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BARK ANATOMY of SOUTHERN SOUTH AMERICAN CUPRESSACEAE by Alberto Antonio De Magistris1 & María Agueda Castro2

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BARK ANATOMY of SOUTHERN SOUTH AMERICAN CUPRESSACEAE by Alberto Antonio De Magistris1 & María Agueda Castro2 IAWA Journal, Vol. 22 (4), 2001: 367–383 BARK ANATOMY OF SOUTHERN SOUTH AMERICAN CUPRESSACEAE by Alberto Antonio De Magistris1 & María Agueda Castro2 SUMMARY Anatomical and macroscopic features of the bark are described for the Cupressaceae indigenous to southern South America: Austrocedrus chilensis, Fitzroya cupressoides and Pilgerodendron uviferum. Macro- scopically there are two different types: 1) with rather deep longitudi- nal and transverse fissures, and short ridges or scales varying in adher- ence (Austrocedrus); and 2) with deep fissures, and elongate and wide ridges, which contain the rhytidome accumulated over several years (Fitzroya and Pilgerodendron). The pattern of the secondary phloem of the three species studied agrees closely with that of the other genera of Cupressaceae, but there are diagnostic differences as well: arrangement of the fibre rows, fibre types, frequency of resin ducts and the ray height. Pilgerodendron uviferum shows rays with erect marginal cells, similar to Strasburger cells, whereas ray sclereids in the fibre crossing zones were only observed in Austrocedrus. Resin ducts are present in Austrocedrus and Fitzroya. The main characteristics are used for an identification key. Anatomical relations with other genera of the family are discussed. Key words: Austrocedrus chilensis, Fitzroya cupressoides, Pilgeroden- dron uviferum, Cupressaceae, bark anatomy, secondary phloem, South America. INTRODUCTION The Cupressaceae are represented in South America by three monospecific genera: Austrocedrus chilensis (D. Don) Florin & Boutelje, Fitzroya cupressoides (Mol.) I.M. Johnston and Pilgerodendron uviferum Florin. These species occur in different eco- logical conditions. Austrocedrus chilensis is adapted to dry rocky sites. It is widely distributed in Argentina and Chile and it is a significant source of construction wood. The two other species grow in wet sites. Fitzroya, more frequent in Chile, is today only found in protected areas. Due to its valuable wood, exploitation was carried out close to extinction. The bark of F. cupressoides was also utilised by shipwrights (Alerceʼs oakum). Pilgerodendron uviferum, a typical peat bog species, is the southern- most conifer of the world (Covas 1938). 1) Cátedra de Botánica, Facultad de Ciencias Agrarias, Universidad Nacional de Lomas de Zamora, Ruta 4 Km 2 (1836) Llavallol, Argentina. 2) Laboratorio de Anatomía Vegetal, Facultad de Ciencias Excatas y Naturales, Universidad de Buenos Aires, Pabellón 2, 4° piso, Ciudad Universitaria (1428) Buenos Aires, Argentina [E-mail: [email protected]] – Corresponding author. Downloaded from Brill.com10/07/2021 07:42:43AM via free access 368 IAWA Journal, Vol. 22 (4), 2001 De Magistris & Castro — Bark anatomy of South American Cupressaceae 369 The wood anatomy of these species has been studied in varying detail (Peirce 1937; Greguss 1955; Tortorelli 1956; Díaz-Vaz 1983, 1985a, b; Guerra et al. 1994). In partic- ular, Roig (1992) described the wood ultrastructure of these species and carried out dendro-ecological comparisons. However, the bark anatomy has not been described so far. Studies on bark structure and ultrastructure in Cupressaceae have been con- ducted in various genera indigenous to North America, Asia and Oceania (Chang 1954; Bamber 1959; Liphschitz et al. 1981; Chan 1985; Lev-Yadun & Liphschitz 1989; Yamanaka 1989). Furthermore, various bark and phloem features were analysed in other conifer families (Chan 1986; Nunes et al. 1996; Franceschi et al. 1998). This study presents a detailed