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Tropical Woods TROPICAL WOODS NUMBER 113 pp. 54-84 OCTOBER, 1960 WOOD ANATOMY OF ASTEREAE (COMPOSITAE) SHERWIN CARLQUIST 54 TROPICAL WOODS 1960 WOOD ANATOMY OF ASTEREAE (COMPOSITAE) SHERWiN ARLQUIST1 Claremont Graduate School, Rancho Santa Aria Botanic Garden, Claremont, California The woody Astereae utilized in this study account for more woody species than have been, or will be, studied in connection with any other single tribe of Compositae. The abundance of wood samples available for Astereae is not the result of a large number of woody genera. Indeed, the Heliantheae or Mutisieae possess larger nuinbers of woody genera. The bulk of Astereae studied here are representa tives of only a few genera: Baccharis, Chrysothamnus, Haplopappus, and Olearia. These large genera, many species of which are woody, have permitted an emphasis on patterns of wood evolution within genera as well as within the tribe as a whole. The Astereae, more than any other tribe, offer a clear picture of the parallelisms in wood evolution which are so abundant in Compositae, and which complicate analysis in terms of systematic relationships. On these accounts, Astereae may be considered a module, albeit with its own special features, of wood evolution in a highly specialized family. Despite the abundance of woody species in Astereae, a remarkable paucity of literature concerned with wood anatomy of members of this tribe exists. Metcalfe and Chalk (1950) refer to a few characteristics in the genera (no species given) Baccharis, Ericameria (treated here under Haplopappus), and Olearia. Webber (1936) offers a very limited amount of data for Chrysothanmus nauseosus ssp. mohavensis and Aster carnosus. In a semi-floristic study, the writer (1958b) offered full descriptions of the woods of Baccharis angustifolia, B. halimifolia, and Haplopappus phyllocephalus var. megacephalus. Because some genera of Astereae have been considered close to, or referable to ‘This research was supported by a grant from the National Science Foundation, NSFG—5428. The writer wishes to express sincerest appreciation for this aid, which is permitting completion of studies on wood anatomy of Compositae and related families. No. 113 TROPICAL WOODS 55 Heliantheae, the writer (1958a) included Eastwoodia elegans and Grindelia stricta ssp. procumbens in his study on wood anatomy of that tribe. Unfortunately the organization of tabular material in that paper appears to group these two species under the heading “Ambrosiinae.” The two species were merely included at the end of the table, and should have been set off by themselves. Textual reference makes it clear that at no time did the writer consider them as mem— hers of Ambrosiinae; the evidence of the present study suggests, in fact, that they are best treated under Astereac. Because tabular data on these two taxa has already been given, they are not included here in table 1. Hardly any material other than the references cited above gives anything but the most scanty data on wood anatomy of Astereae. New World woody Astereae are mostly shrubs character istically found in semi-arid or arid habitats. In this category may be included Baccharis (North and South America). Haplopappus (North and South America), Chrysothamnus (North American deserts), Gutierrezia (southwestern U. S., Mexico, South America), and Lepidosparturn (southern California). Some of these genera (Baccharis, Chrysothamnus, Haplopappus) include herbs as well as woody plants, and some species which must be termed herbs (Aster, Conyza, Grindelia, Heterotheca, and Solidago) have been included in this study because sufficient xylem was available. Conyza canadensis and Heterotheca grandiflora are annuals. Among the Old World genera, Olearia is outstanding in its number of species and in its diversity of habit. Although some species are herbaceous, most are shrubs or trees. Some of the latter are small trees of a definite arboreal woody habit. Although Olearia is best represented in Australia and New Zealand, a few species occur on other South Pacific islands. Tetramoloplum humile is an alpine, mat-like, peren nial herb which occurs on the Hawaiian Islands, as do other species of that genus. Psiadia rotundifolia is one of the several curious extinct or vanishing endemic arboreal Corn— positae of St. Helena Island. Likewise, Remya rnauiensis, one of two species of this endemic Hawaiian genus, is a shrub of great rarity from the rain forests of Maui. 56 TROPICAL WOODS 1960 MATERL4LS AND ACKNOWLEDGMENTS Wood samples from several sources were utilized for this study. The bulk of the material was provided from the Samuel J. Record collection of Yale University through the kindness of Dr. William L. Stern, formerly of that insti— tution. Material of Australian species of Olearia