Can the complex phenotypes that characterize naturally occurring hybrid species be re-created in early-generation artificial hybrids? We address this question with three homoploid hybrid species (Helianthus anomalus, Helianthus deserticola, Helianthus paradoxus) and their ancestral parents (Helianthus annuus, Helianthus petiolaris) that are phenotypically distinct and ecologically differentiated. These species, and two synthetic hybrid populations of the ancestral parents, were characterized for morphological, physiological, and life-history traits in greenhouse studies. Among the synthetic hybrids, discriminant analysis identified a few individuals with the multitrait phenotype of the natural hybrid species: 0.7%-1.1% were H. anomalus-like, 0.5%-13% were H. deserticolalike, and only 0.4% were H. paradoxus-like. These relative frequencies mirror previous findings that genetic correlations are favorable for generating the hybrid species' phenotypes, and they correspond well with phylogeographic evidence that demonstrates multiple natural origins of H. deserticola and H. anomalus but a single origin for H. paradoxus. Even though synthetic hybrids with hybrid species phenotypes are rare, their phenotypic correlation matrices share most of the same principal components (eigenvectors), setting the stage for predictable recovery of hybrid species' phenotypes from different hybrid populations. Our results demonstrate past hybridization could have generated hybrid species-like multitrait phenotypes suitable for persistence in their respective environments in just three generations after initial hybridization.
Keywordshybridization; constraints; niche differentiation; adaptive evolution; speciation; Fleury hierarchy There are two contrasting mechanisms by which a new plant species can arise by hybridization (Grant 1981): allolyploid and homoploid hybrid speciation. Because hybridization must be initiated in sympatry, both forms of hybrid speciation require that reproductive barriers arise in the presence of gene flow. Allopolyploid speciation is straightforward because reproductive isolation is an instantaneous by-product of genome doubling. However, there is no assured means by which homoploid hybrid lineages become isolated from their parental species. Early authors (Stebbins 1957;Grant 1958;Templeton 1980) (McCarthy et al. 1995;Buerkle et al. 2000). Karyotypic divergence and spatial isolation are well documented in natural hybrid species (Gallez and Gottlieb 1982;Rieseberg 1991;Rieseberg et al. 1995;Hirai et al. 2000), and most hybrid species occupy habitats that are different or extreme compared with the parental species (Abbott 1992; Arnold 1997;Rieseberg 1997;Schwarzbach et al. 2001;Rosenthal et al. 2005).The simulation models above assume that habitat differentiation must occur early in the speciation process (Buerkle et al. 2000). Hybridization provides a plausible mechanism for rapid niche differentiation between the parental species and neohybrids. Recombinant hybrids are expected to have more phenotypi...