IAWA Bulletin n.s., Vol. 13 (2).1992: 151-162

COMPARATIVE WOOD ANATOMY OF SOUTHERN S011I'B AMERICAN CUPRESSACEAE

by

FidelA. Roig Laboratorio de Dendrocronologfa. Centro Regional de Investigaciones Cientfficas y Tecnicas. CONICET C.C. 330. 5500 Mendoza. Argentina I

Summary The wood anatomy is described for the mnments in a cold-temperate heavy rainfall Cupressaceae indigenous to southern South climate (1500-3000 rom/year), while A. chi· America: Austrocedrus chilensis. Pi/geroden­ lensis occurs on dry (500 mm/year rainfall) dron uviferum and Fitzroya cupressoides. rocky sites. They grow slowly. are 10ng­ 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 anatomical 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 pW1JOse of this paper is to provide for these dron. but not ofAustrocedrus 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 samples were collected ern South America by three rnonospecific from the stems of trees more than 200 years genera: Austrocedrus Aorin er Boutleje. Pi/· old and growing in Argentina and Chile (see gerodendron Florin and Fitzroya Hooker. listing at beginning of each wood anatomical Figure 1 shows their geographic distribu­ description). Two specimens are from her­ tion. They and Araucaria araucana (Molina) barium collections. Collection codes are as K. Koch (Araucariaceae) are the most eco­ follows: FR. personal collection of the nomically valuable native conifers in the author; MERL, Herbario Ruiz Leal-lADlZA south of Chile and Argentina. Principally (Instituto 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 Pi/gero­ ples were boiled in water and then sectioned dendron uviferum occur in peat bog envi- (20 11m thick) on a sliding microtome. Sec-

I) Current Address: Eidgenossische Forschungsanstalt fUr Wald, Schnee und Landschaft. CH­ 8903 Birmensdorf, Switzerland. 152 IAWA Bulletin n.s., Vol. 13 (2),1992 Roig - Wood anatomy of Cupressaceae 153 tions were stained with safranin and mounted wood traeheids. Occasionally in cross section in Canada balsam. Macerations were made triangular or quadrangular intercellular spaces according to Boodle's method (0'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 earlywooc3 and latewood cross .cheids 37 (30-49) I.fJJl in tangential diameter field, are based on 25 or more measurements and 2365 (1702-3086) I.fJJl in length (mea­ per sample. 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 se<:ond­ species; the values in brackets represent the ary wall and in the pit chambers (Fig. 2F, 1). total range ofindividual 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 ofone square millimetre. For tracheary bordered pits 16 (13-18) J1m in SEM analysis, unstained sections and split diameter, almost exclusively uniseriate oroc­ specimens from a few heartwood samples casionally irregularly aligned in the wider ear­ were mounted on specimen stubs using elec­ lywood tracheids (Fig. 20). Pit membranes tric conductive paste, coated with gold and with a centrally thickened torus with a more examined at 20 kV in a Siemens Autoscan or less smooth edge (Fig. 2H). Pit apenure ISM-U3. Wood anatomical terms are accord­ rounded in earlywood, lenticular in latewood ing to the Multilingual Glossary of the Com­ with axial or radial orientation. Trabeculae minee on Nomenclature (lAWA Committee occasionally present. 1964) and special growth ring terminology Axial parenchyma scarce and diffuse, lo­ from Glock et ai. (1960). cated mainly from the middle to the end sec­ tion ofeach 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 of axial parenchyma Material studied (12 samples). Argentina: smooth, thick, and often oblique. Individual Rio Negro, FR 222, 223; DLM s.n.; Neu­ axial parenchyma cells 179 (120-254) J1m qu~n, FR 235, 236,237,238,239,240,249. long, 27 (21-34) J1m in tangential diameter. Chile: Bellavista FR 245; El Manzano, FR Dark deposits fill axial parenchyma. 246. Rays uniseriate, rarely biseriate, 6 (2-16) Non-(lIIIJlonUcaijeanues: Specific gravity cells high; 36 (31-38) rays per sq.mm (in based on air-dry volume 0.49 g/cm3 (Tono­ tangential section). Rays with body cells ap­ relli 1956); heanwood colour varies from proximately isodiametric in tangential view yellow to shades of brown. Heanwood and (Fig. 2B). End walls ofray 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) Anatomicaijeatures: 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. Oose oblique shading corresponds to Aus­ Irocedrus chi/ensis, triangles to Fitzroya cupressoides and open oblique shading to Pilgeroden­ dron uvijerum. Between 39° and 43 0 SL the natural ranges ofthe 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 ai. 1983; Moore 1983; Roig el ai. 1985. 154 IAWA Bulletin n.s., Vol. 13 (2),1992

