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Comparative Wood Anatomy of Southern South American Cupressaceae

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Comparative Wood Anatomy of Southern South American Cupressaceae IAWA Bulletin n.s., Vol. 13 (2),1992: 151-162 COMPARATIVE WOOD ANATOMY OF SOUTHERN SOUTH AMERICAN CUPRESSACEAE by Fidel A. Roig Laboratorio de Dendrocronolog{a, Centro Regional de lnvestigaciones Cientfficas y Tecnicas, CONICET C. C. 330, 5500 Mendoza, Argentina 1 Summary The wood anatomy is described for the ronments in a cold-temperate heavy rainfall Cupressaceae indigenous to southem South climate (1500-3000 mm/year), whileA. chi­ America: Austrocedrus chilensis, Pilgeroden­ lensis occurs on dry (500 mm/year rainfall) dron uviferum and Fitzroya cupressoides. rocky sites. They grow slowly, are long­ The abundance and distributional pattern of lived, and have been used in dendrochrono­ axial parenchyma within each annual ring, logical studies. Fitzroya cupressoides attains height, and the presence or absence of nod­ ages of more than 1500 years (Boninsegna ules in the end walls of ray parenchyma & Holmes 1985), Pilgerodendron uviferum are all useful anatomie al features for distin­ 500-600 years and Austrocedrus chilensis guishing between the three species. Physical about 1000 years (LaMarche et al. 1979). characteristics such as odour and heartwood The wood anatomy of these species has colour also can be used to separate these been studied in varying detail (peirce 1937; species. Axial parenchyma cell length and Phillips 1948; Greguss 1955; Tortorelli tracheid length show considerable interspeci­ 1956; Diaz-Vaz 1983, 1985a, 1985b). The fic variation. Tracheid lengths of Pilgeroden­ purpose of this paper is to provide for these dron, but not of Austrocedrus and Fitzroya, important southern South American species decrease with increasing latitude. more detailed wood anatomical descriptions Key words: Wood anatomy, Cupressaceae, than now available in the literature, and give Austrocedrus, Pilgerodendron, Fitzroya, features useful in distinguishing between Argentina, Chile. them. Introduction Materials and Methods The Cupressaceae are represented in south­ Fresh heartwood sampies were collected ern South America by three monospecific from the sterns of trees more than 200 years genera: Austrocedrus Florin et Boutleje, Pil­ old and growing in Argentina and Chile (see gerodendron Florin and Fitzroya Hooker. listing at beginning of each wood anatomical Figure 1 shows their geographie distribu­ description). Two specimens are from her­ tion. They andAraucaria araucana (Molina) barium collections. Collection codes are as K. Koch (Araucariaceae) are the most eco­ folIows: FR, personal collection of the nomically valuable native conifers in the author; MERL, Herbario Ruiz Leal-IADIZA south of Chile and Argentina. Prlncipally (lnstituto Argentino de Investigaciones de las because of their decay resistance, these Cu­ Zonas Aridas, Mendoza, Argentina); DLM, pressaceae woods are used in building. These increment core of Dendrochronology Lab­ species occur in different ecological con­ CRICYT, Mendoza, Argentina. Wood sam­ ditions. Fitzroya cupressoides and Pilgero­ pIes were boiled in water and then sectioned dendron uviferum occur in peat bog envi- (20 Jlm thick) on a sliding microtome. Sec- 1) Current Address: Eidgenössische Forschungsanstalt für Wald, Schnee und Landschaft, CH- 8903 Birmensdorf, Switzerland. Downloaded from Brill.com10/05/2021 05:02:39PM via free access 152 IAWA Bulletin n.s., Vol. 13 (2),1992 , 24· .... , / - 32" 44· 48" 52' ~ r rra Fuegc I 1,.. __ ,,_, I . , .......B?' ____ 7.!.o· ____ 7_2' ___ sa- __ J 64· PIJgerodendron uviferum Downloaded from Brill.com10/05/2021 05:02:39PM via free access Roig - Wood anatomy of Cupressaceae 153 tions were stained with safranin and mounted wood tracheids. Occasionally in cross section in Canada balsam. Macerations were made triangular or quadrangular intercellular spaces according to Boodle's method (D'Ambrogio occur between tracheids and between trache­ de Argueso 1986) and mounted in glycerin ids and ray parenchyma (Fig. 2E, G). False jelly. Quantitative data, including the number growth rings not observed. Earlywood tra­ of pits per earlywood and latewood cross cheids 37 (30-49) J.I.ffi in tangential diameter field, are based on 25 or more measurements and 2365 (1702-3086) J.I.ffi in length (mea­ per sampie. The numerical values given in sured in maceration). Tracheids with a warty the descriptions are the mean value for each layer lining the inner surface of the second­ species; the values in brackets represent the ary wall and in the pit chambers (Fig. 2F, J). total range of individual measurements. Data Tracheid cross-sectional outlines are usually on ray frequency are based on at least five rounded, sometimes angular (Fig. 2E). Inter­ counts in areas of one square millimetre. For tracheary bordered pits 16 (13-18) Ilm in SEM analysis, unstained sections and split diameter, almost exclusive1y uniseriate or oc­ specimens from a few heartwood sampies casionally irregularly aligned in the wider ear- were mounted on specimen stubs using elec­ 1ywood tracheids (Fig. 2D). Pit membranes tric conductive paste, coated with