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Vegetative Anatomy of the Hawaiian Species of Santa/Urn (Santalaceae)L

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Vegetative Anatomy of the Hawaiian Species of Santa/Urn (Santalaceae)L Pacific Science (1980), vol. 34, no. 1 © 1981 by The University Press of Hawaii. All rights reserved Vegetative Anatomy of the Hawaiian Species of Santa/urn (Santalaceae)l LANISTEMMERMANN 2 ABSTRACT: Wood and foliar anatomy of the Hawaiian representatives of the genus Santalum are described. No consistent differences in wood anatomy between taxa were found; however, significant anatomical differences in foliar anatomy were observed. Characteristics of leaf anatomy that are of taxonomic value are the bottle-shaped adaxial epidermal cells of S. haleakalae, the papillate nature of the abaxial leaf surface in several taxa, and the presence of adaxial as well as abaxial stomata in S. ellipticum. SEVEN SPECIES OF Santalum from Hawai'i bases of commerce for the Kingdom of have been recognized in the past. Hawai'i after European contact. Nomenclature in this paper follows the The taxa of the Hawaiian sandalwoods recent revision of the genus (Stemmermann have been distinguished by previous authors 1980). The sandalwoods are separated into on gross characteristics of the flowers, fruit, two groups generally thought to be derived leaves, and growth habits of the plants, but from separate introductions to Hawai'i until now no complete anatomical study of (Rock 1916, Skottsberg 1927, 1930, Tuyama the wood of the various taxa has been made. 1939, Fosberg 1948). The freycinetianum Though the wood of at least three Hawaiian group (including S. haleakalae, S. freycine­ taxa, probably S. paniculatum var. pilgeri, tianum,3 S. freycinetianum var. lanaiense, S. freycinetianum, and S. freycinetianum var. S. freycinetianum var. auwahiense, and pyrularium, has been examined in previous S. freycinetianum var. pyrularium) is charac­ studies (Brown 1922, Metcalfe 1935), it is terized by having flowers that are usually impossible to be certain which taxa were reddish, with long perigonal tubes. The ellip­ actually investigated due to confusion of the ticum group (including S. ellipticum, S. ellip­ original diagnoses of S. freycinetianum and ticum var. littorale, S. paniculatum,3 and S. ellipticum. Furthermore, at least four taxa S. paniculatum var. pilgeri) is characterized were not examined at all. Therefore, since by having flowers that are greenish at an­ the wood of the genus Santalum has been thesis, with short, perigonal tubes. Some, significantly important in the economy of perhaps all, of the species were harvested Hawai'i, and review of the literature demon­ during the sandalwood trade in the nine­ strated a paucity of information on the teenth century, and formed one of the first wood anatomy of the Hawaiian taxa, a study was undertaken to determine whether any differences in the wood exist among the 1 Hawaii Agricultural Experiment Station Journal series no. 2387. This work was supported by McIntyre­ species or sections of the genus Santalum in Stennis funds allocated to the Hawaii Agricultural Hawai'i. Experiment Station to support project 677, and by a In addition, though gross foliar features travel grant from Pacific Tropical Botanical Garden to have been used to distinguish among the the author. This paper represents a portion of a thesis submitted to the Graduate Division of the University of Hawaiian taxa of Santalum (Skottsberg Hawaii in partial fulfillment of the requirements for the 1927, Degener 1940), no anatomical descrip­ M.S. degree. Manuscript accepted 10 January 1980. tions or comparisons of the leaves of the 2 Present address: University of Hawaii, Department taxa had been made. This study therefore of Botany, Honolulu, Hawaii 96822. also examined foliar anatomy of all the taxa 3 For S. freyeil7elial7um, S. e/lipliewn, and S. pal7i­ eu/alum, the typical variety is assumed unless another is to determine the anatomical basis for char­ specifically mentioned. acters of apparent taxonomic significance. 55 56 PACIFIC SCIENCE, Volume 34, January 1980 MATERIALS AND METHODS in Craf III, with aspiration when possible. Leaves to be sectioned were dehydrated in Fresh wood samples were collected from an ethanol tertiary butanol series, infiltrated populations of each of the recognized species with Parowax, finally embedded in Para­ of Santalum. Voucher herbarium specimens plast, and sectioned on a rotary microtome. were prepared for each sample and were Cross sections and paradermal sections were deposited in the herbarium at the University cut through the widest part of the lamina of Hawaii (HAW). Wood samples were taken and perpendicular to the midrib. The sec­ from branches ranging from 5 (rarely 4) to 9 tions were mounted and then stained in cm in diameter, fixed in Craf III, and as­ Johansen's quadruple stain (Johansen 1940), pirated to allow for rapid fixation of the toluidine blue ° (Sakai 1973), or various tissues. After at least a week of fixation the schedules of tannic acid-ferric