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Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids II
Ordo: Myrtales

Familia: Combretaceae
Subfamilia: Combretoideae
Tribus: Combreteae
Subtribus: Combretinae
Genus: Combretum
Species:
a

C. aculeatum – C. acuminatum – C. acutifolium – C. acutum – C. adenogonium – C. adrianii – C. alatum – C. albidum – C. albiflorum – C. albopunctatum – C. alfredii – C. ambongense – C. andradae – C. angolense – C. angustipetalum – C. annulatum – C. apetalum – C. aphanopetalum – C. apiculatum – C. argenteum – C. argyrotrichum – C. assimile – C. aureonitens – C. auriculatum
b

C. basilei – C. bauchiense – C. bipindense – C. blepharopetala – C. boinensis – C. bracteosum – C. bracteosum – C. brassiciforme – C. brevistylum – C. butyrosum
c

C. cacouciaC. caffrumC. camporum – C. capitatum – C. capituliflorum – C. capuronii – C. carringtonianum – C. caudatisepalum – C. caudatum – C. celastroides – C. chionanthoides – C. cinereopetalum – C. cinnabarinum – C. clarense – C. coccineum – C. collinum – C. comosum – C. conchipetalum – C. confertum – C. confusum – C. congolanum – C. constrictum – C. contractum – C. coursianum – C. cuspidatum – C. cyclocarpum
d

C. decandrum – C. decaryanum – C. decaryi – C. deciduum – C. demeusei – C. densiflorum – C. discolor – C. dolichopetalum – C. duarteanum
e

C. echirense – C. edwardsii – C. elaeagnoides – C. engleri – C. erianthum – C. eriogynum – C. erosum – C. erythrophyllum – C. esteriense – C. eugeneanum – C. evisceratum – C. exalatum – C. exannulatum – C. exellii
f

C. falcatum – C. farinosum – C. foliatum – C. formosum – C. frangulifolium – C. fruticosum – C. fulvum – C. fuscum
g

C. gabonense – C. ghesquierei – C. gillettianum – C. glabrum – C. glaucocarpum – C. glutinosum – C. goetzei – C. goldieanum – C. goossensii – C. gordonii – C. gossweileri – C. gracile – C. graciliflorum – C. grandidieri – C. grandiflorum – C. griffithii
h

C. harrisii – C. hartmannianum – C. haullevilleanum – C. hensii – C. hereroense – C. hilariana – C. holstii – C. homalioides
i

C. igneiflorum – C. iliairii – C. imberbe – C. indicum – C. inflatum – C. ivanii
k

C. kachinense – C. kasaiense – C. kirkii – C. klossii – C. kraussii
l

C. lanceolatum – C. lasiocarpum – C. latialatum – C. latifolium – C. laurifolium – C. laxum – C. lecardii – C. leprosum – C. lineare – C. lisowskii – C. llewelynii – C. lokele – C. longicollum – C. longipilosum – C. longispicatum – C. louisii – C. lukafuensis – C. luxenii
m

C. macrocalyx – C. malabaricum – C. malifolium – C. mannii – C. marchettii – C. marginatum – C. mellifluum – C. meridionalis – C. micranthum – C. microphyllum – C. mildbraedii – C. mkuzense – C. moggii – C. molle – C. monetaria – C. mooreanum – C. mortehanii – C. mossambicense – C. mucronatum – C. multinervium – C. mweroense
n

C. nanum – C. ndjoleense – C. nelsonii – C. nigrescens – C. nigricans – C. nioroense – C. niphophilum – C. nusbaumeri
o

C. obovatum – C. obscurum – C. octagonum – C. olivaceum – C. oliviforme – C. oudenhovenii – C. oxygonium – C. oxystachyum – C. oyemense
p

C. padoides – C. paniculatum – C. paradoxum – C. paraguariense – C. paucinervium – C. pavonii – C. pecoense – C. pentagonum – C. perakense – C. petrophilum – C. pilosum – C. pisoniiflorum – C. pisonioides – C. platypetalum – C. platypterum – C. polyanthum – C. procursum – C. prostratum – C. psidioides – C. punctatum – C. purpureiflorum – C. pyramidatum – C. pyrifolium
q

