On the evolutionary origins of the cacti. -Taxon 46: 217-232. 1997. -ISSN 0040-0262.Understanding evolutionary responses of plants to desert environments depends upon phylogenetic knowledge of desert plants. The diverse American desert family Cactaceae has been presumed, on the basis of distinctiveness, to be phylogenetically isolated and relatively ancient (> 65 million years old). Using maximum likelihood and parsimony analyses of the rapidly evolving internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA (nrDNA), we show that the cacti are phylogenetically nested among other aridity-adapted lineages of the angiosperm family Portulacaceae. The ITS divergence between pereskioid cacti and the genus Talinum (Portulacaceae) is less than that between many Portulacaceae genera. Synthesis of the ITS data with morphological and chloroplast DNA evidence suggests an origin of cacti in mid-Tertiary, c. 30 million years ago, and a later Tertiary diversification coincident with development of the American desert. This, in tum, implies that the diversification rate in cacti was much higher than in their nearest relatives. The present results illustrate the central role of phylogenetic reconstruction in ecological and evolutionary theory.

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Sequences of the ribosomal DNA internal transcribed spacer (ITS) region and the chloroplast DNA ycf3-trnS intergenic spacer were determined for 183 samples representing Chilean and non-Chilean taxa of western American Portulacaceae and their outgroups. The data refine previous inferences of generic circumscriptions and interrelations. In particular, the data reveal that an earlier circumscription of Cistanthe Spach is polyphyletic and also reveal a North American clade comprising Claytonia L., Lewisia Pursh, Lewisiopsis R. Govaerts, and Montia L. Within the South American genera, two patterns emerge from the data: (1) in some cases, interspecific divergence is remarkably low given the markers employed and estimated number of species; and (2) where divergence of one or both markers offer phylogenetic resolution, there is conflict between them and/or with morphology. The patterns can be evaluated in terms of two phenomena: (1) morphological/ecological radiation proceeding much faster than sequence divergence; and (2) frequent hybridization. In the latter case, the gene tree patterns may distort the true timing and cladistic pattern of morphological and ecological diversification. At present, the degree to which evidence for hybridization among the Chilean Portulacaceae will prove to be the rule or the exception is unclear. Nonetheless, spatial and temporal ecological patterns in Chile generally favor hybrid formation and persistence.