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The polyploid Hawaiian silversword alliance (Asteraceae), a spectacular example of adaptive radiation in plants, was shown previously to have descended from North American tarweeds of the Madia/Raillardiopsis group, a primarily diploid assemblage. The origin of the polyploid condition in the silversword alliance was not resolved in earlier biosystematic, cytogenetic, and molecular studies, apart from the determination that polyploidy in modern species of Madia/Raillardiopsis arose independent of that of the Hawaiian group. We determined that two floral homeotic genes, ASAP3/TM6 and ASAP1, are found in duplicate copies within members of the Hawaiian silversword alliance and appear to have arisen as a result of interspecific hybridization between two North American tarweed species. Our molecular phylogenetic analyses of the ASAP3/TM6 loci suggest that the interspecific hybridization event in the ancestry of the Hawaiian silversword alliance involved members of lineages that include Raillardiopsis muirii (and perhaps Madia nutans) and Raillardiopsis scabrida. The ASAP1 analysis also indicates that the two species of Raillardiopsis are among the closest North American relatives of the Hawaiian silversword alliance. Previous biosystematic evidence demonstrates the potential for allopolyploid formation between members of the two North American tarweed lineages; a vigorous hybrid between R. muirii and R. scabrida has been produced that formed viable, mostly tetraporate (diploid) pollen, in keeping with observed meiotic failure. Various genetic consequences of allopolyploidy may help to explain the phenomenal evolutionary diversification of the silversword alliance.

Two vigorous transoceanic, bigeneric hybrids, Dubautia laevigata (n = 14) X Raillardiopsis muirii (n = 8) and [Dubautia knudsenii X laxa] (n = 14) X Madia bolanderi (n = 6), and one vigorous transoceanic trigeneric hybrid, Dubautia scabra (n = 14) X [M. bolanderi X R. muirii], between mainland tarweeds and Hawaiian silversword allies were artificially produced and subjected to cytogenetic analysis. In addition to univalents, - 46-80% of the microsporocytes scored from these hybrids exhibited from one to four bivalents. However, some of the bivalents scored in the second bigeneric hybrid represented infragenomic association of Dubautia chromosomes. Stainable pollen of these hybrids ranged from 4.4 to 49%, mostly comprising large, tetracolporate, apparently diploid grains. The functionality of such grains was demonstrated in the primary hybrid M. bolanderi X R. muirii that was used to produce the trigeneric hybrid, and suggests the possible mode of origin of the Hawaiian genome via allopolyploidy. Illustrations of parents and Fls indicate that the hybrids produced in this study generally exhibit intermediate character states. However, the phenotypes of the "ray" flowers in hybrids between discoid and radiate species were noticeably unpredictable; in one case intermediacy appeared to be expressed largely in quantitative terms, while in two others intermediacy appeared to be expressed largely in qualitative terms.

Chloroplast DNA restriction-site comparisons were made among 24 species of the Hawaiian silversword alliance (Argyroxiphium, Dubautia, and Wilkesia) and 7 species of North American perennial tarweeds in Adenothamnus, Madia, Raillardella, and Raillardiopsis (Asteraceae-Madiinae). These data and results from intergeneric hybridization indicated surprisingly close genetic affinity of the monophyletic Hawaiian group to two diploid species of montane perennial herbs in California, Madia bolanderi and Raillardiopsis muirii. Of 117 restriction-site mutations shared among a subset of two or more accessions, more than one-fifth (25 mutations) separated the silversword alliance, M. bolanderi, and Raillardiopsis from Adenothamnus and Raillardella. An additional 10 mutations distinguished the silversword alliance, M. bolanderi, and R. muirii from Adenothamnus, Raillardella, and Raillardiopsis scabrida. Phylogenetic analyses of these data and production of vigorous hybrids of the combinations Dubautia laevigata x R. muirii and (Dubautia knudsenii x Dubautia laxa) x M. bolanderi reinforce and refine Carlquist's hypothesis [Carlquist, S. (1959) Aliso 4, 171-236] that the Hawaiian silversword alliance arose from American tarweeds. Ultimate origin of silversword alliance chloroplast DNA from within the Californian-endemic paraphyletic genus Raillardiopsis was supported with high bootstrap confidence. Geologic considerations and the distribution of sporophytic self-incompatibility among these species demonstrate that the tarweed ancestor of the silverswords overcame (i) a dispersal barrier of at least 3900 km of open ocean and (ii) the breeding barrier of self-incompatibility.