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). - C: RLS, cupressoid cross-field pits. - D: Intertracheid bordered pits in uniseriate arrangement. - E: TS, trabeculae traversing several tracheids (arrows). - F: Details of trabeculae. - G: TLS, ray cells of two sizes. - H: SEM, interior ofaxial parenchyma cello - I: SEM, aspirated pit membrane with thick torus and thin margo. - J: SEM, globular contents (arrows) in axial parenchyma cello - K: LTS, SEM, ray parenchyma cell containing amorphous substance. On both sides, tracheids with warty 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. - D: Uniseriate intertracheid bordered pits in the radial walls of earlywood 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, warty layer lining the pit chamber of a tracheid bordered pit. - K: SEM, interior view ofaxial parenchyma cell wall. - L: SEM, tracheid with warty layer lining showing half-bordered pit pairs with an adjacent radial parenchyma cell (arrow).

Downloaded from Brill.com10/05/2021 05:02:39PM via free access Roig - Wood anatomy of Cupressaceae 155

Downloaded from Brill.com10/05/2021 05:02:39PM via free access 156 IAWA Bulletin n.s., Vol. 13 (2),1992

Downloaded from Brill.com10/05/2021 05:02:39PM via free access Roig - Wood anatomy of Cupressaceae 157

Downloaded from Brill.com10/05/2021 05:02:39PM via free access 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. 3 I). Bordered J.Lm in tangential diameter and 2401 (1564- pits in earlywood tracheids have a rounded 3443) J.Lm in length (measured in macera­ to slightly radialoriented pit aperture, and tion). Warty layer present in both the inner lentic-ular shape with axial or radial orien­ surface of tracheids 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) J.Lm. 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. 4D). The torus with a con­ tional outline angular, near isodiametric 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 of the end wall. Individ­ Axial parenchyma scarce, diffuse and ual axial parenchyma cells 219 (161-305) llIl1 mainly in the latewood (Fig. 2A), occasion­ in length and 27 (20-33) J.Lm 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 cell 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 isodiametric in tangential poorly developed. Individual axial parenchy­ view. Ray parenchyma without nodular end ma cells 311 (230-395) J.Lm in length and walls and with barely noticeable indentures. 31 (25-35) J.Lm 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 (1-10) 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) lohnston (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 Rio Negro, DLM s.n., FR 234, DLM s.n.; soid, with 2 (1-5) pits per cross-field in the Chubut, FR 229. Chile: Osorno, FR 247; earlywood and 2 (1-3) pits with extended Valdivia, FR 248; Continental Chiloe, FR aperture in the latewood (Fig. 4C). Dark de­ 210; Insular Chiloe, FR 204, 205. posits common in ray parenchyma cells. Ray Non-anlltomicalfeatures: Specific gravity tracheids absent. No crystals observed in the based on air-dry volume 0.49 g/cm3 (Torto­ 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. Features useful in distinguishing the species Anatomicalfeatures: Annual growth rings The wood anatomy of Austrocedrus chi­ distinct (Fig. 4A). Earlywood to latewood lensis, Pilgerodendron uviferum and Fitzroya transition 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).

Downloaded from Brill.com10/05/2021 05:02:39PM via free access Roig - Wood anatomy of Cupressaceae 159

tracheid length (j.un) from the middle to the end part of each growth 3500 ring; in Fitzroya parenchyma is more or less 0 A. chilensis I ro F. aJpressoides abundant and restrieted nearly exclusive1y to • P. UViferum the latewood. In Austrocedrus and Fitzroya 0 c axial parenehyma is absent in some rings, but 2500 c @Co. rf present in others. In Pilgerodendron the axial I 0, clll! I o1§, parenchyma occurs in every growth ring, al­ 0 though the amount varies from year' to year • and from site to site. In all three species the 1500 I axial parenehyma is easily detectable on ac­ @ ...." • • count of its dark etmtents . ..• Ray height is also a useful variable for distinguishing between the species. Ray 500L------~------~--~ 30 40 50 height in Austrocedrus averages 6 cells, 1atitude while ray heights in Fitzroya and Pilgero­ dendron average 3 and 2 eells, respectively Fig. 5. Scatter diagram showing the relation (Fig. 6). Rays as tall (up to 20 eells high) between the ttacheid length value against the as those mentioned by Greguss (1955) and provenance. In Pilgerodendron there is an Peiree (1937) were not found in any of the inverse relation between these two variables. Pilgerodendron sampies used in this study. The broken lines join different tracheid length Peiree included P. uviferum in Libocedrus values obtained from different trees growing (as Libocedrus uvifera) and aeeording to his at the same collection site. description of Libocedrus, ray height ranges from 1 to 24 eells. The ray heights reported by Diaz-Vaz (1983, 1985a, 1985b) for these altitude (masl) species agree with the values obtained in the 1500 present study. I0 A. chilensis The end (vertieal) walls of the ray paren­ 0'0 0 F. rupressoides I • P. uvfferum ehyma eells ean be smooth (Austrocedrus and