Pi/gerodendron uviferum ( D. Don.) Aorin from earlywood to latewood (Fig. 3A). Some (Fig. 3). wide rings show density fluctuations (bands Material studied (14 samples). Argentina: of varying cell wall thickness and cell size) Rio Negro, FR 114, 224; Chubul, FR 230; within both the earlywood and latewood. Ab­ Santa Cruz, FR 228. Chile: Osomo, FR 241; rupt reductions in ring width (Schweingruber Continental Chiloe, FR 214, 212, 242, 243, 1986) persisting for a variable number of 244; Insular Chiloe, 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 traeheids. False growth rings Non-anatomicalfeanues: Specific gravity present. Earlywood traeheids 31 (20-43) IJ.11l based on air-dry volume 0.50 g/cm3 (Tono­ in tangential diameter and 1439 (909-2034) relli 1956); heartwood colour reddish-brown, IJ.11l in length. Inner walls oftracheids and pit sapwood yellowish. 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 sttong resinous odour. Hean­ Bordered pits 13 (9-15) IUJl in diameter, uni­ wood fluorescence yellowish-green. seriate and nearly exclusively on the radial Analomicalfeanues: Annual growth rings walls (Fig. 3D). Pit membranes with well 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: TI.S, axial parenchyma cell (arrow) containing amorphous substance. - C: RLS, cupressoid cross-field pits. - 0: Uni­ seriate intertr8cheid bordered pits. - E: TS showing intercellular axial spaces (arrows). - F: RLS, SEM, middle lamella and primary wall, secondary wall and a warty layer lining two contiguous tracheids. - G: TI.S, 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. - 1: TS, fluorescentlighl, scattered axial parenchyma cells (ar­ rows). - J: SEM, lower pan of a tracheid, showing pit membranes in aspirated condition. and pits in an alternate pattern. A warty layer covers the lumen side ofthe tracheid.

Fig. 3. A-K: Pi/gerodendron uviferum. - A: TS. annual growth rings with diffuse axial paren­ chyma tissue (black points). - B: TI.S, axial parenchyma cell (arrow). - C: RLS, cupressoid cross-field pits. - 0: Intenracheid bordered pits in uniseriate arrangement. - E: TS. trabeculae traversing several tracheids (arrows). - F: Details of trabeculae. - G: TI.S. ray cells of two sizes. - H: SEM, interior ofaxial parenchyma cell. - I: SEM, aspirated pit membrane with thick torus and thin margo. - J: SEM, globular contents (arrows) in axial parenchyma cell. - K: LTS, SEM. ray parenchyma cell containing amorphous substance. On both sides. tracheids with wany layer lining.

