Euphorbiaceae

Manioc Manihot esculenta, copyright Arthur Chapman.


Belongs within: Malpighiales.
Contains: Glochidion, Macaranga, Sauropus, Bertya, Croton, Plukenetieae, Acalypha, Euphorbia.

The Euphorbiaceae are a diverse group of flowering plants, often containing latex. Many members of the family are succulent, and some can bear a strong superficial resemblance to cacti. Notable members include Manihot, a genus of Neotropical herbs, shrubs and trees with white latex and often deeply palmate leaves, of which manioc or cassava M. esculenta is widely grown as a food plant. The castor oil plant Ricinus communis is a large herb or shrub with palmately lobed leaves that is originally native to a range between eastern Africa and India but which has becomed naturalised in most tropical and subtropical regions of the world. Hevea is a Neotropical genus of trees with trifoliolate leaves, of which the Pará rubber tree H. brasiliensis is the most commercially significant source of latex for rubber.

Characters (from Flora of China): Trees, shrubs, or herbs, rarely woody or herbaceous lianas, monoecious or dioecious, indumentum of simple, branched, stellate, or gland-tipped hairs, peltate or glandular scales or stinging hairs, latex often present, clear, white, or colored; roots woody, rarely roots tuberous and stems succulent, sometimes spiny. Leaves alternate or opposite, rarely whorled; stipules usually present, often free, sometimes modified into spines or glands, deciduous or persistent; petioles long to short, sometimes with glands at apex or base; leaf blade simple, sometimes palmately lobed, rarely compound, or reduced to scales, margins entire or toothed, sometimes with distinct glands along margin and/or on surface, venation pinnate or palmate. Inflorescences axillary or terminal, flowers in cymes or fascicles, these often arranged along an elongated axis, branched or unbranched, forming a thyrse, in congested heads, or in a flowerlike cyathium with very reduced flowers enclosed within a ± cupular involucre; bracts sometimes petaloid. Flowers unisexual, within same inflorescence or in separate inflorescences, actinomorphic. Sepals (1-)3-6(-8), free or connate into calyx tube, valvate or imbricate, rarely absent. Petals free, often reduced or absent. Disk present or absent. Male flowers with disk intrastaminal or extrastaminal, entire to dissected. Stamens one to very many, hypogynous; filaments free or connate; anthers 2(-4)-locular, mostly dehiscing longitudinally, rarely transversely or by pores, introrse or extrorse; rudimentary ovary sometimes present. Female flowers rarely with staminodes; ovary superior, (1-)2-5(-20)-locular; placentation axile; ovules 1 or 2 per locule, anatropous or hemitropous; styles free or connate, entire or lobed, or multifid, lobes erect, horizontal or curved; stigma capitate, linear, fimbriate, fan-shaped or pinnatilobate. Fruit typically a capsule elastically dehiscent into 2-valved cocci from a persistent columella, sometimes a berry or drupe. Seeds 1 or 2 per locule; seed coat thin to indurate, sometimes fleshy to form a sarcotesta; caruncle sometimes present; aril sometimes present; endosperm present or absent; embryo straight to curved or folded; cotyledons usually broader than radical. x = 6-14.

<==Euphorbiaceae [Acalyphoideae]
    |--Neoscortechinia DL07
    |    |--N. angustifolia H03
    |    |--N. philippinensis H03
    |    `--N. sumatrensis H03
    `--+--Jatropha XR12
       |    |--J. curcas BB07
       |    `--J. gossypiifolia LK14
       |--+--Micrandra DL07 [incl. Cunuria E03]
       |  |    `--M. siphonoides DL07
       |  |--Hevea DL07
       |  |    |--H. brasiliensis FP01
       |  |    `--H. pauciflora DL07
       |  `--Manihot XR12
       |       |--M. aipi C55
       |       |--M. epruinosa RS98
       |       |--M. esculenta BB07
       |       |--M. flabellifolia H90
       |       |--M. glaziovii RS98
       |       |--M. pseudoglaziovii Mo99
       |       |--M. quinquepartita GA97
       |       `--M. utilissima [=Mandioca utilissima] MM96
       |--+--Suregada Rottler 1803 XR12, A-R03 [incl. Gelonium Willdenow 1806 non Gaertner 1791 A-R03]
       |  |--Crotonoideae GA97
       |  |    |  i. s.: Aleurites moluccana GA97, C06 [incl. A. triloba C06]
       |  |    |--Croton XR12
       |  |    `--+--Codiaeum variegatum XR12, DL07
       |  |       |--Ostodes paniculata DL07
       |  |       `--Trigonostemon XR12
       |  `--+--Omphalea Linnaeus 1759 XR12, GA97 [Omphaleae]
       |     |    |--*O. triandra Linnaeus 1759 GA97
       |     |    `--O. diandra Linnaeus 1759 [incl. O. megacarpa Hemsley 1897] GA97
       |     `--+--Tetrorchidium XR12
       |        `--Endospermum XR12
       |             |--E. formicarum N91
       |             |--E. medullosum NC91
       |             `--E. moluccanum XR12
       `--+--+--Moultonianthus XR12
          |  `--+--Conceveiba XR12
          |     `--+--+--Lasiocroton XR12
          |        |  `--Plukenetieae XR12
          |        `--+--Ricinus XR12
          |           |    |--R. communis DL07
          |           |    `--R. zanzibarensis P09
          |           `--+--Spathiostemon XR12
          |              `--Acalypha XR12
          `--+--Pimelodendron zoanthogyne XR12, DL07
             `--Euphorbioideae GA97
                  |  i. s.: Senefeldera GA97
                  |         Mabea GA97
                  |           |--M. brasiliensis H11
                  |           |--M. salicoides GA97
                  |           `--M. taquari L-M-HG08
                  |--Euphorbia XR12
                  `--+--Hura XR12
                     |--Maprounea guianensis DL07
                     `--Homalanthus Juss. 1824 XR12, A61 [=Omalanthus H90]
                          |--H. novo-guineensis LK14
                          |--H. polyandrus (Hooker) Cheeseman 1906 (see below for synonymy) A61
                          |--H. populifolius C16 [=Carumbium populifolium C16, Omalanthus populifolius H90]
                          |--H. populneus DL07
                          `--‘Omalanthus’ stillingifolius H90