c Pilgerodendron) or have a nodular appear­ 1000 ,p 0 anee (Fitzroya). Nodular end walls were also reported in Fitzroya by Greguss (1955), and • • o 0 co c perhaps by Peirce (1937: 49) who referred to 0 0 "ray tangential walls with delieate local thiek­ 500 • 0 00 0 enings." The presenee of these bead-like thiek­ enings in the ray parenchyma end walls is a o useful eharacter for distinguishing the wood .. of Fitzroya. oL-__~ __•~L- •__ -- __----+---~ 500 1000 1500 2000 2500 3000 3500 Smooth end walls in axial parenehyma are eommon in the three speeies, but poorly de­ tracheid length (J.IID) veloped nodules ean oceur in Pilgerodendron Fig. 6. Tracheid length plotted against alti­ and Fitzroya. tude. No relation can be found. Traeheid and axial parenehyma length also differ. In Austrocedrus and Fitzroya traeheid length is greater than 2000 J..LIIl, while in Pil­ gerodendron it is less than 2000 J..LIIl (Figs. 5, Different distribution al patterns ofaxial 7). Axial parenehyma eelliength is greater in parenchyma within the growth ring character­ Fitzroya than in the other two genera. ise each species. In Pilgerodendron axial pa­ Typieally, the three species have uniseriate renchyma is abundant, and seattered through intertraeheary bordered pits, abundant in the entire growth ring; in Austrocedrus paren­ radial walls, but in some wide earlywood chyma is less abundant and oceurs frequently tracheids, the pattern of pits changes from

Downloaded from Brill.com10/05/2021 05:02:39PM via free access 160 IAWA Bulletin n.s., Vol. 13 (2),1992

ray height (nwnber of ceIls) woods do not have a noticeable odour. Fitz­ 10~------~~======~ roya has a reddish heartwood; Austrocedrus a a ~ :: :::~es I and Pilgerodendron have a brownish OT pale 8 I • P. uviferum brownish heartwood.

a Ecological trends 6 The very wide natural range of these spe­ a "rIl a eies presents an opportunity to determine if 4 a a there is ecological variation in their wood anatomy. The range of Pilgerodendron and o .:)()oeooOoo• Austrocedrus extends over more than 12 de­ 2 •• -:.eo .eo grees latitude. In some woods, xylem ele­ • • ment lengths decrease with increasing latitude o~----~----~----~----~--~ and altitude (Nylinder & Hagglund 1954; 10 20 30 40 50 60 Baas 1973; Van der Graaff & Baas 1974; tangential ray / mm2 Van den Oever et 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 rays/ ited, some preliminary ecological inferences tangential mm2• The broken line separates can be made. Austrocedrus from Pilgerodendron-Firzroya It appears that in Pilgerodendron tracheid data. length decreases as latitude increases, but in Austrocedrus and Fitzroya latitude and tra­ cheid length apparently are not related (Fig. 5). However, there is considerable variability in uniseriate to open-alternate biseriate. The tracheid length between different individuals structure 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 et al. (1960) found signifi­ or indented margin torus in Fitzroya. cant variation in the tracheid length of Pinus Ray tracheids were not observed in any of taeda individuals growing in similar environ­ the sampies 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 sampies are cheids in the three species. In all the three needed. Latitudinal variation in tracheid length species, but most commonly in Pilgeroden­ must also be considered independently of dron, there was 'traumatic tissue' probably variations in tracheid length related to the formed in response to freezing during the cambial age and distance from the pith. The period of active growth. Such tissue contains sampies fOT this study were obtained from the radially oriented tracheids, which might be outermost heartwood of tree sterns of similar confused with ray tracheids. ages. No relationship was found between Quadratic or triangular intercellular spaces altitude and tracheid length (Fig. 6). were observed in transverse sections of nor­ There is an inverse relation between ray mal tissue (Austrocedrus), but such spaces height and number of ray/sq.mm. This re­ are also associated with compression wood. lation is more apparent in Austrocedrus than Compression wood was frequently observed in Pilgerodendron and Fitzroya (Fig. 7). Sim­ in Pilgerodendron whose sterns 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 colour and 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 increasein ray number per surface odour, while Fitzroya and Austrocedrus area.

