Rhamnaceae

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

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

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. Both of these two tribes 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. The two tribes Rhamneae and Zizipheae are less firmly defined than the smaller tribes; the Rhamneae is found in most parts of the family's distribution except extratropical South America whereas the 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. denticulata P03
    |    |--V. ecorollata B00
    |    |--V. lucens SK93
    |    |--V. madraspatana SK93
    |    |--V. neocaledonica SK93
    |    |--V. viminalis SK93
    |    `--V. vitiensis SK93
    |--Gouanieae SK93
    |    |--Gouania SK93
    |    |--Reissekia smilacina SK93
    |    |--Helinus SK93
    |    |    |--H. mystacinus SK93
    |    |    `--H. scandens SK93
    |    `--Crumenaria SK93
    |         |--C. choretroides SK93
    |         |--C. glaziovii SK93
    |         `--C. polygaloides SK93
    |--Colletieae SK93
    |    |--Talguenea quinquenervia SK93
    |    |--Retanilla ephedra SK93
    |    |--Trevoa trinervia SK93
    |    |--Colletia SK93
    |    |    |--C. spinosa D03
    |    |    `--C. spinosissima SK93
    |    |--Discaria Hook. 1830 SK93, A61
    |    |    |--D. americana SK93
    |    |    |--D. australis A61
    |    |    |--D. discolor D03
    |    |    |--D. nitida H90
    |    |    |--D. pubescens SK93
    |    |    `--D. toumatou Raoul 1844 (see below for synonymy) A61
    |    `--Adolphia SK93
    |         |--A. californica SK93
    |         `--A. infesta SK93
    |--Rhamneae [Pomaderreae] SK93
    |    |--Rhamnus SK93
    |    |--Ampeloziziphus amazonicus SK93
    |    |--Macrorhamnus faraloatra SK93
    |    |--Ceanothus SK93
    |    |--Pomaderris SK93
    |    |--Cryptandra SK93
    |    |--Siegfriedia darwinioides SK93
    |    |--Alphitonia SK93
    |    |--Colubrina SK93
    |    |--Noltea SK93
    |    |    |--N. africana SK93
    |    |    `--N. natalensis SK93
    |    |--Scutia SK93
    |    |    |--S. buxifolia SK93
    |    |    |--S. capensis SK93
    |    |    `--S. circumscissa SK93
    |    |--Hovenia SK93
    |    |    |--H. acerba D07
    |    |    |--H. dulcis SK93
    |    |    `--H. tomentella SK93
    |    |--Lasiodiscus SK93
    |    |    |--L. fasciculiflorus SK93
    |    |    |--L. mannii SK93
    |    |    `--L. mildbraedii SK93
    |    |--Spyridium SK93
    |    |    |--S. cinereum H90
    |    |    |--S. eriocephalum H90
    |    |    |--S. globulosum SK93
    |    |    |--S. mucronatum G04b
    |    |    |--S. parvifolium SK93
    |    |    `--S. spathulatum Z94
    |    |--Trymalium SK93
    |    |    |--T. angustifolium SK93
    |    |    |--T. daltonii SK93
    |    |    |--T. floribundum M06
    |    |    |--T. ledifolium OS04
    |    |    |    |--T. l. var. ledifolium OS04
    |    |    |    `--T. l. var. rosmarinifolium OS04
    |    |    `--T. myrtillus G04b
    |    |--Sageretia SK93
    |    |    |--S. brandrethiana SK93
    |    |    |--S. elegans SK93
    |    |    |--S. michauxii SK93
    |    |    |--S. minutiflora SK93
    |    |    |--S. oppositifolia SK93
    |    |    |--S. pycnophylla SK93
    |    |    |--S. theezans SK93
    |    |    `--S. wrightii SK93
    |    `--Phylica SK93
    |         |--P. arborescens SK93
    |         |--P. buxifolia SK93
    |         |--P. cephalantha SK93
    |         |--P. cuspidata SK93
    |         |--P. paniculata SK93
    |         |--P. parviflora SK93
    |         |--P. virgata SK93
    |         `--P. willdenowiana SK93
    `--Zizipheae SK93
         |--Ziziphus SK93
         |--Reynosia SK93
         |--Doerpfeldia cubensis SK93
         |--Krugiodendron ferreum SK93
         |--Berchemiella berchemiaefolia SK93
         |--Chaydaia tonkinensis SK93
         |--Microrhamnus ericoides SK93
         |--Maesopsis eminii SK93
         |--Condaliopsis SK93
         |    |--C. lycioides SK93
         |    |--C. obtusifolia SK93
         |    `--C. parryi SK93
         |--Rhamnella SK93
         |    |--R. franguloides SK93
         |    |--R. obovalis SK93
         |    `--R. wilsonii SK93
         |--Rhamnidium SK93
         |    |--R. elaeocarpum SK93
         |    |--R. ellipticum SK93
         |    `--R. pruinosum SK93
         |--Auerodendron SK93
         |    |--A. acuminatum SK93
         |    |--A. cubense SK93
         |    |--A. glaucescens SK93
         |    `--A. reticulatum SK93
         |--Sarcomphalus SK93
         |    |--S. domingensis SK93
         |    |--S. havanensis SK93
         |    |--S. laurinus SK93
         |    |--S. microdictyus SK93
         |    `--S. reticulatus SK93
         |--Paliurus SK93
         |    |--P. hemsleyanus SK93
         |    |--P. hirsutus SK93
         |    |--P. orientalis SK93
         |    |--P. ramosissimus SK93 [incl. P. aubletia LO98]
         |    `--P. spina-christi SK93
         |--Berchemia SK93
         |    |--B. edgeworthii O88
         |    |--B. flavescens SK93
         |    |--B. giraldiana SK93
         |    |--B. philippinensis SK93
         |    `--B. scandens SK93
         |--Karwinskia SK93
         |    |--K. bicolor SK93
         |    |--K. glandulosa SK93
         |    |--K. humboldtiana SK93
         |    |--K. oblongifolia SK93
         |    |--K. orbiculata SK93
         |    |--K. rocana SK93
         |    `--K. subcordata SK93
         `--Condalia SK93
              |--C. buxifolia SK93
              |--C. globosa H93
              |    |--C. g. var. globosa H93
              |    `--C. g. var. pubescens H93
              |--C. lineata SK93
              |--C. megacarpa A19
              |--C. mexicana SK93
              |--C. microphylla SK93
              `--C. obovata SK93