Euphorbiaceae incertae sedis:
  Mercurialis PT01
    |--M. annua PT98
    `--M. perennis PT01
  Antidesma E03
    |--A. diandrum P03
    |--A. ghaesembilla VT13
    `--A. montanum K03
  Glochidion UB06
  Dichostemma H03
  Kairothamnus E03
  Gymnanthes lucida E03, J87
  Strophioblachia E03
  Agrostistachys capitata E03, H03
  Doryxylon E03
  Shirakiopsis sanchezii E03, H03
  Distichirops E03
  Homonoia E03
  Chaetocarpus Thwaites 1849 (nom. cons.) [Chaetocarpeae] KC01
  Baccaurea E03
    |--B. lanceolata K03
    `--B. puberula K03
  Macaranga E03
  Sauropus E03
  Acidocroton YY22
  Actinostemon E03
  Adenochlaena YY22
  Adenocline YY22
  Adenopeltis YY22
  Afrotrewia YY22
  Agyneja impubes YY22, V98
  Alcoceria YY22
  Algernonia YY22
  Alphandia E03
  Amanoa E03
  Anabaenella YY22
  Andrachne E03
  Angostylis YY22
  Anthostema YY22
  Aonikena YY22
  Apodiscus YY22
  Argithamnia YY22
  Argomuellera E03
  Aristogeitonia E03
  Avellantia YY22
  Baliospermum montanum E03, SR07
  Bertya YY22
  Blachia E03
  Bonania YY22
  Breynia E03
    |--B. cernua HM03
    |--B. nivosa B00a
    `--B. oblongifolia MM09
  Calpigyne YY22
  Cephalocroton E03
  Cephalocrotonopsis YY22
  Cephalomappa E03
  Chiropetalum YY22
  Chloradenia YY22
  Chondrostylis E03
  Chrozophora E03
    |--C. brocchiana AGF98
    |--C. plicata AGF98
    |--C. rottelari VB02
    `--C. tinctoria Y98
  Cladogynos E03
  Clavistylis YY22
  Cluytiandra YY22
  Cnesmone E03
  Colliguaya YY22
  Corythea YY22
  Crotonopsis YY22
  Dalembertia YY22
  Dicoelia E03
  Ditaxis E03
    |--D. californica H93
    |--D. clariana H93
    |--D. lanceolata H93
    |--D. neomexicana DL07
    `--D. serrata H93
  Dysopsis E03
  Eleutherostigma YY22
  Epiprinus E03
  Erismanthus E03
  Fragariopsis YY22
  Grimmeodendron YY22
  Hippomane [Hippomaneae] E03
    `--H. mancinella RJ11
  Hypocoton YY22
  Joannesia YY22
  Julocroton YY22
  Klaineanthus YY22
  Lasiococca E03
  Leidesia YY22
  Longetia YY22
  Lortia YY22
  Mareya E03
  Megistostigma E03
  Micrococca mercurialis E03, SR07
  Monadenium YY22
  Necepsia E03
  Neopalisaya YY22
  Neopycnocoma YY22
  Neoroepera YY22
  Nepenthandra YY22
  Ophthalmoblapton YY22
  Pachystroma YY22
  Pachystylidium E03
  Paracroton pendulus E03, SR07
    |--P. p. ssp. pendulus SR07
    `--P. p. ssp. zeylanicus SR07
  Pedilanthus tithymaloides E03, K03
  Petalodiscus YY22
  Platygyne YY22
  Poranthera Rudge 1811 A61
    |--P. alpina Cheesem. ex Hooker 1881 A61
    |--P. coerulea LK14
    |--P. corymbosa C08
    |--P. ericifolia H90
    |--P. huegelii GK00
    `--P. microphylla Brong. 1828 A61
  Pseudanthus H90
    |--P. divaricatissimus H90
    |--P. orientalis H90
    |--P. ovalifolius H90
    |--P. pimeleoides H90
    `--P. virgatus GK00
  Pterococcus Hasskarl 1842 (nom. cons.) FT93
  Pycnocoma E03
  Ramelia YY22
  Ricinocarpos LK14
    |--R. bowmanii H90
    |--R. marginatus LK14
    |--R. pinifolius L09
    |--R. rosmarinifolius LK14
    |--R. speciosus H90
    `--R. trichophyllus LK14
  Sagotia E03
  Schizostigma YY22
  Sebastiania YY22
  Seidelia YY22