Fig. 4. A-L. Fitzroya cupressoides. - A: TS, annual growth rings with strong concentration of axial parenchyma in the latewood. - B: TLS. axial parenchyma cell (arrow). - C: RLS, cupres­ soid cross-field pits. - 0: Uniseriate intertracheid bordered pits in the radial walls ofearlywood tracheids. - E: Trabeculae (arrow) traversing three earlywood tracheids. - F: TLS, helical fis­ sures in tracheid cell walls, compression wood. - G: RLS. nodules in end walls ofray cells. - H: SEM, aspirated scalloped torus. - I: SEM. tracheid-parenchyma pit membrane seen from the paren­ chyma side. showing excrescences. - J: SEM, wany layer lining the pit chamber of a tracheid bordered pit. - K: SEM. interior view ofaxial parenchyma cell wall. - L: SEM, tracheid with wany layer lining showing half-bordered pit pairs with an adjacent radial parenchyma cell (arrow). Roig - Wood anatomy of Cupressaceae 155 156 IAWA Bulletin n.s., Vol. 13 (2), 1992 Roig - Wood anatomy of Cupressaceae 157 158 IAWA Bulletin n.s., Vol. 13 (2), 1992 from which fibrillar bundles occasionally ex­ common. Earlywood tracheids 39 (28-57) tend across the margo (Fig. 31). Bordered Ilm in tangential diameter and 2401 (1564­ pits in earlywood tracheids have a rounded 3443) Ilm in length (measured in macera­ to slightly radial oriented pit apenure, and tion). Warty layer present in both the inner lentic-ular shape with axial or radial orien­ surface oftracheids and the pit chamber (Fig. tation in latewood ones. Trabeculae present 4L). Tracheid cross-sectional outlines angu­ (Fig. 3E, F). lar. Bordered pit diameter 13 (10-16) Ilm. Axial parenchyma abundant, diffuse or Radial walls with abundant pits disposed in with a strong concentration in the middle of one row or irregularly aligned in earlywood each ring. Axial parenchyma cells' cross sec­ tracheids (Fig. 40). The torus with a con­ tional outline angular, near isodiarnetric in spicuous scalloped edge. Bordered pit aper­ earlywood, radially flattened in latewood. ture rounded or slightly radially oriented Axial parenchyma cells thin-walled; inden­ in earlywood tracheids, and lenticular shape tures and nodules in end walls absent or oc­ with oblique orientation in latewood ones. casionally a solitary poorly-developed nodule Trabeculae present (Fig. 4E). located in the middle ofthe end wall. Individ­ Axial parenchyma scarce. diffuse and ual axial parenchyma cells 219 (161-305) Jl11l mainly in the latewood (Fig. 2A). occasion­ in length and 27 (20-33) Ilm in tangential ally loosely grouped in tangential bands near diameter. Amorphous or spheroidal dark de­ or at the beginning of the latewood. Cross­ posits attached to the inner cen wall. sectional outlines angular, radially flattened. Rays uniseriate, occasionally biseriate, 2 Axial parenchyma cells with thin longitudinal (1-10) cells high; 46 (44-52) rays per sq.mm walls; thick end walls, smooth or occasion­ (in tangential section). Rays with body cells ally with small nodules. indentures absent or elongated or near isodiarnetric in tangential poorly developed Individual axial parenchy­ view. Ray parenchyma without nodular end ma cells 311 (230-395) Ilm in length and walls and with barely noticeable indentures. 31 (25-35) IlID in tangential diameter. Cells Cross-field pits typically cupressoid (Fig. commonly filled by dark-coloured contents. 3C), with 3 (1-6) pits per cross-field in the Rays uniseriate. 3 0-10) cells high. 42 earlywood and 2 (1-4) pits in the latewood. (32-43) rays per sq.mm (in tangential view). Amorphous dark-coloured deposits common Ray having body cells elongated or approxi­ in the ray parenchyma (Fig. 3K). Ray tra­ mately isodiametric in tangential view. End cheids absent. No crystals in the ray cells. walls of ray parenchyma cells characterised by conspicuous nodules, indentures absent Fitzroya cupressoides (Molina) Johnston (Fig. 40). Axial and radial parenchyma with (Fig. 4). numerous simple pits with round or elliptic Material studied (9 samples) Argentina: shape. Cross-field pitting typically cupres­ RCo Negro, DLM S.n.• FR 234, DLM s.n.; soid, with 2 (1-5) pits per cross-field in the Chubut, FR 229. Chile: Osomo, FR 247; earlywood and 2 0-3) pits with extended Valdivia. FR 248; Continental Chiloe. FR apenure in the latewood (Fig. 4C). Dark de­ 210; Insular Chiloe. FR 204. 205. posits common in ray parenchyma cells. Ray Non-anatomicalfeatures: Specific gravity tracheids absenL No crystals observed in the based on air-dry volume 0.49 g/cm3 (Tono­ ray cells. relli 1956); heartwood colour darker (basical­ ly red) than yellow sapwood. Wood without Discussion and conclusions distinct odour. HeartWood fluorescence yel­ lowish-green. FeatlUeS weful in distinguishing the species Anatomicalfeanues: Annual growth rings The wood anatomy ofAwrrocedrw chi­ distinct (Fig. 4A). Earlywood to latewood lensis, Pilgerodendron uviferum and Fitzroya Iransition gradual. but occasionally abrupt. cupressoides seems at first sight to be quite Ring boundaries slightly wavy. The abrupt homogeneous. However. some features are growth changes with similar characteristics useful for separating woods of these three as in Pilgerodendron. False growth rings un- species (see key in the Appendix). Roig - Wood anatOmy of Cupressaceae 159 lnlCheid length aun> from the middle ro the end pan ofeach growth 3500 ring; in Fitzroya parenchyma is more or less abundant Md restricted nearly exclusively to o II II the latewood. m Austrocedrus and Fitzroya axial parenchyma is absent in some rings, but ~ 2500 II @CIo • present in others. In Pilger~ndronthe axial 00 I .th I ll~ parenchyma occurs in every growth ring, al­ " though the amount varies from year to year • and from site to site. In all three species the 1500 ...or axial parenchyma is easily detectable on ac· @ • • • countofits dark contents. ..• Ray height is also a useful variable for sool.- -"-_-J distinguishing between the species. Ray 30 40 50 height in Austrocedrus averages 6 cells. Ialitudc while ray heights in Fitzroya and Pi/gero­ dendron average 3 and 2 cells. respectively Fig. 5. Scatter diagram showing the relation (Fig. 6). Rays as tall (up to 20 cells high) between the tracheid length value against the as those mentioned by Greguss (1955) Md ~mm~.m~~u~~onth~isM Peirce (1937) were not found in any of the inverse relation between these two variables. Pilgerodendron samples used in this study. The broken lines join different tracheid length Peirce included P. uviferum in Libocedrus values obtained from different trees growing (as Ubocedrus uvifera) Md according to his at the same collection sire. description ofUbocedrus. ray height rMges from I to 24 cells. The ray heights reported by Diaz-Vaz (1983. 1985a, 1985b) for these altitude (m~l) species agree with the values obtained in the 1500 present study. !Po're0 F.A.___...... 1 The end (vertical) walls of the ray paren­ • P.""'"""" chyma ~lls can be smooth (Ausrrocedrus Md

