ADAPTIVE RADIATION IN THE HAWAIIAN SILVERSWORD ALLIANCE (COMPOSITAE-MADIINAE). I. CYTOGENETICS OF SPONTANEOUS HYBRIDS

The silversword alliance of the Hawaiian Islands comprises one of the most spectacular arrays of life forms known in a relatively small, clearly natural plant group. Perhaps the best known member of this alliance is the Haleakala silversword (Argyroxiphium macrocephalum Gray). However, A. macrocephalum is only one of six species in a genus which includes plants called greenswords as well as those called silverswords (cf. St. John, 1973). They ordinarily form large basal rosettes of attractive silver or green leaves for a number of years before they finally produce an elongated inflorescence. Argyroxiphium macrocephalum normally does not branch and dies after producing its massive inflorescence (cf. Kobayashi, 1973a). It is thus, under normal conditions, an essentially herbaceous, monocarpic plant. The floral heads of sword plants are characteristically large (>3 cm) and radiate. Argyroxiphium occurs primarily as a pioneer plant on volcanic pumice or in bogs and is restricted to the islands of Hawaii and Maui. In marked contrast to Argyroxiphium the related endemic genus Dubautia of this alliance (sensu Keck, 1936) is comprised largely of woody, branched shrubs that produce small (<1.5 cm), discoid heads year after year. However, the genus shows a truly remarkable spectrum of variation from somewhat herbaceous, low-growing forms through woody shrubs to trees. Moreover, the recently rediscovered D. latifolia (Gray) Keck from Kauai is a large liana. Altogether there are about 25 species of Dubautia, some of which, like Argyroxiphium may also be found as pioneers on volcanic pumice or in bogs, but others are found in dry forests or in rain forests. Its distribution extends throughout the entire system of major islands from Hawaii to Kauai. A third, related Hawaiian endemic genus, Wilkesia, is comprised of two species, one of which is often monocarpic and produces a rosette of leaves at the summit of a usually unbranched woody stem (cf. St. John, 1971). The second species branches rather freely from the base and presumably flowers repeatedly. The heads of Wilkesia are fairly large and discoid. Wilkesia is found on dry slopes, primarily in Waimea Canyon, only on the island of Kauai. The remarkable array of life forms within the silversword alliance exploits almost every conceivable terrestrial habitat in Hawaii. Its representatives occur from near sea level to timberline and from areas that receive less than 45 cm of annual precipitation to perhaps the wettest place on earth, receiving about 1,300 cm of annual precipitation. These habitats range from very recent lava flows on Hawaii to mature rain forests on Kauai. In spite of the spectacular morphological and ecological diversity exhibited by the members of the silversword alliance, the occurrence of many natural intergeneric, intersubgeneric, and interspecific hybrids (Sherff, 1935, 1944; Kobayashi, 1973b; and Carr, 1978a) attests to the fact that they form a natural, genetically cohesive group that has in all probability resulted from rapid evolutionary differentiation after a single colonization by a progenitor, possibly in less than 10 million years (see Macdonald and Abbott, 1970 for a discussion of Hawaiian geology). Collectively, these plants constitute what may be considered an unparalleled ex-