  Speranskia YY22
  Sphaerostylis YY22
  Stenadenium YY22
  Stillingia E03
    |--S. linearifolia H93
    |--S. paucidentata H93
    `--S. spinulosa H93
  Sumbavia YY22
  Sumbaviopsis E03
  Synadenium YY22
  Syndyophyllum E03
  Tannodia E03
  Tetraplandra YY22
  Tragiella E03
  Trisyngyne YY22
  Wetriaria YY22
  Wielandia YY22
  Adelia E03
  Adenophaedra E03
  Adriana KM08
    |--A. glabrata H90
    |    |--A. g. var. glabrata [incl. A. glabrata var. heterophylla] H90
    |    `--A. g. var. subglabra [incl. A. glabrata var. acerifolia] H90
    |--A. quadripartita SM06
    |--A. tomentosa B00b
    |    |--A. t. var. tomentosa B00b
    |    `--A. t. var. hookeri B00b [incl. A. hookeri var. glabriuscula H90]
    `--A. urticoides KM08
  Alchorneopsis portoricensis E03, SWK87
  Alcinaeanthus YY22
  Aparisthmium cordatum E03, L-M-HG08
  Aporosella YY22
  Astrocasia E03
  Athroandra YY22
  Baccaureopsis YY22
  Blumeodendron E03
  Bocquillonia YY22
  Buraeavia YY22
  Caelebogyne YY22
  Caryodendron E03
  Chascotheca YY22
  Cheilosa E03
  Chlamydojatropha YY22
  Choriophyllum YY22
  Clarorivinia YY22
  Coccoceras YY22
  Cocconerion YY22
  Coelediscus YY22
  Cometia YY22
  Conceveibastrum YY22
  Crotonogyne YY22
  Cyathogyne YY22
  Cyrtogonone YY22
  Cyrtogyne YY22
  Deuteromallotus YY22
  Discocarpus E03
  Discoclaoxylon E03
  Discocleidion YY22
  Discoglypremna E03
  Ditta myricoides J87, SWK87
  Everettiodendron YY22
  Fontainea E03
    |--F. australis H90
    `--F. oraria H90
  Gavarettia E03
  Givotia YY22
  Grossera YY22
  Hamilcoa YY22
  Hasskarlia YY22
  Hemicyclia YY22
  Holstia YY22
  Lepidoturus YY22
  Leptonemea YY22
  Leucocroton YY22
  Maesobotrya E03
  Manniophyton YY22
  Martretia YY22
  Mettenia YY22
  Mischodon YY22
  Neoboutonia E03
  Neojatropha YY22
  Neomanniophyton YY22
  Nulbraedia YY22
  Paivaeusa YY22
  Pausandra E03
  Plagiostyles YY22
  Platystigma YY22
  Podadenia YY22
  Poggeophyton YY22
  Polydragma YY22
  Protomegabaria YY22
  Pseudagrostistachys E03
  Pseudocroton YY22
  Pseudolachnostylis E03
  Richeria E03
  Ricinodendron E03
  Savia E03
  Secretania YY22
  Spondianthus E03
  Staphysora YY22
  Tetracarpidium YY22
  Thecacoris E03
  Trewia nudiflora E03, HU08
  Trigonopleura E03
  Uapaca YY22
  Uranthera YY22
  Veconcites YY22
  Wetria E03
  Actephila E03
    |--A. latifolia H09
    `--A. lindleyi H90
  Alchornea [Alchornieae] E03
    |--A. costaricensis OB11
    |--A. glandulosa MP00
    |--A. ilicifolia [incl. Coelebogyne ilicifolia] H90
    `--A. latifolia SWK87
  Amperea GK00
    |--A. simulans GK00
    |--A. volubilis GK00
    `--A. xiphoclada [incl. A. spartioides] H90
  Angostylidium YY22
  Baloghia H90
    |--B. inophylla [incl. B. lucida] H90
    `--B. marmorata H90
  Beyeria KM08
    |--B. brevifolia G04b
    |--B. cinerea KM08
    |--B. lasiocarpa H90
    |--B. leschenaulti G76
    |--B. opaca H90
    `--B. viscosa N91
  Caperonia E03
  Claoxylon H90
    |--C. australe H90
    `--C. hillii LK14
  Cleidion E03
  Cleistanthus UB06
    |--C. collinus S02
    |--C. cunninghamii H90