II Pilgerodendron) or have a nodular appear­ 1000 II ance (Fitzroya). Nodular end walls were also reported in Fitzroya by Greguss (1955), and .. "" • " II> a perhaps by Peirce (1937: 49) who referred to " " " "ray tangential walls with delicate local thick­ SOO a enings." The presence of these bead-like thick­ enings in the ray parenchyma end walls is a • useful character for distinguishing the wood •• " o L-. • ...e.l.-L-_- --J of Fitzroya. 500 HIOO ISOD 2000 2500 3000 3500 Smooth end walls in axial parenchyma are common in the three species. but poorly de­ uacheid length (IUD) veloped nodules can occur in Pitgerodendron Fig. 6. Tracheid length plotted against alti­ and Fitzroya. tude. No relation can be found. Trachcid and axial parenchyma length also differ. In Austrocedrus and Fituoya tracheid length is greater than 2000 j1ID, while in Pit­ gerodendron it is less than 2000 I.lJll (Figs. 5, Different distributional patterns of axial 7). Axial parenchyma cell length is greater in parenchyma within the growth ring character­ Fitzroya than in the other two genera. ise each species. m Pilgerodendron axial pa­ Typically. the three species have uniseriate renchyma is abundant. Md scattered through intertracheary bordered pits, abundant in the entire growth ring; in Austrocedrus paren­ radial walls, but in some wide earlywood chyma is less abundant and occurs frequently lracheids. the pattern of pits changes from 160 IAWA Bulletin n.s.• Vol 13 (2).1992

ray height (nwnbcr of cells) woods do not have a noticeable odour. Fitz­ 10 r------;::::::=:=:=::::;} roya has a reddish heartWood; Austrocedrus a Q I ~ ~::.":.IO·1 and Pilgerodendron have a brownish or pale 8 • P.lMlonun brownish heartwood.

