Rhamnaceae

 Flowering branch of wild Irishman Discaria toumatou, photographed by P. B. Pelser.

Belongs within: Rosales.
Contains: Gouania, ZiziphusReynosiaCondalia, Rhamnus, Ceanothus, Pomaderris, Cryptandra, Alphitonia, Colubrina.

The Rhamnaceae are a family of plants with simple leaves and small flowers, in which the anthers are inserted opposite the petals (obhaplostemony). Most species are woody though the genus Crumenaria in the tribe Gouanieae is herbaceous. Other members of the Gouanieae and the Ventilagineae are climbers, with the Ventilagineae further characterised by the presence of a wing-like apical appendix on the fruit. The supple jack Ventilago viminalis of Australia may develop into a tree at maturity (Harden 1990). Both Gouanieae and Ventilagineae are tropical, with the Gouanieae pantropical and the Ventilagineae palaeotropical.

The Colletieae are a group of strongly armed trees and shrubs found mostly in extratropical South America, with Adolphia found in south-west North America and Discaria in Australasia. Discaria species are rigid shrubs with small, opposite leaves that are often shed early (Harden 1990).

The two tribes Rhamneae and Zizipheae are less firmly defined than the smaller tribes. Rhamneae are found in most parts of the family’s distribution except extratropical South America whereas Zizipheae are found mostly in the West Indies and southern Asia (Schirarend & Köhler 1993). Rhamneae have fruits that are berries or schizocarps with a thick testa whereas fruits of Zizipheae are drupes with a thin papery testa.

Characters (from the Angiosperm Phylogeny Website): Woody; chelidonic acid present; saponins, biflavonyls, benzylisoquinoline alkaloids present; myricetin, ellagic acid absent; libriform fibers present; lysigenous mucilage cavities present; leaves opposite or spiral, conduplicate(or -plicate) or involute, colleters present; flowers small, 4-5(-6)-merous; hypanthium present, calyx with a longitudinal median ridge adaxially, corolla cucullate (sometimes absent), enfolding androecium; infratectum granulate-intermediate; gynoecium more or less inferior, opposite sepals or odd member adaxial, style present or styles separate, stigma papillate; ovules epitropous or apotropous, outer integument 4-10 cells across, inner integument 3-4 cells across, parietal tissue 4-7 cells across, nucellar cap to 6 cells across, nucellus more or less protruding through micropyle, hypostase present; archesporium often multicellular, antipodals degenerate; calyx often deciduous; seeds often laterally flattened; testa and tegmen more or less multiplicative, endotegmen of cuboid cells, with scalariform thickenings (sometimes slightly lignified); endosperm present or absent, polyembryony common, embryo green.