gold and with a central1y thickened torus with a more examined at 20 kV in a Siemens Autoscan or less Slnooth edge (Fig. 2H). Pit aperture JSM-U3. Wood anatomical terms are accord­ rounded in earlywood, lenticu1ar in 1atewood ing to the Multilingual Glossary of the Com­ with axial or radial orientation. Trabeculae mittee on Nomenclature (lAWA Committee occasionally present 1964) and special growth ring terminology Axial parenchyma scarce and diffuse, 10- from Glock et al. (1960). cated mainly from the middle to the end sec­ tion of each growth ring (Fig. 21). Cells have Descriptions isodiametric cross-sectional outline or are Austrocedrus chilensis (D. Don) Florin et radially flattened with thin walls and rounded Boutelje (Fig. 2). simple pits. End walls ofaxial parenchyma Material studied (12 sampies). Argentina: smooth, thick, and often oblique. Individual RIO Negro, FR 222, 223; DLM s.n.; Neu­ axial parenchyma cells 179 (120-254) Ilm quen, FR 235, 236, 237, 238, 239, 240, 249. long, 27 (21-34) Ilm in tangential diameter. Chile: Bellavista FR 245; EI Manzano, FR Dark deposits fill axial parenchyma. 246. Rays uniseriate, rarely biseriate, 6 (2-16) Non-anatormcaIJeatures: Specific gravity cells high; 36 (31-38) rays per sq.mm (in based on air-dry volume 0.49 g/cm3 (Torto­ tangential section). Rays with body cells ap­ relli 1956); heartwood colour varies from proximately isodiametric in tangential view yellow to shades of brown. Heartwood and (Fig. 2B). End walls of ray parenchyma cells sapwood have similar coloration. Wood with­ thin, without indentures, and only occasion­ out distinctive odour. Heartwood fluores­ ally with small and isolated nodules. Cross­ cence yellowish-green. field pits cupressoid (Fig. 2C), with 2 (1-4) AnatomicalJeatures: Growth rings distinct pits in the earlywood (Fig. 2C), and 2 (1-3) (Fig. 2A). Gradual transition from early­ pits in the latewood. Some ray cells filled with wood to latewood; narrow latewood marked dark-coloured deposits. Rays lack ray tra­ by radial flattening of the last formed late cheids. No crystals observed. Fig. 1. Natural range of the studied Cupressaceae. Close oblique shading corresponds to Aus­ trocedrus chilensis, triangles to Fitzroya cupressoides and open oblique shading to Pilgeroden­ dron uviferum. Between 39° and 43° SL the natural ranges of the three species are superimposed. Areas with x represent the ice-caps. Sources: Florin 1930; Covas 1938; Bernath 1953; Quinta­ nilla 1977; Martinez Miranda 1981; Pisano 1983; Rodriguez et al. 1983; Moore 1983; Roig et al. 1985. Downloaded from Brill.com10/05/2021 05:02:39PM via free access 154 IAWA Bulletin n.s., Vol. 13 (2), 1992 Pilgerodendron uviferum ( D. Don.) Florin from earlywood to latewood (Fig. 3A). Some (Fig. 3). wide rings show density fluctuations (bands Material studied (14 sampies). Argentina: of varying cell wall thickness and ceIl size) Rio Negro, FR 114, 224; Chubut, FR 230; within both the earlywood and latewood. Ab­ Santa Cruz, FR 228. Chile: Osorno, FR 241; rupt reductions in ring width (Schweingruber Continental Chiloe, FR 214, 212, 242, 243, 1986) persisting for a variable number of 244; Insular Chilo€, FR 227, 198; Isla Piazzi, years occur, especially in older trees. These Caleta Ocasi6n, MERL 44163; Ultima Espe­ periods with very narrow rings of 1-2 rows ranza, MERL 44178. of thin walled tracheids. False growth rings Non-anatomical features: Specific gravity present. Earlywood tracheids 31 (20-43) lJlD based on air-dry volume 0.50 g/cm3 (Torto­ in tangential diameter and 1439 (909-2034) relli 1956); heartwood colour reddish-brown, lJlD in length. Inner walls of tracheids and pit sapwood yeIlowish. Thick bands of denser chamber with a fine warty layer. Tracheid and darker compression wood often present. cross-sectional outlines rounded to angular. Wood with strong resinous odour. Heart­ Bordered pits 13 (9-15) lJlD in diameter, uni­ wood fluorescence yeIlowish-green. seriate and nearly exclusively on the radial Anatomical features: Annual growth rings walls (Fig. 3D). Pit membranes with weIl distinct. Gradual to semi-gradual transition developed torus with a relatively smooth edge (text continued on page 158) Legends of Figures 2-4: Fig. 2. A-J. Austrocedrus chilensis. - A: TS, annual growth rings. - B: TLS, axial parenchyma cell (arrow) containing amorphous substance. - C: RLS, cupressoid cross-field pits. - D: Uni­ seriate intertracheid bordered pits. - E: TS showing interceIlular axial spaces (arrows). - F: RLS, SEM, middle lamella and primary wall, secondary wall and a warty layer lining two contiguous tracheids. - G: TLS, SEM, ray parenchyma cell with contents. The arrows show intercellular radial spaces. - H: SEM, an aspirated pit membrane showing a thick torus and margo with thick and radially aligned microfibrils. - I: TS, fluorescent light, scattered axial parenchyma cells (ar­ rows). - J: SEM, lower part of a tracheid, showing pit membranes in aspirated condition, and pits in an alternate pattern. A warty layer covers the lumen side of the tracheid. Fig. 3. A-K: Pilgerodendron uviferum. - A: TS, annual growth rings with diffuse axial paren­ chyma tissue (black points). - B: TLS, axial parenchyma ceIl (arrow).
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