chloride, samples were transferred to alcohol and sec­ safranin 0, and fast green. tioned on a sliding microtome. Sections were Leaf clearings were prepared by placing stained in tannic acid-ferric chloride­ fixed leaves in the paraffin oven in a solution lacmoid according to Cheadle, Gifford, and of 50 g chloral hydrate, 25 ml liquid phenol, Esau (1953) and mounted in synthetic resin. and 35 mllactic acid, until clearing was com­ Macerations were prepared by placing plete. The cleared leaves were rehydrated slivers of wood in a solution of 1 part 30 and stained with tannic acid-ferric chloride percent hydrogen peroxide, 4 parts water, and safranin °or safranin and celestine blue and 5 parts glacial acetic acid (Gifford et al. (Gray and Pickle 1956), dehydrated, and 1956). Macerating fluid was changed at 3­ mounted in resin. day intervals until the maceration process To determine silica body integrity and was complete, usually after about 5 days. shape, a few macerations of leaves were pre­ Material was then rinsed, stained with a pared using the same solution and staining tannic acid mordant followed by ferric technique used for wood maceration. In chloride and safranin 0, dehydrated in an addition to this, the percent weight of silica ethanol series to xylene, and mounted in bodies in leaves of a few samples was de­ synthetic resin. termined by recording the dry weight of The wood morphology of Santalum ellip­ leaves, ashing them in a muffle furnace in ticum var. littorale was not studied for sev­ porcelain crucibles, reweighing them, and eral reasons. That taxon is on proposed lists recording the ashed weight. The ash was of endangered species for Hawai'i (Fosberg treated with approximately 10 percent con­ and Herbst 1975, U. S. Department of the centrated H 2 S04 , 90 percent concentrated Interior 1976), and as the plant is a low HN03 , and a few drops of 30 percent H 2 0 2 . shrub, with no sizable branches except for The treated ash was then left on a hot plate the central axis of the plant, most of an for a week until the carbon had been re­ entire plant would have had to be taken to moved, and the remaining substance was do an analysis that could be comparable filtered on ashless filter paper, re-ashed, and with the other taxa examined. Since analysis reweighed. of wood morphology of the other taxa Basic statistics for wood morphology were showed no consistent differences of taxo­ based on 20 measurements per sample for nomic significance; because the habit of the soine attributes: extreme vessel body length, plant is such that the wood could not be of fiber length, vessel diameter, ray height, and economic importance; and since it is found vessel distribution. Ten measurements per in semiarid conditions where regrowth is sample were made for the other examined slow, it was felt that any information gath­ wood characteristics and for most foliar ered on its wood morphology would not be characteristics examined. 4 of sufficient interest to warrant the taking of a plant. 4 Refer to M.S. thesis deposited at the University of Leaf samples were collected from the same Hawaii for actual numerical analyses (Stemmermann populations as the wood samples and fixed 1977). FIGURE I. Wood morphology-vessel characteristics of Hawaiian Santalum. A, semi-ring porous arrangement of vessels in S. haleakalae (L. Stemmermann 737) x 29; B, diffuse porous arrangf;ment of vessels in S. haleakalae (L. Stemmermann 750) x 29; C, tailed and transverse to slightly oblique vessel end walls in one sample of S. paniculatum (L. Stemmermann 799) x 177; D, ring porous arrangement of vessels in a sample of S. paniculatum var. pilgeri (Pat Conant s.n.) x 29; E, single-tailed vessel member of S. paniculatwn (L. Stemmermann 799) notice circular structure in the side wall as discussed in the text x 177; F, double-tailed vessel member of S. paniculatum (L. Stemmermann 791 B) with circular structure in side wall x 177; G. tyloses with resinous contents in vessel of S. jreycinetianum (L. Stemmermann 801) x 177. 58 PACIFIC SCIENCE, Volume 34, January 1980 RESULTS on that analysis, the vessels were again found to be short to medium sized, ranging from 141 to 576 These data are in Wood .um. agreement with previous reports for vessel GROSS MORPHOLOGY: The sapwood of all lengths for Santalacean species (Swamy the examined species of Santalurn is yellow­ 1949) and for S. freycinetianurn var. pyru­ ish in color, while the heartwood becomes lariurn (Brown 1922). brown, sometimes with a reddish hue. Only Both radial and tangential vessel dia­ the heartwood is aromatic, and in samples meters were measured, and vessels were taken for this research from branches less found to be round to oval and to range from than 9 cm in diameter, little or no heartwood being very small to small, or rarely extremely had developed. small or moderate sized. Radial diameters The wood of all the examined species of range from 31 to 113 .um, and tangential Santalurn is diffuse porous (Figure IE) or diameters range from 22 to 56 .um.
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