Rhamnaceae

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


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

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, while the Zizipheae are found mostly in the West Indies and southern Asia (Schirarend & Köhler 1993).

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. 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. pubescens SK93
    |         `--D. toumatou Raoul 1844 (see below for synonymy) A61
    |--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 D01
    |    |    |--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 [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
    |    |--Condalia SK93
    |    |    |--C. buxifolia SK93
    |    |    |--C. lineata SK93
    |    |    |--C. mexicana SK93
    |    |    |--C. microphylla SK93
    |    |    `--C. obovata SK93
    |    `--Karwinskia SK93
    |         |--K. bicolor SK93
    |         |--K. glandulosa SK93
    |         |--K. humboldtiana SK93
    |         |--K. oblongifolia SK93
    |         |--K. orbiculata SK93
    |         |--K. rocana SK93
    |         `--K. subcordata SK93
    `--Rhamneae [Pomaderreae] SK93
         |--Rhamnus D01
         |--Ampeloziziphus amazonicus SK93
         |--Macrorhamnus faraloatra SK93
         |--Ceanothus SK93
         |--Pomaderris SK93
         |--Cryptandra SK93
         |--Siegfriedia darwinioides SK93
         |--Noltea SK93
         |    |--N. africana SK93
         |    `--N. natalensis SK93
         |--Scutia D01
         |    |--S. buxifolia SK93
         |    |--S. capensis SK93
         |    `--S. circumscissa SK93
         |--Hovenia SK93
         |    |--H. acerba D07
         |    |--H. dulcis D01
         |    `--H. tomentella SK93
         |--Lasiodiscus SK93
         |    |--L. fasciculiflorus SK93
         |    |--L. mannii SK93
         |    `--L. mildbraedii SK93
         |--Spyridium SK93
         |    |--S. globulosum SM06
         |    |--S. mucronatum G04b
         |    `--S. parvifolium SK93
         |--Frangula SK93
         |    |--F. alnus F05
         |    |--F. californica SK93
         |    |--F. palmeri SK93
         |    |--F. polymorpha SK93
         |    |--F. purshiana SK93
         |    `--F. sphaerosperma SK93
         |--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 D01
         |    |--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
         |--Alphitonia SK93
         |    |--A. excelsa M65
         |    |--A. franguloides SK93
         |    |--A. incana C78
         |    |--A. macrocarpa SK93
         |    |--A. petriei SK93
         |    |--A. philippinensis SK93
         |    |--A. ponderosa SK93
         |    |--A. xerocarpa SK93
         |    `--A. ziziphoides SK93
         `--Colubrina SK93
              |--C. arborescens SK93
              |--C. asiatica SK93
              |--C. berteroana SK93
              |--C. ehrenbergii SK93
              |--C. elliptica SK93
              |--C. greggii SK93
              |--C. oppositifolia SK93
              |--C. texensis SK93
              `--C. triflora SK93

Rhamnaceae incertae sedis:
  Mailothia YY22
  Apteron YY22
  Pleuranthodes YY22
    |--P. hillebrandii SK93
    `--P. orbiculare SK93
  Stenanthemum KM08
    |--S. divaricatum KM08
    |--S. liberum G04b
    `--S. stipulosum G04a
  Adolphia SK93
    |-¬-A. californica SK93
    `--A. infesta SK93
  Kentrothamnus SK93
  Emmenosperma SK93
    |--E. alphitonioides SK93
    `--E. micropetala SK93
  Hybosperma SK93
    |--H. spinosum SK93
    `--H. verrucosum SK93
  Nesiota elliptica SK93
  Schistocarpaea johnsonii SK93
  Smythea lanceata SK93

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

*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).

[C78] Clunie, N. M. U. 1978. The vegetation. In Handbooks of the Flora of Papua New Guinea vol. 1 (J. S. Womersley, ed.) pp. 1-11. Melbourne University Press: Carlton South (Australia).

[D07] Dash, S. S. 2007. Useful plants of Kabi Sacred Grove, Sikkim. Bulletin of the Botanical Survey of India 49 (1-4): 79-88.

[D01] Doweld, A. B. 2001. The systematic relevance of fruit and seed structure in Bersama and Melianthus (Melianthaceae). Plant Systematics and Evolution 227: 75-103.

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

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261-282.

[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.

[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.

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

[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.

[M65] Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bulletin of the American Museum of Natural History 130: 1-362.

[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 The Himalayan Plants vol. 1 (H. Ohba & S. B. Malla, eds) 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 (1-4): 121-142.

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

[SM06] Semeniuk, C. A., L. A. Milne, P. Ladd & V. Semeniuk. 2006. Pollen in the surface sediments of wetlands in the Becher Point area, southwestern Australia: a baseline for use in interpreting Holocene sequences. Journal of the Royal Society of Western Australia 89 (1): 27-43.

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

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