i’as fur nished by Dr. H. D. Ingle of the Division of Forest Products, Commonwealth Scientific and Industrial Research Organi zation, South Melbourne, Australia. Dr. H. R. Orman of the New Zealand Forest Service, Whakarewarewa, New Zea land, sent samples of Olearia species in that region. The sample of Grindelia cuneifolia was provided by the late Mr. Thomas G. Robbins, staff member of the Jepson Flerharium, University of California, Berkeley. The generosity of these individuals is gratefully acknowledged. A number of addi tional samples were collected in the field by the writer, since woody Astereae arc abundant in southern California. Still other samples were derived from specimens cultivated in the Rancho Santa Ana Botanic Garden, or from her barium specimens. In table 1, specimens are identified by (1) collector’s name and number, if such were known, to gether with the herbarium abbreviation according to Lan jouw and Stafleu (1959); and (2) wood collection numbers. cited in accordance with the abbreviations of Stern and Chambers (1959). Nomenclature of species which occur in the California flora follows that of Munz and Keck (1959); this has influenced treatment of other species (e.g., segregate genera of Haplopappus are not recognized). Most of the sections for this study were prepared by Mr. Charles F. Quibell or Mr. Alfred G. Diboll. METHODS Methods of preparation are the same as those described earlier by the writer (1958a). A set of slides of the species studied here has been sent to the Division of Woods, Smith sonian Institution, Washington, D. C. Fig. 1—4.—Fig. 1, 2. Baccbaris angustifolia.—Fig. 1. Transection. Note band of thick-walled fibers, prominent zones of parenchyma around vessels was well as apotracheal parenchyma.—Fig. 2. Tangential section. Note wide zone of parenchyma at right of vessel—Fig. 3, 4. Raccharis rosrnarinifolia.—Fig. 3. Transection. Note zonation of vessels in rings and patches.—Fig. 4. Tangential section, showing storied fibers. All, )< 65. No. 113 TROPICAL WOODS 57 ANATOMICAL DESCRIPTIONS Table 1 contains a summary of characteristics considered to be significant in comparing species of Astereae with each other and with other Compositae. Features difficult to sum marize in chart form, or which occur in a few species only, are discussed under appropriate histological headings below. Explanation of symbols in table 1: cb = coarse bands on vessel walls cg = continuous grooves, i.e., grooves which form extended helices around vessel walls f = libriform fibers fb = fine bands on vessel walls g = grooves interconnecting two or several pits in a helix my = more numerous vessels p = axial parenchyma r = vascular rays tf = thin-walled fibers v = vessel elements Vt = vascular tracheids wv = wider vessels + = present, characteristic — = present to a limited extent 0 = absent VESSELS I)imensions.—A figure for average diameter of vessel elements is given in table 1. .‘Vhere vascular tracheids occur, this figure is naturally very difficult to determine. Measure ments of vessel-element width were made at the widest point, and included the wall of vessels seen in transection. The figure for average vessel diameter illustrates the rela tively small diameter of vessels in most Astereae: species in which the average diameter is less than 50 are in the majority. This contrasts with Cichorieae (Cariquist, 1960), hut not with Helenieae (Cariquist, 1959). If the diameter of the widest vessel is determined (first column, table 1), this figure is found to reflect the average diameter hut also 58 TROPICAL WOODS 1960 Table I. WooD CHARACTERS n AsrEiuAR SPECIES COLLECTION Aster spinosus Benth. Detweiler 36 (F) (Yw-26691) Baccharis angustifolia Michx. Stern& Chambers 250 (Y,RSA) (Yw-51462) B. cassiniaefolia DC. Rimbach 837 (Y) (Yw-34188) B. concava Pers. (Yw-34037) B. glomerulifolia Pers. MacDonald 5267 (Y) (Yw325o4) B. halimifolia L. Stern & Brizicky 263 (Y,RSA) (Yw-5 1101) B. halirnifolia L. Stern & Brizicky 292 (Y,RSA) (Yw-51122) B. halimifolia L. Stern & Brizicky 398 (Y,RSA) (Yw-51199) B. halimifolia L. Stern & Brizicky 424 (Y,RSA) (Yw-5 1219) B. lanceolata H.B.K. (Yw-16940) B. myrsinites (Pers.)Lam. (Yw-7262) B. sicglecta Britt. Wilson 1477 (F) (Yw-50570) B. obtusifolia H.B.K. Rimbach 156 (Y) (Yw-24095) B. patagonica H.&A. (Yw-1770) B. pilularis DC. ssp. Carlquist 492 (RSA) consanguinea (DC.) C. B. Wolf B. Plummerae Gray Wolf 4404 (RSA) B. polyantha H.B.K. (Yw-16910) B. rosmarinifolia H.&A. (Yw-49852) B. sergiloides Gray Carlquist 430 (RSA) B. thesioides H.B.K. Detweiler 32 (Y) (Yw-14757) B. viminea DC. Cariquist 289 (RSA) Chrysothamnus latisquarnaeus (Gray) Derweiler 22 (F) (Yw-26677) Greene C. nauseosus (Pall.)Britt. ssp. Cariquist 436 (RSA) consimilis (Greene)Hall & Clem. C. nauseosus (Pall.) Britt. ssp.
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