C. quadrangulare – C. quadratum
r

C. rabiense – C. racemosum – C. razianum – C. recurvatum – C. relictum – C. robustum – C. robynsii – C. rochetianum – C. rohrii – C. rotundifolium – C. rovirosae – C. roxburghii – C. rupestre – C. rupicola
s

C. sanjappae – C. scandens – C. schumannii – C. schweinfurthii – C. sericeum – C. sordidum – C. spinosum – C. stenopterum – C. stocksii – C. stylesii – C. subglabratum – C. subumbellatum – C. sulcatum – C. sundaicum – C. sylvicola
t

C. tanaense – C. tarquense – C. tenuipetiolatum – C. ternatum – C. tetragonocarpum – C. tetralophoides – C. tetralophum – C. tibatiense – C. tomentosum – C. trichophyllum – C. trifoliatum
u

C. ulei – C. umbricola
v

C. velutinum – C. vendae – C. vernicosum – C. villosum – C. violaceum – C. viscosum
w

C. wallichii – C. wattii – C. wildemanii – C. wilksii – C. winitii – C. woodii
x

C. xanthothyrsum
y

C. yuankiangense
z

C. zenkeri – C. zeyheri
Source(s) of checklist:
Name

Combretum Loefl., Iter Hispan. 308. (1763) nom. cons.

Typus: C. fruticosum
(Loefl.) Stuntz

Synonyms

Heterotypic
Aetia Adans., Fam. 2: 84, 513. 1763.
Bucholzia Stadtm. ex Willemet, Ann. Bot. (Usteri) 18: 54 (1796)
Bureava Baill.
Cacoucia Aubl.
Campylogyne Welw. ex Hemsl.
Chrysostachys Pohl
Cristaria Sonn.
Embryogonia Blume
Forsgardia Vell.
Gonocarpus Ham.
Grislea L.
Hambergera Scop.
Kleinia Crantz, Inst. 2: 488. 1766, nom. illeg. non Mill. (1754).
Mekistus Lour. ex Gomes, Mem. Acad. Sc. Lisb. Cl. Sc. Pol. Mor. Bel.-Let. n. s. iv. (I), 29. 1868; Merrill in Trans. Am. Phil. Soc. n. s. xxiv. II. 16, 282. 1935, in syn.
Physopodium Desv.
Poivrea Comm. ex Thouars
Quisqualis L., Sp. Pl., ed. 2, 1: 556. 1762.
Schousboea Willd.
Seguiera Rchb. ex Oliv.
Sheadendron G.Bertol.
Sphalanthus Jack
Udani Adans., Fam. 2: (22). 1763.

References

Löfling, P. 1758. Iter Hispanicum 308.

Additional references

Maurin, O., Turner, I.M., Boatwright, J.S. & Christenhusz, M.J. 2020. New combinations in Combretaceae subtribe Combretinae from Africa and Asia. Phytotaxa 451(3): 231-237. DOI: 10.11646/phytotaxa.451.3.6 Paywall Reference page.

Links

Hassler, M. 2019. Combretum. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2019. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: 2019 October 24. Reference page.
Govaerts, R. et al. 2021. Combretum in Kew Science Plants of the World online. The Board of Trustees of the Royal Botanic Gardens, Kew. Published online. Accessed: 2021 October 4. Reference page.
International Plant Names Index. 2016. Combretum. Published online. Accessed: Apr. 7 2016.
Tropicos.org 2016. Combretum. Missouri Botanical Garden. Published online. Accessed: 7 Apr. 2016.
USDA, ARS, Germplasm Resources Information Network. Combretum in the Germplasm Resources Information Network (GRIN), U.S. Department of Agriculture Agricultural Research Service.

