It is estimated that 5 million years of evolution separate Arabidopsis thaliana from its close relative Arabidopsis lyrata. The two taxa differ by many characteristics, and together they exemplify the differentiation of angiosperms into self-fertilizing and cross-fertilizing species as well as annual and perennial species. Despite their disparate life histories, the two species can be crossed to produce viable and vigorous hybrids exhibiting heterotic effects. Although pollen sterile, the hybrids produce viable ovules and were used as female parent in backcrosses to both parental species. The resulting backcross plants exhibited transgressive variation for a number of interesting developmental and growth traits as well as negative nuclear/ cytoplasmic interactions. Moreover, the genesis of a fertile amphidiploid neospecies, apparently by spontaneous somatic doubling in an interspecific hybrid, was observed in the laboratory. The mechanisms responsible for the generation of amphiploids and the subsequent evolution of amphiploid genomes can now be studied through direct observation using the large arsenal of molecular tools available for Arabidopsis.Plant growth and development have traditionally been studied by generating relevant mutations or by analyzing naturally occurring variants within a species. In only a few cases has the tremendous interspecies variation that was generated over the millions of years of evolution been used. In recent years, it has been increasingly recognized that natural variability is a major resource that could complement traditional approaches. Thus, in the model plant Arabidopsis, intraspecific genetic variation has been noted among different geographical isolates, and this variation, which is largely quantitative in nature, is being subjected to analytical methods developed for the analysis of quantitative trait loci in crop plants (for review, see Alonso-Blanco and Koornneef, 2000). However, the enormous store of natural variation that is manifest in interspecies differences has remained largely untapped.Wide crosses and interspecific hybridizations have been used to investigate the genetic basis of complex traits that differentiate varieties within a species as well as related species in several plant families (Doebley et al