    `--C. sankunnianus Sivar. & Indu Balach. 1985 UB06
  Cordemoya YY22
  Dimorphocalyx australiensis LK14
  Garcia E03
  Lautembergia YY22
  Lebidieropsis YY22
  Melanolepis E03
  Reverchonia YY22
  Sapium E03
    |--S. buchii J87
    |--S. caribaeum SWK87
    |--S. haematospermum F11
    |--S. haitiense J87
    |--S. indicum R35
    |--S. jamaicense SWK87
    |--S. japonicum [=Excoecaria japonica] LO98
    |--S. laurocerasus SWK87
    `--S. sebiferum MH98
  Securinega BB07
    |--S. leucopyrus RaS02
    `--S. virosa BB07
  Tanodia YY22
  Astraea lobata [=Croton lobatum] C55
  Mallotus E03
    |--M. claoxyloides H90
    |--M. discolor H90
    |--M. dispersus LK14
    |--M. japonicus LO98
    |--M. nesophilus VT13
    |--M. paniculatus B00b
    |--M. philippensis HU08
    |--M. polyadenos B00a
    `--M. repandus P03
  Triadica E03
  Microstachys chamaelea LK14
  Margaritaria dubium-traceyi E03, LK14
  Ptychopyxis E03
  Balakata luzonica E03
  Leptopus SK05
    |--L. australis SK05
    |--L. decaisnei Ma99
    `--L. sanjappae Sumathi, Karthigeyan et al. 2005 SK05
  Annesijoa E03
  Borneodendron E03
  Botryophora E03
  Choriceras E03
  Clonostylis E03
  Elateriospermum E03
  Loerzingia E03
  Neotrewia E03
  Octospermum E03
  Reutalis E03
  Richeriella E03
  Tapoides E03
  Vernicia fordii [=Aleurites fordii] H90
  Meineckia parvifolia E03, SR07
  Spirostachys E03
    |--S. africana Ru-S02
    `--S. patagonica O06
  Cavacoa E03
  Crotonogynopsis E03
  Mildbraedia E03
  Neoholstia E03
  Paranecepsia E03
  Sclerocroton E03
  Cnidoscolus E03
    |--C. aconitifolius M-FR98
    |--C. quercifolius BV09
    `--C. vitifolius C-SQ08
  Argythamnia E03
  Dendrothrix E03
  Sandwithia E03
  Senefelderopsis croizatii GA97
  Celianella E03
  Glycydendron E03
  Jablonskia E03
  ‘Diplomorpha’ Griffith 1854 non Meisner 1841 FT93
  Hyeronima MP00
    |--H. alchorneoides MP00
    |--H. clusioides SWK87
    `--H. domingensis J87
  Calycopeplus LK14
    |--C. collinus LK14
    `--C. paucifolius G04a
  Erythrococca trichogyne DL07
  Monotaxis DL07
    |--M. bracteata DL07
    |--M. linifolia N91
    |--M. macrophylla H90
    |--M. occidentalis GK00
    `--M. tenuis LK14
  Koilodepas DL07
    |--K. batamense DL07
    `--K. longifolium K03
  Bernardia DL07
    |--B. incana BT72
    `--B. myricifolia DL07
  Crepetocarpon perkinsii XR12
  Excoecaria LK14
    |--E. agallocha LK14
    |--E. dallachyana H90
    |--E. ovalis LK14
    `--E. parvifolia LK14
  Gavarretia GA97
  Eremocarpus setigerus H93
  Securinegoxylon biseriatum Mädel 1962 CBH93
  Crotonopollis burdwanensis CBH93
  Retitricolporites irregularis CBH93
  Friesia Spreng. 1818 KC01
  Galorrheus Haw. 1812 KC01
  Heterochlamys Turcz. 1843 KC01
  Siphonia Rich. ex Schreb. 1791 KC01
  Trichosterigma Klotzsch & Garcke 1859 KC01
  Zygospermum Thwaites ex Baill. 1858 KC01

Homalanthus polyandrus (Hooker) Cheeseman 1906 [=Carumbium polyandrum Hooker 1864; incl. H. nutans Hooker 1856 (preoc.)] A61

*Type species of generic name indicated

REFERENCES

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