<==Rhamnaceae
    |--Ventilago [Ventilagineae]SK93
    |    |--V. denticulataP03
    |    |--V. ecorollataB00
    |    |--V. lucensSK93
    |    |--V. madraspatanaSK93
    |    |--V. neocaledonicaSK93
    |    |--V. viminalisSK93
    |    `--V. vitiensisSK93
    |--GouanieaeSK93
    |    |--GouaniaSK93
    |    |--Reissekia smilacinaSK93
    |    |--HelinusSK93
    |    |    |--H. mystacinusSK93
    |    |    `--H. scandensSK93
    |    `--CrumenariaSK93
    |         |--C. choretroidesSK93
    |         |--C. glazioviiSK93
    |         `--C. polygaloidesSK93
    |--ColletieaeSK93
    |    |--Talguenea quinquenerviaSK93
    |    |--Retanilla ephedraSK93
    |    |--Trevoa trinerviaSK93
    |    |--ColletiaSK93
    |    |    |--C. spinosaD03
    |    |    `--C. spinosissimaSK93
    |    |--AdolphiaSK93
    |    |    |--A. californicaSK93
    |    |    `--A. infestaSK93
    |    `--Discaria Hook. 1830SK93, A61
    |         |--D. americanaSK93
    |         |--D. australisA61
    |         |--D. discolorD03
    |         |--D. nitidaH90
    |         |--D. pubescensSK93
    |         `--D. toumatou Raoul 1844 [=Notophoena toumatou Miers 1860; incl. D. australis var. apetala Hooker 1852]A61
    |--ZizipheaeSK93
    |    |--ZiziphusSK93
    |    |--ReynosiaSK93
    |    |--Doerpfeldia cubensisSK93
    |    |--Krugiodendron ferreumSK93
    |    |--Berchemiella berchemiaefoliaSK93
    |    |--Chaydaia tonkinensisSK93
    |    |--Microrhamnus ericoidesSK93
    |    |--Maesopsis eminiiSK93
    |    |--CondaliaSK93
    |    |--CondaliopsisSK93
    |    |    |--C. lycioidesSK93
    |    |    |--C. obtusifoliaSK93
    |    |    `--C. parryiSK93
    |    |--RhamnellaSK93
    |    |    |--R. franguloidesSK93
    |    |    |--R. obovalisSK93
    |    |    `--R. wilsoniiSK93
    |    |--RhamnidiumSK93
    |    |    |--R. elaeocarpumSK93
    |    |    |--R. ellipticumSK93
    |    |    `--R. pruinosumSK93
    |    |--AuerodendronSK93
    |    |    |--A. acuminatumSK93
    |    |    |--A. cubenseSK93
    |    |    |--A. glaucescensSK93
    |    |    `--A. reticulatumSK93
    |    |--SarcomphalusSK93
    |    |    |--S. domingensisSK93
    |    |    |--S. havanensisSK93
    |    |    |--S. laurinusSK93
    |    |    |--S. microdictyusSK93
    |    |    `--S. reticulatusSK93
    |    |--PaliurusSK93
    |    |    |--P. hemsleyanusSK93
    |    |    |--P. hirsutusSK93
    |    |    |--P. orientalisSK93
    |    |    |--P. ramosissimusSK93 [incl. P. aubletiaLO98]
    |    |    `--P. spina-christiSK93
    |    |--BerchemiaSK93
    |    |    |--B. edgeworthiiO88
    |    |    |--B. flavescensSK93
    |    |    |--B. giraldianaSK93
    |    |    |--B. philippinensisSK93
    |    |    `--B. scandensSK93
    |    `--KarwinskiaSK93
    |         |--K. bicolorSK93
    |         |--K. glandulosaSK93
    |         |--K. humboldtianaSK93
    |         |--K. oblongifoliaSK93
    |         |--K. orbiculataSK93
    |         |--K. rocanaSK93
    |         `--K. subcordataSK93
    `--Rhamneae [Pomaderreae]SK93
         |--RhamnusSK93
         |--Ampeloziziphus amazonicusSK93
         |--Macrorhamnus faraloatraSK93
         |--CeanothusSK93
         |--PomaderrisSK93
         |--CryptandraSK93
         |--Siegfriedia darwinioidesSK93
         |--AlphitoniaSK93
         |--ColubrinaSK93
         |--NolteaSK93
         |    |--N. africanaSK93
         |    `--N. natalensisSK93
         |--ScutiaSK93
         |    |--S. buxifoliaSK93
         |    |--S. capensisSK93
         |    `--S. circumscissaSK93
         |--HoveniaSK93
         |    |--H. acerbaD07
         |    |--H. dulcisSK93
         |    `--H. tomentellaSK93
         |--LasiodiscusSK93
         |    |--L. fasciculiflorusSK93
         |    |--L. manniiSK93
         |    `--L. mildbraediiSK93
         |--SpyridiumSK93
         |    |--S. cinereumH90
         |    |--S. divaricatumK90
         |    |--S. eriocephalumH90
         |    |--S. globulosumSK93
         |    |--S. mucronatumG04b
         |    |--S. parvifoliumSK93
         |    `--S. spathulatumZ94
         |--TrymaliumSK93
         |    |--T. angustifoliumSK93
         |    |--T. daltoniiSK93
         |    |--T. floribundumM06
         |    |--T. ledifoliumOS04
         |    |    |--T. l. var. ledifoliumOS04
         |    |    `--T. l. var. rosmarinifoliumOS04
         |    `--T. myrtillusG04b
         |--SageretiaSK93
         |    |--S. brandrethianaSK93
         |    |--S. elegansSK93
         |    |--S. michauxiiSK93
         |    |--S. minutifloraSK93
         |    |--S. oppositifoliaSK93
         |    |--S. pycnophyllaSK93
         |    |--S. theezansSK93
         |    `--S. wrightiiSK93
         `--PhylicaSK93
              |--P. arborescensSK93
              |--P. buxifoliaSK93
              |--P. cephalanthaSK93
              |--P. cuspidataSK93
              |--P. paniculataSK93
              |--P. parvifloraSK93
              |--P. virgataSK93
              `--P. willdenowianaSK93
Rhamnaceae incertae sedis:
  MailothiaYY22
  ApteronYY22
  PleuranthodesSK93
    |--P. hillebrandiiSK93
    `--P. orbiculareSK93
  StenanthemumKM08
    |--S. divaricatumKM08
    |--S. liberumG04b
    `--S. stipulosumG04a
  KentrothamnusSK93
  EmmenospermaSK93
    |--E. alphitonioidesSK93
    |--E. cunninghamiiLK14
    `--E. micropetalaSK93
  HybospermaSK93
    |--H. spinosumSK93
    `--H. verrucosumSK93
  Nesiota ellipticaSK93
  Schistocarpaea johnsoniiSK93
  Smythea lanceataSK93
  EorhamnidiumCBH93
  RhamnoxylonCBH93
  ‘Tricolporopollenites’ haanradensisCBH93
  Scypharia Miers 1860KC01