Vernacular names
Afrikaans: Boswilge
Esperanto: Kombreto
suomi: Kombretumit
中文: 使君子属

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.
Contents

1 Ecology
2 Use by humans
3 Species
4 References
5 External links

Ecology
Combretum albidum in Kinnerasani Wildlife Sanctuary, Andhra Pradesh, India

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.[2]

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.[3] 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. The class of chemical compounds known as combretastatins were first isolated from South African Bushwillow (Combretum caffrum), 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:[1]

Combretum aculeatum Vent.
Combretum acuminatum Roxb.
Combretum acutifolium Exell
Combretum acutum M.A.Lawson
Combretum adenogonium Steud. ex A.Rich.
Combretum adrianii Jongkind
Combretum afzelii G.Don
Combretum alatum Craib
Combretum albiflorum (Tul.) Jongkind
Combretum albopunctatum Suess.
Combretum alfredi Hance
Combretum andongense Engl. & Diels
Combretum andradae Exell & J.G.García
Combretum angolense Welw. ex M.A.Lawson
Combretum angustipetalum Chiov.
Combretum annulatum Craib
Combretum apetalum Wall. ex Kurz
Combretum aphanopetalum Engl. & Diels
Combretum apiculatum Sond.
Combretum argenteum Bertol.
Combretum argyrotrichum Welw. ex M.A.Lawson
Combretum assimile Eichler
Combretum atropurpureum Engl. & Diels
Combretum aureonitens Engl. & Gilg
Combretum auriculatum Engl. & Diels
Combretum barbatum G.Don
Combretum batesii Exell
Combretum bauchiense Hutch. & Dalziel
Combretum baumii Engl. & Gilg
Combretum bipindense Engl. & Diels
Combretum blepharopetalum Wickens
Combretum boinensis Jongkind
Combretum bracteatum (M.A.Lawson) Engl. & Diels
Combretum bracteosum (Hochst.) Engl. & Diels
Combretum brassiciforme Exell
Combretum brevistylum Eichler
Combretum brunneum Engl. & Diels
Combretum butyrosum (G.Bertol.) Tul.
Combretum cacoucia Exell ex Sandwith
Combretum caffrum (Eckl. & Zeyh.) Kuntze
Combretum camporum Engl.
Combretum capitatum De Wild. & Exell
Combretum capituliflorum Fenzl ex Schweinf.
Combretum capuronii Jongkind
Combretum carringtonianum Exell & J.G.García
Combretum caudatisepalum Exell & J.G.García
Combretum celastroides Welw. ex M.A.Lawson
Combretum chinense G.Don
Combretum chionanthoides Engl. & Diels
Combretum cinereopetalum Engl. & Diels
Combretum cinnabarinum Engl. & Diels
Combretum clarense Jongkind
Combretum coccineum (Sonn.) Lam.
Combretum collinum Fresen.
Combretum comosum G.Don
Combretum conchipetalum Engl. & Diels
Combretum confertum (Benth.) M.A.Lawson
Combretum × confusum Merr. & Rolfe
Combretum congolanum Liben
Combretum constrictum (Benth.) M.A.Lawson
Combretum contractum Engl. & Diels
Combretum coriifolium Engl. & Diels
Combretum coursianum (H.Perrier) Jongkind
Combretum cuspidatum Planch. ex Benth.
Combretum decandrum Jacq.
Combretum decaryi Jongkind
Combretum deciduum Collett & Hemsl.
Combretum demeusei De Wild.
Combretum discolor Taub.
Combretum dolichopodum Gilg
Combretum duarteanum Cambess.
Combretum echirense Jongkind
Combretum edwardsii Exell
Combretum elaeagnoides Klotzsch
Combretum engleri Schinz, De Wild. & T.Durand
Combretum eriogynum (H.Perrier) Jongkind
Combretum erosum Jongkind
Combretum erythrophloeum Gilg & Ledermann ex Engl.
