In a tristylic species three types of individuals, distinguished by their flower morphology, occur. One form has a long style, while the anthers are in two whorls, one very low down, and one about half way between the stigma and the low anther whorl. The three positions of the sex organs in this form (called long or L) define three positions that can be occupied by anthers or stigmas in the two other forms. In short (S) individuals, the style is very short, and the anthers occupy mid (m) and long (1) positions; in mid (M) plants, the stigma occupies the middle position and the anthers the 1 and s positions. Darwin (1877) showed in several tristylic species that pollen from the different anther positions differed in its incompatibility reactions in such a way that a form with stigmas at a given level can be successfully fertilized only by pollen from anthers at the same level; this pollen will necessarily come from another individual, but it may come from either of the two forms other than that of the stigma-bearing plant in question. Tristyly is known in three families only (Lythraceae, Oxalidaceae and Pontederiaceae). Genera with tristylic species often also contain dimorphic and sometimes monomorphic populations (e.g., Darwin, 1877; Ornduff, 1964; Mulcahy, 1964; Lewis and Rao, 1971; Weller and Denton, 1976; and Barrett, 1979). It seems unlikely that these represent stages in the evolution of tristyly, because in many such cases there are features that appear to be residual tristylic characters (e. g., Lewis and Rao, 1971; Weller, 1976; and Barrett, 1979). In Lythrum, nearly all Old World species are trimorphic with two whorls of stamens, or monomorphic annuals with one stamen set, while all New World species are dimorphic (probably L and M, according to a personal communication from Ornduff) and have one stamen set. Here again, it seems most likely that trimorphism is the ancestral condition, since the presence of two anther whorls is characteristic of all three families with tristyled members (Lewis and Rao, 1971). In the present paper, the non-tristylic species in tristyled genera are therefore regarded as secondarily derived from tristyly. Probably the commonest type of breakdown of tristyly is loss of the mid form, giving a dimorphic (L and S) population, but various other forms of breakdown are known in Oxalis (reviewed by Ornduff, 1972); and in the Pontederiaceae (Barrett, 1979), and the dimorphic Lythrums may represent another exception. The genetics of tristyly has been worked out in several species. The genetic control is basically the same in Lythrum salicaria (Fisher and Mather, 1943), in section Ionoxalis of the genus Oxalis (Weller, 1976a) and in Eichhornia crassipes (Barrett, 1978a), although Lythrum and Eichhornia are polyploids, while most of the Oxalis species are not. 0. rosea may, however, have a different system (Von Ubisch, 1926; Fyfe, 1956). In section Ionoxalis, two genes control tristyly. The L form has the genotype ssmm. The dominant allele M determines mid-level stigmas in ss plants, and S (which is dominant to s and epistatic to M) determines the short stigma position. Thus S plants could have any one of six possible genotypes, though several of these will not be found in natural tristylic populations. While it is clear that this genetic system gives a tristylic equilibrium population (Fisher, 1941; Finney, 1952), it is by no means obvious how it could evolve. It
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