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A Remnant Greensword Population from Pu'u 'Alaea, Maui, with Characteristics of Argyroxiphium Virescens (Asteraceae)L

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A Remnant Greensword Population from Pu'u 'Alaea, Maui, with Characteristics of Argyroxiphium Virescens (Asteraceae)L Pacific Science (1998), vol. 52, no. 1: 61-68 © 1998 by University of Hawai'i Press. All rights reserved A Remnant Greensword Population from Pu'u 'Alaea, Maui, with Characteristics of Argyroxiphium virescens (Asteraceae)l 2 2 GERALD D. CARR AND ARTHUR C. MEDEIROS ,3 ABSTRACT: Two unusual greenswards occurring on Pu'u 'Alaea in 1989 re­ portedly possessed vegetative features characteristic of the presumed extir­ pated species Argyroxiphium virescens Hillebr. One of these Pu'u 'Alaea plants flowered in August 1989, allowing detailed comparisons with preserved speci­ mens of A. virescens as well as other species and hybrids of Argyroxiphium native to East Maui. These comparisons suggest that the unusual Pu'u 'Alaea greenswords represent remnants of hybridization between the now presumably extinct A. virescens and the more common HaleakaHi silversword, A. sandwi­ cense subsp. macrocephalum (A. Gray) Meyrat, that still occurs at and adjacent to this site. The estimated pollen fertility of 62% in the Pu'u 'Alaea plant is consistent with this interpretation. Recovery ofa few embryos from fruits ofthe plant that flowered in 1989 and the possibility of tissue culture of the remaining living plant at Pu'u 'Alaea apparently represent the last opportunities to con­ serve any vestige of A. virescens. THE HIGHER SLOPES and summit areas of Ha­ Originally described by Hillebrand (1888), leakala, East Maui, compose the aboriginal Argyroxiphium virescens was collected about home of three distinctive species of the genus a dozen times between 1840 and 1945, from Argyroxiphium. The most common and con­ Ko'olau Gap and adjacent slopes, and from spicuous of these is the well-known Halea­ the eastern summit of Haleakala. The most kala silversword, A. sandwicense subsp. mac­ recent specimen known is that of St. John rocephalum (A. Gray) Meyrat. This striking & Mitchell 21153, taken in 1945, from the plant is widespread around the summit area pass north of Kuiki [2135 m elev.] (Carr and is particularly abundant on the cinder 1985). Thirty years after its last documented cones within the crater. More restricted in observation in nature, A. virescens was con­ distribution is the greensword A. grayanum sidered possibly extinct (Fosberg and Herbst (Hillebr.) Degener, which, nevertheless, is 1975). Fifteen years later, with no evidence locally abundant, primarily in bogs and ad­ to the contrary, the federal government listed jacent areas of the northeastern slopes of the species as extinct in the wild (Anony­ Haleakala and in and around the summit mous 1990). Because there was no known bogs ofWest Maui. A second greensword, A. cultivation or seed bank of this species, it virescens Hillebr., was once found on the seemed that A. virescens would never again upper slopes of Haleakala. This unique East be seen. Maui greensword has been seen by few living Knowing the plight of the ill-fated green­ persons. sword, a few local botanists have been in­ trigued for several years by two unusual plants on Pu'u 'Alaea, Haleakala, that seemed to bear a striking resemblance to A. virescens. 1 Manuscript accepted 30 April 1997. In 1989, one of these plants flowered and 2Department of Botany, University of Hawai'i at material was collected for this study. In this Manoa, 3190 Maile Way, Honolulu, Hawai'i 96822­ 2279. paper we document the similarity of the Pu'u 3 National Biological Service, Haleakala National 'Alaea specimen to A. virescens and conclude Park Field Station, P.O. Box 369, Makawao, Hawai'i. that it likely represents a hybrid product be- 61 62 PACIFIC SCIENCE, Volume 52, January 1998 tween previously existing A. virescens and the mens (Figure 1). Although Degener made Haleakala silversword, A. sandwicense subsp. extensive collections (Degener 3528, Degener macrocephalum, which is still found in the 7489) from what may have been the last same vicinity. These plants may represent the substantial stronghold of A. virescens, his only hope of saving some of the genes that discussion of the species (Degener 1930) and were once embodied in the unique greensword his failure to acknowledge the existence of A. A. virescens. grayanum on Haleakala suggest that he con­ fused the two greenswords occurring on East Maui at that time. The brevity of Hille­ brand's original description of A. virescens MATERIALS AND METHODS and his restricted view ofA. grayanum, which The materials from the field that were he described as a new species of Wilkesia, used in this study were collected by A.C.M. probably fostered similar confusion among from a single flowering plant at Pu'u 'Alaea, other early collectors as well. However, the Haleakala, East Maui, in August 1989. The extremely narrow green leaves attributed to material collected included 14 basal leaves, A. virescens by Hillebrand (1888) in his orig­ 8 cauline leaves, and 7 flowering heads with inal description of this species are virtually peduncles partially or wholly intact. These diagnostic within the entire silversword alli­ structures were photographed and then pre­ ance and leave little doubt regarding proper served as dried, pressed herbarium material. application of this name. Carr (1985) pub­ Later, a representative flowering head was lished an expanded description of A. vires­ rehydrated and dissected to determine num­ cens that provided a better means to iden­ bers and sizes of component structures. The tify this poorly understood species. At the detailed description of the plant is based on same time he reported the occurrence of the these materials augmented by interpretations putative hybrids A. grayanum x A. virescens from photographs taken in the field. Herbar­ and A. sandwicense subsp. macroc;ephalum x ium specimens available at the B. P. Bishop A. virescens. Museum (BISH) and at the University of Ha­ wai'i Herbarium (HAw) provided a means for Description ofthe Pu'u'Alaea Greensword comparing the Pu'u 'Alaea plant with other representatives of Argyroxiphium. Pollen At first glance the Pu'u 'Alaea greensword from the heads of the Pu'u 'Alaea plant and (Figure 2) appears similar to small indi­ a nearby individual of A. sandwicense subsp. viduals of A. sandwicense subsp. macro­ macrocephalum collected on the same day cephalum, but with reduced pubescence re­ was mounted on a microscope slide in cotton sulting in a distinctly greener (versus silver) blue (Maneval 1936). The stain reaction of overall appearance. However, inspection of more than 500 grains was scored, and stain­ the leaves (Figures 3-7), flowering heads ability was considered an estimate of pollen (Figure 8), and florets (Figures 9-10) reveals fertility in these plants. The herbarium mate­ a number of features by which the Pu'u rial of the Pu'u 'Alaea plant will be deposited 'Alaea greensword differs from any extant at BISH. species. The very narrow leaves with hairy, but not silvery-floccose surfaces; the well­ developed, nearly white rays; the nearly total absence of pappus; and other, less obvious RESULTS AND DISCUSSION attributes compose a unique combination of features not known in any other member Argyroxiphium virescens of the genus Argyroxiphium. To facilitate de­ There is no convincing evidence that this tailed comparisons, a comprehensive char­ greensword was ever very common or widely acterization of the Pu'u 'Alaea greensword distributed. Its historical occurrence in only a in the same format as existing species de­ few sites is documented by herbarium speci- scriptions (Carr 1985) is presented below. See MAMALU "'- lAY '000 CONTOUR INTERVAL = 1000 FEET 0 5 10 15 20 MILES I I i i I I I I II I I 0 8 18 24 32 KM * Argyroxiphium sandwicense SSp. macrocephalum o Argyroxiphium grayanum o Argyroxiphium virescens -I- PUIU I Alaea greensward • Other putative A. s. ssp. m. x A. v. hybrids FIGURE 1. Locations of populations of Argyroxiphium on the summit of East Maui. 64 PACIFIC SCIENCE, Volume 52, January 1998 FIGURES 3-7. Morphology of leaves of East Maui species of Argyroxiphium: (3) A. grayanum, narrowly el­ liptic-ligulate, broadest above the middle; (4) A. vires­ cens, linear-ligulate, attenuate near the base; (5-6) Pu'u 'Alaea greensword: (5) whole leaf, linear-ligulate; (6) en­ larged portion ofleaf (note visibility ofveins through the layer of hairs); (7) A. sandwicense subsp. macrocephalum, linear-Ianceolate, dilated toward base. FIGURE 2. Pu'u 'Alaea greensword, in situ, August 90, disposed in a relatively lax racemiform 1989. capitulescence about 58 cm long and 24 cm wide, the peduncles ascending, about 8-12 cm long, distally recurved, glandular villous Materials and Methods for clarification of and also glandular puberulent to subfarina­ the material sampled. ceous; phyllaries (involucral bracts) mostly Erect, unbranched monocarpic rosette 9-12, about 14-17 mm long, linear-elliptic to shrub about 85 cm tall at flowering time. linear-lanceolate, glandular hirsutulous with Leaves closely disposed in a helical fashion, trichomes to 2 mm long, proximally adnate 19-24 cm long, 4-5 mm wide at midpoint, to the receptacular cup, the margins enfold­ linear-ligulate, straight or slightly arcuate­ ing the ray achenes; ray florets mostly 9-12, ascending, scarcely succulent, nearly flat, co­ the corolla narrowly tubular for 3-4.5 mm piously sericeous on both surfaces but not proximally, distally broadened into a limb totally obscuring the veins, pale greenish 11.5-13 mm long and about 4 mm wide when fresh, drying silvery gray; margins bearing 2-3 apical lobes 2-3 mm in length, entire, sericeous; apex acute, attenuate from the tubular portion sparsely puberulent and above the midpoint; base linear from mid­ sometimes minutely glandular, pale greenish point or above, abruptly dilated near point below and reddish above, the limb very of insertion; venation of 5-7 parallel nerves, pale greenish white, the pappus obsolete, the partially obscured by vesture.
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