Combretum erythrophyllum (Burch.) Sond.
Combretum esteriense Jongkind
Combretum evisceratum (H.Perrier) Jongkind
Combretum exalatum Engl.
Combretum exannulatum (O.Hoffm.) Engl. & Diels
Combretum excelsum Keay
Combretum exellii Jongkind
Combretum falcatum (Welw. ex Hiern) Jongkind
Combretum farinosum Kunth
Combretum flammeum Welw. ex Hiern
Combretum foliatum Craib
Combretum frangulifolium Kunth
Combretum fruticosum (Loefl.) Stuntz
Combretum fulvum Keay
Combretum fuscum Planch. ex Benth.
Combretum gabonense Exell
Combretum germainii Liben
Combretum ghesquierei Liben
Combretum gillettianum Liben
Combretum glabrum DC.
Combretum glaucocarpum Mart.
Combretum glutinosum Perr. ex DC.
Combretum goetzei Engl. & Diels
Combretum goldieanum F.Muell.
Combretum goossensii De Wild. & Exell
Combretum gordonii Jongkind
Combretum gossweileri Exell
Combretum gracile Schott
Combretum graciliflorum Stace
Combretum grandidieri Drake
Combretum grandiflorum G.Don
Combretum griffithii Van Heurck & Müll.Arg.
Combretum harmsianum Diels
Combretum harrisii Wickens
Combretum hartmannianum Schweinf.
Combretum haullevilleanum De Wild.
Combretum hensii Engl. & Diels
Combretum hereroense Schinz
Combretum hilarianum D.Dietr.
Combretum holstii Engl.
Combretum homalioides Hutch. & Dalziel
Combretum igneiflorum Rendón & R.Delgad.
Combretum illairii Engl.
Combretum imberbe Wawra
Combretum indicum (L.) DeFilipps
Combretum inflatum Jongkind
Combretum ivanii Jongkind
Combretum kasaiense Liben
Combretum kirkii M.A.Lawson
Combretum klossii Ridl.
Combretum klotzschii Welw. ex M.A.Lawson
Combretum kostermansii Exell
Combretum kraussii Hochst.
Combretum lanceolatum Pohl ex Eichler
Combretum lanuginosum G.Don
Combretum latialatum Engl.
Combretum latifolium Blume
Combretum laxum Jacq.
Combretum lecardii Engl. & Diels
Combretum leprosum Mart.
Combretum lindense Exell & Mildbr.
Combretum lisowskii Jongkind
Combretum littoreum (Engl.) Engl.
Combretum llewelynii J.F.Macbr.
Combretum lokele Liben
Combretum longicollum Jongkind
Combretum longipilosum Engl. & Diels
Combretum longispicatum (Engl.) Engl. & Diels
Combretum longistipitatum Jongkind
Combretum louisii Liben
Combretum lukafuensis De Wild.
Combretum luxenii Exell
Combretum macrocalyx (Tul.) Jongkind
Combretum malabaricum (Bedd.) Sujana, Ratheesh & Anil Kumar
Combretum mannii G.Lawson ex Engl. & Diels
Combretum marginatum Engl. & Diels
Combretum marquesii Engl. & Diels
Combretum mellifluum Eichler
Combretum meridionalis (H.Perrier) Jongkind
Combretum micranthum G.Don
Combretum microphyllum Klotzsch
Combretum mkuzense J.D.Carr & Retief
Combretum moggii Exell
Combretum molle R.Br. ex G.Don
Combretum monetaria Mart.
Combretum mooreanum Exell
Combretum mortehanii De Wild. & Exell
Combretum mossambicense (Klotzsch) Engl.
Combretum mucronatum Schumach. & Thonn.
Combretum multinervium Exell
Combretum mussaendiflorum Engl. & Diels
Combretum mweroense Baker
Combretum nanum Buch.-Ham. ex D.Don
Combretum ndjoleense Jongkind
Combretum nigrescens King
Combretum nigricans Lepr. ex Guill. & Perr.
Combretum nioroense Aubrév. ex Keay
Combretum niphophilum Gilg & Ledermann ex Engl.
Combretum nusbaumeri Jongkind & L.Gaut.
Combretum oatesii Rolfe
Combretum obovatum F.Hoffm.
Combretum obscurum Tul.
Combretum octagonum (H.Perrier) Jongkind