*Type species of generic name indicated

References

[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[D07] Dash, S. S. 2007. Useful plants of Kabi Sacred Grove, Sikkim. Bulletin of the Botanical Survey of India 49: 79–88.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8. Botany pp. 1–34. The University: Princeton (New Jersey).

[G04a] Gibson, N. 2004a. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35–47.

[G04b] Gibson, N. 2004b. Flora and vegetation of the Eastern Goldfields Ranges: part 7. Middle and South Ironcap, Digger Rock and Hatter Hill. Journal of the Royal Society of Western Australia 87 (2): 49–62.

[H90] Harden, G. J. (ed.) 1990. Flora of New South Wales vol. 1. New South Wales University Press.

[K90] Keighery, G. J. 1990. Vegetation and flora of Shark Bay, Western Australia. In: Berry, P. F., S. D. Bradshaw & B. R. Wilson (eds) Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee pp. 61–87. Western Australian Museum.

[KM08] Keighery, G. J., & W. Muir. 2008. Vegetation and vascular flora of Faure Island, Shark Bay, Western Australia. Records of the Western Australian Museum Supplement 75: 11–19.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[LO98] Lack, H. W., & H. Ohba. 1998. Die Xylothek des Chikusai Kato. Willdenowia 28: 263–276.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[M06] McCaw, W. L. 2006. Asplenium aethiopicum recolonises karri forest following timber harvesting and burning. Journal of the Royal Society of Western Australia 89 (3): 119–122.

[OS04] Obbens, F. J., & L. W. Sage. 2004. Vegetation and flora of a diverse upland remnant of the Western Australian wheatbelt (Nature Reserve A21064). Journal of the Royal Society of Western Australia 87 (1): 19–28.

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[P03] Paul, T. K. 2003. Botanical observations on the Purulia pumped storage hydropower project area, Bagmundi Hills, Purulia district, West Bengal. Bulletin of the Botanical Survey of India 45: 121–142.

[SK93] Schirarend, C., & E. Köhler. 1993. Rhamnaceae Juss. World Pollen and Spore Flora 17/18: 1–53.

[YY22] Yampolsky, C., & H. Yampolsky. 1922. Distribution of sex forms in the phanerogamic flora. Bibliotheca Genetica 3: 1–62.

[Z94] Zimmerman, E. C. 1994. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May. CSIRO Australia.

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