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(Last Modified On 8/12/2013)
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(Last Modified On 8/12/2013)
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Family
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CAPPARACEAE
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Contributor
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W. G. D'ARCY
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Description
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Erect or sprawling herbs, shrubs, or small trees (Lagerheim), fetid, glabrate; sap clear; stems branching. Leaves alternate, trifoliolate, the leaflets narrow, pinnately veined, subsessile; petiole long and slender; exstipulate. Inflorescences lax terminal racemes, the pedicels slender, subtended by somewhat distant narrow bracts. Flowers perfect, green, white, or yellow (6-)8(-9)-merous, the calyx lobes free, lanceolate to elliptic, glabrous, pulverulent or ciliate, petals oblong, stipitate; stamens inserted on a rudimentary disc, alternating with minute nec- taries, the filament pubescent, sometimes expanded and differentiated basally, sometimes unequal, the anthers ovate, basifixed, somewhat hastate, the apex (not the connective) sometimes differentiated into a point, papillose or sometimes minutely pubescent, longitudinally dehiscent; pollen tricolporate; ovary ellipsoidal to globose, imperfectly 6-8 locular, the placenta axile, protruding as arms into the locule, the ovules campylotropous, numerous; style stout or obsolete, the stigmas 6-8, short, dark hornlike processes. Fruit a subglobose berry, the style and -the stigmas usually persistent, the calyx usually deciduous; seeds small, numerous, reniform, the embryo curved, surrounded by scant endosperm.
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Habit
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herbs, shrubs, or small trees
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Distribution
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The Tovariaceae comprises one genus, Tovaria, widespread in tropical Amer- ica, in the Antilles known only from Jamaica.
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Note
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Plants of this group were first published in the Capparaceae, and were recognized as a tribe, Tovarieae, of that family by Triana and Planchon (1862). Eichler (1889) assigned the genus to the Papaveraceae, and Pax (1891) placed it in a family of its own where most textbooks now treat it, although Metcalfe and Chalke (1950) accepted Tovaria as a member of the Capparaceae without noting any reservations. It supposedly differs from other Capparaceae in its pleiomorphic central placenta, in its globose fruit, and in anatomical details alluded to but not contrasted by Lagerheim (1892). In aspect and form of the leaves and inflorescence it is similar to other genera of the Capparaceae, e.g., Cleome, Polanisia, etc., tending to become woodier than these but not more so than other members of the family. The central placenta is approached in the axile placenta of Styxis (= Roydsia Roxb.) (see Lotsy 1911) which has 3 locules in contrast to the usual 2 locules in Capparaceae and 6-8 locules in Tovaria. Lagerheim's study showed the seeds and embryogenesis to be fundamentally similar in the Capparaceae and Papaveraceae with a trend to reduction of nucellus becoming more pronounced in the Cruciferae and Resedaceae, two other related families. In this respect he found Tovaria to be somewhat intermediate between the Capparaceae and Papaver- aceae. Meager pollen information (Erdtmann 1952) to date and chemical data (Hegnauer 1973, Gibbs 1974) support affinity with or placement within the Capparaceae and argue against proximity to the Papaveraceae. Tovaria finds many plants of similar general appearance in the Capparaceae and strong odors are also common in that family. No members of the Papaveraceae have even vaguely similar form. The Papaveraceae are also distinguished from the Cap- paraceae including Tovaria by colored sap, bluish overall cast, lack of myrosin cells, and other features which Cronquist (1968) believed reflect a considerable phylogenetic distance between the two families. Mauritzon (1935) suggested that secretory cells in the parenchyma of Tovaria might be homologous with the latex tubes of the Papaveraceae, but this suggestion has not met subsequent endorsement. Goldblatt (1978) found the chromosomes of Tovaria to be n = 14, a number reported in two genera of Capparaceae. On balance, there is good evidence for including Tovaria in the Capparaceae and little for placing it near the Papaveraceae. Its affinities are best indicated by recognizing it as a sub- family, Tovarioideae, of the Capparaceae.
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Reference
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Cronquist, A. 1968. The evolution and classification of flowering plants. Boston. Eichler, A. G. 1889. Capparidaceae, Cruciferae, Fumariaceae, Papaveraceae in C. F. P. de Martius, Flora of Brasil 13(1): 238-343. Erdtmann, G. 1952. Pollen morphology and plant taxonomy. Vol. 1, Angio- sperms. Stockholm. Fawcett, W. & A. B. Rendle. 1914. Tovariaceae. Flora of Jamaica 3: 246-247. Gibbs, R. D. 1974. Chemotaxonomy of flowering plants. 4 vols. Montreal. Goldblatt, P. 1978. Chromosome numbers in three cytologically unknown New World families, Tovariaceae, Trigoniaceae, and Vivianaceae. Ann. Missouri Bot. Garden 65: 776-777. Hegnauer, R. 1973. Tovariaceae in Chemotaxonomie der Pflanzen 6: 530-531. Lagerheim, G. de. 1892. Zur Kenntniss der Tovariaceen. Ber. Deutsch. Bot. Ges. 10: 163-169. Lotsy, J. P. 1911. Vortrage fiber Botanische Stammesgeschichte 3(1): 915. Mauritzon, J. 1935. Embryologie einiger Capparidaceen sowie von Tovaria pendula. Ark. Bot. 26A (15): 1-14. Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the Dicotyledons. Oxford. Pax, F. 1891. Tovariaceae in A. Engler & K. Prantl, Die Natiirlichen Pflanzen- familien 3(2): 207-208. Triana, J. J. & J. E. Planchon. 1862. Prodr. Fl. Novo-Granat. Ann. Sci. Nat. ser. 4. 17: 88.
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