Combretum olivaceum Engl.
Combretum oudenhovenii Jongkind
Combretum ovalifolium Roxb. ex G.Don
Combretum oxygonium (Tul.) Jongkind
Combretum oxystachyum Welw. ex M.A.Lawson
Combretum oyemense Exell
Combretum padoides Engl. & Diels
Combretum paniculatum Vent.
Combretum paradoxum Welw. ex M.A.Lawson
Combretum paraguariense (Eichler) Stace
Combretum patelliforme Engl. & Diels
Combretum paucinervium Engl. & Diels
Combretum pavonii G.Don
Combretum pecoense Exell
Combretum pellegrinianum Exell
Combretum pentagonum M.A.Lawson
Combretum petrophilum Retief
Combretum phaeocarpum Mart.
Combretum pilosum Roxb. ex G.Don
Combretum pisoniiflorum (Klotzsch) Engl.
Combretum pisonioides Taub.
Combretum platypterum (Welw.) Hutch. & Dalziel
Combretum polyanthum Jongkind
Combretum polystictum Hiern
Combretum procursum Craib
Combretum psidioides Welw.
Combretum punctatum Blume
Combretum purpureiflorum Engl.
Combretum pyramidatum Desv.
Combretum pyrifolium (C.Presl) Kurz
Combretum quadrangulare Kurz
Combretum quadratum Craib
Combretum rabiense Jongkind
Combretum racemosum P.Beauv.
Combretum razianum K.G.Bhat
Combretum recurvatum Sujana, Ratheesh & Anil Kumar
Combretum relictum Hutch. & Dalziel
Combretum rhodanthum Engl. & Diels
Combretum riggenbachianum Gilg & Ledermann ex Engl.
Combretum robustum Jongkind
Combretum robynsii Exell
Combretum rochetianum A.Rich. ex A.Juss.
Combretum rohrii Exell
Combretum rotundifolium Rich.
Combretum rovirosae Exell
Combretum roxburghii Spreng.
Combretum rupestre Jongkind & Texier
Combretum rupicola Ridl.
Combretum sanjappae Chakrab. & Lakra
Combretum scandens Liben
Combretum schumannii Engl.
Combretum sericeum G.Don
Combretum shivannae Gholave, Kambale, Lekhak & S.R.Yadav
Combretum sordidum Exell
Combretum sphaeroides (Tul.) Jongkind
Combretum spinosum Bonpl.
Combretum stenopterum Exell
Combretum stocksii Sprague
Combretum struempellianum Gilg & Ledermann ex Engl.
Combretum stylesii O.Maurin, Jordaan & A.E.van Wyk
Combretum subglabratum De Wild.
Combretum subumbellatum (Baker) Jongkind
Combretum sundaicum Miq.
Combretum sylvicola O.Maurin
Combretum tarquense Clark
Combretum tenuipetiolatum Wickens
Combretum tessmannii Gilg ex Engl.
Combretum tetragonocarpum Kurz
Combretum tetralophoides Slooten
Combretum tetralophum C.B.Clarke
Combretum teuschii O.Hoffm.
Combretum tibatiense Gilg & Ledermann ex Engl.
Combretum tomentosum G.Don
Combretum towaense Engl. & Diels
Combretum trichophyllum Baker
Combretum trifoliatum Vent.
Combretum ulei Exell
Combretum umbricola Engl.
Combretum vendae A.E.van Wyk
Combretum vernicosum Rusby
Combretum villosum (Tul.) Jongkind
Combretum violaceum (Tul.) Jongkind
Combretum virgatum Welw. ex M.A.Lawson
Combretum viscosum Exell
Combretum wallichii DC.
Combretum wattii Exell
Combretum wilksii Jongkind
Combretum winitii Craib
Combretum xanthothyrsum Engl. & Diels
Combretum youngii Exell
Combretum zenkeri Engl. & Diels
Combretum zeyheri Sond.

References

"Combretum Loefl". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 11 April 2021.
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. doi:10.1016/S0169-5347(96)10067-7. ISSN 0169-5347. PMID 21237993.
"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". clinicaltrials.gov.

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