum baumii
- Combretum bipindense
- Combretum blepharopetalum
- Combretum boinensis
- Combretum bracteatum
- Combretum bracteosum
- Combretum brassiciforme
- Combretum brevistylum
- Combretum brunneum
- Combretum butyrosum
- Combretum cacoucia
- Combretum afrum
- Combretum camporum
- Combretum capitatum
- Combretum capituliflorum
- Combretum capuronii
- Combretum carringtonianum
- Combretum caudatisepalum
- Combretum celastroides
- Combretum chinense
- Combretum chionanthoides
- Combretum cinereopetalum
- Combretum cinnabarinum
- Combretum clarense
- Combretum coccineum
- Combretum collinum
- Combretum comosum
- Combretum conchipetalum
- Combretum confertum
- Combretum × confusum
- Combretum congolanum
- Combretum constrictum
- Combretum contractum
- Combretum coriifolium
- Combretum coursianum
- Combretum cuspidatum
- Combretum decandrum
- Combretum decaryi
- Combretum deciduum
- Combretum demeusei
- Combretum discolor
- Combretum dolichopodum
- Combretum duarteanum
- Combretum echirense
- Combretum edwardsii
- Combretum elaeagnoides
- Combretum engleri
- Combretum eriogynum
- Combretum erosum
- Combretum erythrophloeum
- Combretum erythrophyllum
- Combretum esteriense
- Combretum evisceratum
- Combretum exalatum
- Combretum exannulatum
- Combretum excelsum
- Combretum exellii
- Combretum falcatum
- Combretum farinosum
- Combretum flammeum
- Combretum foliatum
- Combretum frangulifolium
- Combretum fruticosum
- Combretum fulvum
- Combretum fuscum
- Combretum gabonense
- Combretum germainii
- Combretum ghesquierei
- Combretum gillettianum
- Combretum glabrum
- Combretum glaucocarpum
- Combretum glutinosum
- Combretum goetzei
- Combretum goldieanum
- Combretum goossensii
- Combretum gordonii
- Combretum gossweileri
- Combretum gracile
- Combretum graciliflorum
- Combretum grandidieri
- Combretum grandiflorum
- Combretum griffithii
- Combretum harmsianum
- Combretum harrisii
- Combretum hartmannianum
- Combretum haullevilleanum
- Combretum hensii
- Combretum hereroense
- Combretum hilarianum
- Combretum holstii
- Combretum homalioides
- Combretum igneiflorum
- Combretum illairii
- Combretum imberbe
- Combretum indicum
- Combretum inflatum
- Combretum ivanii
- Combretum kasaiense
- Combretum kirkii
- Combretum klossii
- Combretum klotzschii
- Combretum kostermansii
- Combretum kraussii
- Combretum lanceolatum
- Combretum lanuginosum
- Combretum latialatum
- Combretum latifolium
- Combretum laxum
- Combretum lecardii
- Combretum leprosum
- Combretum lindense
- Combretum lisowskii
- Combretum littoreum
- Combretum llewelynii
- Combretum lokele
- Combretum longicollum
- Combretum longipilosum
- Combretum longispicatum
- Combretum longistipitatum
- Combretum louisii
- Combretum lukafuensis
- Combretum luxenii
- Combretum macrocalyx
- Combretum malabaricum
- Combretum mannii
- Combretum marginatum
- Combretum marquesii
- Combretum mellifluum
- Combretum meridionalis
- Combretum micranthum
- Combretum microphyllum
- Combretum mkuzense
- Combretum moggii
- Combretum molle
- Combretum monetaria
- Combretum mooreanum
- Combretum mortehanii
- Combretum mossambicense
- Combretum mucronatum
- Combretum multinervium
- Combretum mussaendiflorum
- Combretum mweroense
- Combretum nanum
- Combretum ndjoleense
- Combretum nigrescens
- Combretum nigricans
- Combretum nioroense
- Combretum niphophilum
- Combretum nusbaumeri
- Combretum oatesii
- Combretum obovatum
- Combretum obscurum
- Combretum octagonum
- Combretum olivaceum
- Combretum oudenhovenii
- Combretum ovalifolium
- Combretum oxygonium
- Combretum oxystachyum
- Combretum oyemense
- Combretum padoides
- Combretum paniculatum
- Combretum paradoxum
- Combretum paraguariense
- Combretum patelliforme
- Combretum paucinervium
- Combretum pavonii
- Combretum pecoense
- Combretum pellegrinianum
- Combretum pentagonum
- Combretum petrophilum
- Combretum phaeocarpum
- Combretum pilosum
- Combretum pisoniiflorum
- Combretum pisonioides
- Combretum platypterum
- Combretum polyanthum
- Combretum polystictum
- Combretum procursum
- Combretum psidioides
- Combretum punctatum
- Combretum purpureiflorum
- Combretum pyramidatum
- Combretum pyrifolium
- Combretum quadrangulare
- Combretum quadratum
- Combretum rabiense
- Combretum racemosum
- Combretum razianum
- Combretum recurvatum
- Combretum relictum
- Combretum rhodanthum
- Combretum riggenbachianum
- Combretum robustum
- Combretum robynsii
- Combretum rochetianum
- Combretum rohrii
- Combretum rotundifolium
- Combretum rovirosae
- Combretum roxburghii
- Combretum rupestre
- Combretum rupicola
- Combretum sanjappae
- Combretum scandens
- Combretum schumannii
- Combretum sericeum
- Combretum shivannae
- Combretum sordidum
- Combretum sphaeroides
- Combretum spinosum
- Combretum stenopterum
- Combretum stocksii
- Combretum struempellianum
- Combretum stylesii
- Combretum subglabratum
- Combretum subumbellatum
- Combretum sundaicum
- Combretum sylvicola
- Combretum tarquense
- Combretum tenuipetiolatum
- Combretum tessmannii
- Combretum tetragonocarpum
- Combretum tetralophoides
- Combretum tetralophum
- Combretum teuschii
- Combretum tibatiense
- Combretum tomentosum
- Combretum towaense
- Combretum trichophyllum
- Combretum trifoliatum
- Combretum ulei
- Combretum umbricola
- Combretum vendae
- Combretum vernicosum
- Combretum villosum
- Combretum violaceum
- Combretum virgatum
- Combretum viscosum
- Combretum wallichii
- Combretum wattii
- Combretum wilksii
- Combretum winitii
- Combretum xanthothyrsum
- Combretum youngii
- Combretum zenkeri
- Combretum zeyheri
Notes and References
- C.Michael Hogan. 2012. Kunene River. Topic ed. P.Saundry. Encyclopedia of Earth. National Council for Science and the Environment. Washington DC.
- Carthew. S. 1997. Non-flying mammals as pollinators. Trends in Ecology & Evolution. 12. 3. 104–108. 0169-5347. 10.1016/S0169-5347(96)10067-7. 21237993. 1997TEcoE..12..104C.
- Calopyxis . Dictionnaire des noms malgaches de végétaux . Editions Alzieu . 1999 . Pierre Boiteau . I . 196 . fr . Pierre . Boiteau .
- Web site: Combretastatin A4 However there is the possibility that a modified version of an isomer of aombretastatin-A4 can be rendered effective. After replacing adjacent carbon atoms with nitrogen atoms, using blue light, the isomer can be switched to a form that attacks microtubules of cancer cells. So far this has been tested in vitro with breast cancer cells. Phosphate in Treating Patients With Advanced Anaplastic Thyroid Cancer . 10 June 2010 . clinicaltrials.gov.