Bruguiera cylindrica, photographed by Teo Siyang.

Belongs within: Malpighiales.

The Rhizophoraceae are a largely pantropical family of trees and shrubs, with more southerly representatives in South America and Africa. As used herein, the family encompasses members of the Erythroxylaceae, treated by others as a separate taxon; as the sister relationship between Erythroxylaceae and Rhizophoraceae sensu stricto is regarded as well established, their separation or conjunction is largely a matter of taste. 'Erythroxylaceae' are shrubs and trees with entire leaves, at least partially interpetiolar stipules and usually fasciculate inflorescences with small flowers (Angiosperm Phylogeny website). Three of the four genera of Eythroxylaceae are restricted to Africa; the genus Erythroxylum is pantropical, and includes the two South American species (E. coca and E. novogranatense) from which cocaine is derived (Conn & Kerenga 1995). The Rhizophoraceae sensu stricto are shrubs and trees with opposite, often serrate leaves and large interpetiolar stipules (Angiosperm Phylogeny website). The best known members of the Rhizophoraceae sensu stricto are mangroves, with stilt roots allowing growth in anoxic muds and viviparous seeds germinating before the fruit is shed from the parent plant.

Synapomorphies (from the Angiosperm Phylogeny website): Tropane and pyrrolidine alkaloids, non-hydrolysable tannins present; sieve tube plastids with protein crystalloids; mucilage cells common; stomata paracytic; leaves involute, colleters present; inflorescence cymose; calyx valvate, corolla conduplicate, petals enclosing a stamen(s); filaments connate basally; median gynoecium adaxial, ovules epitropous, micropyle endostomal, endothelium present; antipodal cells ephemeral; fruit a septicidal capsule, calyx persistent; seeds arillate, exotestal cells enlarged, thick-walled, more or less tanniniferous; endosperm starchy, embryo green.

<==Rhizophoraceae [Rhizophoreae]
    |  i. s.: Endosteira T00
    |         Crossostylis T00
    |         Blepharistemma YY22
    |         Kandelia (de Candolle) Wight & Arnott 1834 [=Rhizophora sect. Kandelia de Candolle 1828] SLY03
    |           |--K. candel (Linnaeus) Druce 1914 (see below for synonymy) SLY03
    |           `--K. obovata Sheue, Liu & Yong 2003 SLY03
    |         Rhizophora KN03
    |           |--R. apiculata P88
    |           |--R. mangle SWK87
    |           |--R. mucronata KN03
    |           |--R. racemosa C55
    |           `--R. stylosa H03
    |         Ceriops KN03
    |           |--C. candolleana KN03
    |           `--C. tagal WF01
    |--Erythroxylaceae CK95
    |    |--Nectaropetalum CK95
    |    |--Aneulophus CK95
    |    |--Pinacopodium CK95
    |    `--Erythroxylum DL07
    |         |  i. s.: E. coca CK95
    |         |         E. novogranatense CK95
    |         |         E. obtusifolium VM03
    |         `--E. sect. Coelocarpus CK95
    |              |--E. cuneatum (Miq.) Kurz 1874 CK95
    |              `--E. ecarinatum Burck 1893 (see below for synonymy) CK95
    `--+--Carallia DL07
       |    |--C. brachiata C78
       |    `--C. integerrima H09
       `--Bruguiera DL07
            |--B. caryophylloides H09
            |--B. cylindrica KN03
            |--B. gymnorhiza [incl. B. rheedi] KN03
            |--B. hainesii H03
            `--B. parviflora H09

Erythroxylum ecarinatum Burck 1893 [incl. E. ecarinatum var. ledermannii Schulz 1924, E. salomonense White 1950] CK95

Kandelia candel (Linnaeus) Druce 1914 [=Rhizophora candel Linnaeus 1753, *K. rheedii Wight & Arnott 1834 (nom. illeg.)] SLY03

*Type species of generic name indicated


[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[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).

[CK95] Conn, B. J., & K. Kerenga. 1995. Erythroxylaceae. In Handbooks of the Flora of Papua New Guinea vol. 3 (B. J. Conn, ed.) pp. 56-60. Melbourne University Press: Carlton (Australia).

[DL07] Davis, C. C., M. Latvis, D. L. Nickrent, K. J. Wurdack & D. A. Baum. 2007. Floral gigantism in Rafflesiaceae. Science 315: 1812.

[H03] Heads, M. 2003. Ericaceae in Malesia: Vicariance biogeography, terrane tectonics and ecology. Telopea 10 (1): 311-449.

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

[KN03] Komai, F., & Y. Nasu. 2003. Four species of Olethreutinae (Lepidoptera: Tortricidae) associated with viviparous seedlings of the mangrove Rhizophoraceae in the Ryukyu Islands, Japan. Invertebrate Systematics 17: 75-87.

[P88] Polunin, I. 1988. Plants and Flowers of Malaysia. Times Editions: Singapore.

[SLY03] Sheue, C.-R., H.-Y. Liu & J. W. H. Yong. 2003. Kandelia obovata (Rhizophoraceae), a new mangrove species from eastern Asia. Taxon 52: 287-294.

[SWK87] Snyder, N. F. R., J. W. Wiley & C. B. Kepler. 1987. The Parrots of Luquillo: Natural history and conservation of the Puerto Rican parrot. Western Foundation of Vertebrate Zoology: Los Angeles.

[T00] Thorne, R. F. 2000. The classification and geography of the flowering plants: Dicotyledons of the class Angiospermae (subclasses Magnoliidae, Ranunculidae, Caryophyllidae, Dilleniidae, Rosidae, Asteridae, and Lamiidae). The Botanical Review 66: 441-647.

[VM03] Viswanathan, M. B. & U. Manikandan. 2003. A new species of Balsaminaceae, Impatiens tirunelvelica, from peninsular India. Bulletin of the Botanical Survey of India 45 (1-4): 189-194.

[WF01] Woinarski, J. C. Z., A. Fisher, K. Brennan, I. Morris & R. Chatto. 2001. Patterns of bird species richness and composition on islands off Arnhem Land, Northern Territory, Australia. Austral Ecology 26: 1-13.

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

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