Rhamnaceae incertae sedis:
  Mailothia YY22
  Apteron YY22
  Pleuranthodes SK93
    |--P. hillebrandii SK93
    `--P. orbiculare SK93
  Stenanthemum KM08
    |--S. divaricatum KM08
    |--S. liberum G04b
    `--S. stipulosum G04a
  Kentrothamnus SK93
  Emmenosperma SK93
    |--E. alphitonioides SK93
    |--E. cunninghamii LK14
    `--E. micropetala SK93
  Hybosperma SK93
    |--H. spinosum SK93
    `--H. verrucosum SK93
  Nesiota elliptica SK93
  Schistocarpaea johnsonii SK93
  Smythea lanceata SK93
  Eorhamnidium CBH93
  Rhamnoxylon CBH93
  ‘Tricolporopollenites’ haanradensis CBH93
  Scypharia Miers 1860 KC01

Discaria toumatou Raoul 1844 [=Notophoena toumatou Miers 1860; incl. D. australis var. apetala Hooker 1852] A61

*Type species of generic name indicated

REFERENCES

[A19] Acosta, L. E. 2019. A relictual troglomorphic harvestman discovered in a volcanic cave of western Argentina: Otilioleptes marcelae, new genus, new species, and Otilioleptidae, new family (Arachnida, Opiliones, Gonyleptoidea). PLoS One 14 (10): e0223828.

[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.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[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 9^th 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.

Last updated: 12 September 2020.