a Ecological trends 6 The very wide natural range ofthese spe­ CI ~Q cies presents an opponunity to determine if 4 o o there is ecological variation in their wood anatomy. The range of Pilgerodendron and o eoo 0 • a a DO 0 Ausrrocedrus extends over more than 12de­ 2 • .": • .0 • eo grees latitude. In some woods. xylem ele­ • OL.- • ...... ment lengths decrease with increasing latitude and altitude (Nylinder & Hagglund 1954; , 0 20 30 40 50 60 Baas 1973; Van der Graaff & Baas 1974; tangential ray I mm2 Van den Dever er al. 1980). Although the Fig. 7. Scatter diagram showing the relation material used in this study is somewhat lim­ between the mean ray height against raysl ited. some preliminaIy ecological inferences tangential rmn 2• The broken line separateS can be made. Austrocedrus from Pilgerodendron-Fitzroya It appears that in Pilgerodendron traeheid data. length decreases as latitude increases. but in Ausrrocedrus and Firzroya latitude and tra­ cheid length apparently are not related (Fig. 5). However. there is considerable variability in uniseriate to open-alternate biseriate. The traeheid length between different individuals seructure of the bordered pit membrane var­ growing in the same site and under the same ies. from a more or less smooth torus in Pil­ ecological conditions (see Fig. 5). Kramer gerodendron and Austrocedrus to a scalloped (1957) and Zobel er al. (1960) found signifi­ or indented margin torus in Firzroya. cant variation in the tracheid length ofPinus Ray tracheids were not observed in any of taeda individuals growing in similar environ­ the samples studied. This observation agrees mental conditions. Consequently. in order to with Greguss's (1955) descriptions, but not make valid generalisations about latitudinal with Peirce's (1937) who reponed ray tra­ and altitudinal variation. many samples are cheids in the three species. In all the three needed. Latitudinal variation in traeheid length species. but most commonly in Pilgeroden­ must also be considered independently of dron. there was 'traumatic tissue' probably variations in tr8cheid length related to the formed in response to freezing during the cambial age and distance from the pith. The period ofactive growth. Such tissue contains samples for this study were obtained from the radially oriented tracheids. which might be outennost heartwood of tree stems of similar confused with my tracheids. ages. No relationship was found between Quadratic or triangular intercellular spaces altitude and tracheid length (Fig. 6). were observed in aansverse sections of nor­ There is an inverse relation between ray mal tissue (Ausrrocedrus), but such spaces height and number of ray/sq.mm. This re­ are also associated with compression wood. lation is more apparent in Ausrrocedrus than Compression wood was frequently observed in Pilgerodendroll and Firzroya (Fig. 7). Sim­ in Pilgerodendron whose stems are frequent­ ilar relations have been recorded by Van der ly affected by movements when peat bog soils Graaff and Baas (1974). An explanation for freeze and thaw seasonally. this could be the necessity of maintaining a HeartWood colourand odour are also use­ certain proportion of ray parenchyma cells. ful diagnostic features. Freshly cut surfaces and thus balancing a decrease in ray height of Pilgerodendron wood have a pleasant with an increase in ray number per surface odour. while Fitzroya and Ausrrocedrus area. Roig - Wood anatomy of Cupressaceae 161