Downloaded from Brill.com10/05/2021 05:02:39PM via free access Roig - Wood anatomy of Cupressaeeae 161

Aclmowledgements Greguss, P. 1955. Identifieation of living The author is grateful to Dr. E. Aneibor of gymnosperms on the basis of xylotomy. the Fae. de Ciene. Exaetas y Naturales (UBA) Akaderniai Kiado, Budapest. for valuable suggestions and eomments and IAWA Committee on Nomenelature.1964. to the Instituto de Histologfa y Embriologfa Multilingual glossary of terms used in (UNC) for the use of a seanning electron mi­ wood anatomy. Konkordia, Winterthur. croseope. The author would like to extemi" his Kramer, P.R. 1957. Tracheid 1ength varia­ appreciation to Drs. Fritz H. Schweingruber tion in Loblolly pine. Texas For. Servo and Elisabeth Wheeler who supplied valuable Tech. Report 10. eomments during the revision of the manu­ LaMarche, V.C., R.L. Holmes, P.W. Dun­ seript. widdie & L.G. Drew. 1979. Tree-ring This study was supported by grant PID chronologies of the Southern Hernisphere. N° 9780 from CONICET. 2. Chile. Chronology Series V, Labora­ tory of Tree-Ring Research, Tucson, Ari­ zona. References Martinez Miranda, O. 1981. Flora y fitoso­ Baas, P. 1973. The wood anatomie al range ciologfa de un relicto de Pilgerodendron in Ilex (Aquifoliaceae) and its eeologieal uvifera (D. Don) Florin en el fundo San and phylogenetic significanee. Blumea 21: Pablo de Tregua (Valdivia-Chile). Bosque 193-251. 4:3-11. Bernath, E.L. 1953. Coniferous forest trees Moore, D.M. 1983. Flora of Tierra deI 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 Ameriean ehronology. Tree­ fluence of site and tree properties on yield Ring Bull. 45: 37-42. and quality by production of sulphite pulp Covas, G. 1938. Las eonfferas indfgenas de from spruce. Statens Skogsforskningsin­ la Republica Argentina. Rev. Fae. Agr. de stitut 44 (11): 184. La Plata 21: 201-223. Oever, L. van den, P. Baas & M. Zandee. D' Ambrogio de Argueso, A. 1986. Manual de 1980. Quantitative wood anatomy and teenicas en histologfa vegetal. Ed. Hemi­ provenance in the genus Symploeos. In: sferio Sur, Buenos Aires. J. Bauch (ed.), Natural variations of Diaz-Vaz, J.E. 1983. Fitzroya eupressoides. wOod properties: 49-60. Proceedings of Descripci6n anat6mica. Bosque 5: 47-49. IUFRO Working Party, Oxford. Diaz-Vaz, J. E. 1985a. Austrocedrus ehilen­ Peirce, A.S. 1937. Systematie anatomy of sis. Descripci6n anat6miea. Bosque 6: the woods of the Cupressaeeae. Tropical 49-50. Woods 49: 5-21. Diaz-Vaz, J.E. 1985b. Pilgerodendron uvi­ Phillips, E.W.J. 1948. Identification of soft­ fera. Descripci6n anat6rniea. Bosque 6: woods by their rnicroseopic strueture. 123-124. Forest Prod. Res. Bull. 22: 1-56. Florin, R. 1930. Pilgerodendron, eine neue Pisano, E. 1983. The Magellanic tundra com­ Koniferengattung aus Süd-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. Agerter. 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 Srnithsononian Inst. Misc. Coll. 140: 1- phytogeographical study of temperate 294. Chile. Biogeographica 8: 31-4l. Graaff, N.A. van der & P. Baas. 1974. Wood Rodriguez, R., O. Mattei & M. Quesada anatornical variation in relation to latitude (eds.). 1983. Flora arb6rea de Chile. Uni­ and altitude. Blumea 22: 101-121. versidad de Concepci6n.

Downloaded from Brill.com10/05/2021 05:02:39PM via free access 162 IAWA Bulletin n.s., Vol. 13 (2),1992

Roig, F.A., J. Anehorena, O. Dollenz, A.M. Tortorelli, L.A. 1956. Maderas y bosques Faggi & E. Mendez. 1985. Las eomunida­ argentinos. ACME, Buenos Aires. des vegetales de la Transeeta Botaniea de Zobel, B., E. Thorbjornsen & F. Henson. la Patagonia Austral. In: Boeleke, Moore & 1960. Geographie, site and individual tree Roig (eds.), Transecta Botaniea de la Pata­ variation in wood properties of Loblolly gonia Austral: 350-456. Buenos Aires. Pine. Silvae Genetiea 9 (6): 149-158. Sehweingruber, F.H. 1986. Abrupt growth ehanges in eonifers. IAWA Bull. n. s. 7: 277-283.

APPENDIX

Key to southern South American Cupressaceae

1. Axial parenehyma diffuse, or with strong eoneentration in the middle of the ring, low rays (average height = 2 eells); average traeheid length less than 2000 11m; ray parenehyma end walls without nodules; heartwood strongly aromatie ...... Pilgerodendron uviferum 1. Axial parenehyma distributed ehiefly in or near the latewood but also present in the middle of the earlywood, tracheid length greater than 2000 IJ.ID; heartwood without notieeable odour 2 2. Ray parenehyma end walls with nodules; ray height averaging 3 eells; torus with sealloped margin torus; heartwood reddish ...... Fitzroya cupressoides 2. Ray parenehyma end walls without nodules; ray height averaging 6 eells; torus with smooth margin; heartwood brownish or pale brownish ...... Austrocedrus chilensis

Downloaded from Brill.com10/05/2021 05:02:39PM via free access