Acknowledgements Greguss, P. 1955. Identification of living The author is grateful to Dr. E. Ancibor of gymnosperms on the basis of xylotomy. the Fac. de Ciene. Exactas y Naturales (UBA) Akaderniai Kiado, Budapest. for valuable suggestions and comments and IAWA Committee on Nomenclature. 1964. to the Instituto de Histologia y Embriologfa Multilingual glossary of terms used in (UNC) for the use ofa scanning elecuon mi­ wood anatomy. Konkordia, Wintenhur. croscope. The author would like to extend his Kramer, P.R. 1957. Tracheid length varia­ appreciation to Drs. Fritz H. Schweingruber tion in Loblolly pine. Texas For. Servo and Elisabeth Wheeler who supplied valuable Tech. Report 10. comments during the reYision of the manu­ LaMarche, V.C., R.L. Holmes, P.W. Dun­ script. widdie & L.G. Drew. 1979. Tree-ring This study was supponed by gran! PID chronologies of the Southern Hemisphere. N° 9780 from CONICET. 2. Chile. Chronology Series V, Labora­ tory of Tree-Ring Research, Tucson, Ari­ zona. Ref'erences Martinez Miranda, O. 1981. Flora y fitoso­ Baas, P. 1973. The wood anatomical range ciolog!a de un relicto de Pilgerodendron in Hex (Aquifoliaceae) and its ecological uvifera (D. Don) Florin en el fundo San and phylogenetic significance. Blumea 21: Pablo de Tregua (Valdivia-Chile). Bosque 193-251. 4:3-11. Bernath, E.L. 1953. Coniferous forest ttees Moore, O. M. 1983. Flora of Tierra del Fue­ of Chile. Tropical Woods 52: 19-26. go. Cambridge University Press, London, Boninsegna, J. A. & R. L. Holmes. 1985. Missouri. Fitzroya cupressoides yields 1534-year Nylinder, P. & E. Hagglund. 1954. The in­ long South American chronology. Tree­ fluence of she and tree properties on yield Ring Bull. 45: 37-42. and quality by production of sulphite pulp Covas, G. 1938. Las confferas indigenas de from spruce. Statens Skogsforskningsin­ la Republica Argentina. Rev. Fac. Agr. de stitut 44 (ll): 184. La Plata 21: 201-223. Dever, L. van den, P. Baas & M. Zandee. 0'Ambrogio de Argueso, A. 1986. Manual de 1980. Quantitative wood anatomy and tecnicas en histologfa vegetal. Ed. Hemi­ provenance in the genus Symplocos. In: sferio Sur, Buenos Aires. J. Bauch (ed.), Natural variations of Diaz-Vaz, J.E. 1983. Fitzroya cupressoides. wood properties: 49-60. Proceedings of Descripci6n anat6mica. Bosque 5: 47-49. IUFRO Working Party, Oxford. Diaz-Vaz, J.E. 1985a. Austroeedrus chilen­ Peirce, A. S. 1937. Systematic anatomy of sis. Descripci6n anatomica. Bosque 6: the woods of the Cupressaceae. Tropical 49-50. Woods 49: 5-21. Diaz-Vaz, J.E. 1985b. Pilgerodendron uvi­ Phillips. E.W.J. 1948. Identification of soft­ fera. Descripcion anatomica. Bosque 6: woods by their microscopic structure. 123-124. Forest Prod. Res. Bull. 22: 1-56. Florin, R. 1930. Pilgerodendron, eine neue Pisano, E. 1983. The Magellanic tundra com­ Koniferengattung aus Sud-Chile. Svensk plex. In: A. Gore (ed.), Swamp. bog, fen Bot. Tidskrift 24: 133-135. and moor. Regional Studies Chapter 10: Glock, W.S., R.A. Studhalter & S. Agener. 295-329. Elsevier Sc. Pub!.. Amster­ 1960. Classification and multiplicity of dam. growth layers in the branches of trees. Quintanilla. V. 1977. A contribution to the Smithsononian Inst. Misc. ColI. 140: 1­ phytogeographical study of temperate 294. Chile. Biogeographica 8: 31-41. Graaff, N.A. van der & P. Baas. 1974. Wood Rodriguez, R., O. Mattei & M. Quesada anatomical variation in relation to latitude (eds.). 1983. Flora arb6rea de Chile. Uni­ and altitude. Blumea 22: 101-121. versidad de Concepci6n. 162 IAWA Bulletin n.s., Vol. 13 (2), 1992

Roig, F.A., J. Anchorena, O. Dollenz, A.M. Tonorelli, L.A. 1956. Maderas y bosques Faggi & E. Mendez. 1985. Las comunida­ argentinos. ACME, Buenos Aires. des vegeUlles de la Transecta Boranica de Zobel, B., E. Thorbjomsen & F. Henson. Ia Patagonia Austral. In: Boelcke, Moore & 1960. Geographic, site and individual tree Roig (cds.), Transecta Bouinica de 1a Pata­ vllJ'iation in wood properties of Loblolly gonia Austral: 350-456. Buenos Aires. Pine. Silvae Genetica 9 (6): 149-158. Schweingruber, F. H. 1986. Abrupt growth changes in conifers. IAWA Bull. n.s. 7: 277-283.

APPENDIX

Key to southern South American Cupressaceae

1. Axial parenchyma diffuse. or with strong concentration in the middle of the ring, low rays (average height =2 cells); average tracheid length less than 2000 11m; ray parenchyma end walls without nodules; heanwood strongly aromatic Pilgerodendron uvi/erum 1. Axial parenchyma distributed chiefly in or near the latewood but also present in the middle of the earlywood, traeheid length greater than 2000 )1m; heartwood without noticeable odour 2 2. Ray parenchyma end walls with nodules; ray height averaging 3 cells; torus with scalloped margin torus; heartwood reddish ...... Fitzroya cupressoides 2. Ray parenchyma end walls without nodules; ray height averaging 6 cells; torus with smooth margin; heanwood brownish or pale brownish ...... ••. .. Austrocedrus chilensis