account of the bark anatomy of these South American taxa. MATERIALS AND METHODS Fresh bark samples including the vascular cambium zone were obtained from mature trees at breast height, and immediately fixed in FAA in Nahuel Huapi and Los Alerces National Parks (Patagonia, Argentina). Foliage and FAA-fixed material are deposited in the Gaspar Xuarez Herbarium (BAA), at Buenos Aires University (see listing at each bark description). Bark portions about 8–10 mm in width were embedded in polyethyleneglycol 1540 (PEG) to obtain transverse and longitudinal sections of approximately 10 μm thick with a sliding microtome, following the technique described by Rupp (1964), modified by Richter (1981), and adapted by De Magistris et al. (2000). The sections were stained with tannic acid and resorcin blue (Cheadle et al. 1953), dehydrated, and mounted with syn- thetic medium. Additional sections without staining were mounted in glycerine 50% in order to observe the presence and distribution of tannic-resinous substances as well as the starch grains distribution with Lugol. Material, macerated according to the method de- scribed by Boodle (1916), was stained with aqueous safranin, and mounted in glycerin- jelly. Data, including shape, length and height of the phloem elements, are based on 30 or more measurements per sample. Polarised light was used for observation of secondary walls and calcium oxalate crystals. For SEM study, sections, 25–30 μm thick, were ob- tained with a sliding microtome from PEG-embedded samples (from which the embed- ding medium was removed in water), dehydrated, coated with gold, and examined in a JEOL JSM 25SII microscope. For macromorphological descriptions, measurements of fissure length and depth, rhytidome ridges width and total bark thickness were made in the field, on mature trees, over 50 years old. Nomenclature for microscopic and macro- scopic features follows Trockenbrodt (1990) and Junikka (1994), respectively. DESCRIPTIONS Austrocedrus chilensis (D. Don) Florin & Boutelje. – Common name: Ciprés de la Cordillera (Fig. 1A). Material studied: De Magistris 22, VI-17-98, Lago Guillelmo, near Huinca-Ruca, Nahuel Huapi National Park, Río Negro province; De Magistris 30, IV-25-98, between El Mirador and Huincalú, Traful, Nahuel Huapi National Park, Neuquén province; De Magistris 33, IV- 25-98, near Coa-Co, Traful, Nahuel Huapi National Park, Neuquén province; De Magistris 20, I-08-98, Lago Futalaufquen, Los Alerces National Park, Chubut province, all in Ar- gentina. Downloaded from Brill.com10/07/2021 07:42:43AM via free access 368 IAWA Journal, Vol. 22 (4), 2001 De Magistris & Castro — Bark anatomy of South American Cupressaceae 369 Macroscopic features Bark fissured. Fissures, 7.3 (3–12) mm deep, longitudinal and transverse; the lon- gitudinal: elongated, vertical to slightly oblique, compound, usually spiralled in their entirety; transverse fissures when present are short. In general, ridges, 50–150 mm in length, 21.9 (14–31) mm in width, with rounded surface and more or less sharp ends, usually interlaced (Fig. 1B), sometimes rectangular adherent scales with flattened to hollow surface. Bark thickness 8–25 mm. Consistency very firm and fibrous. Rhyti- dome with imbricated layers. Pale brown to greyish brown externally. With pleasant resinous scent. Microscopic features The phloem consists of sieve cells, axial and ray parenchyma, and fibres arranged in regular tangential rows. Each row of axial parenchyma cells always has a row of sieve cells on either side. These composite rows (sieve cell–parenchyma cell–sieve cell) are separated by a row of fibres (Fig. 2C). This regular pattern is maintained in the conducting phloem, but is gradually lost towards the non-conducting phloem. The latter is characterised by an increase of ray and axial parenchyma cell size in TS, which causes the collapse of entire tissue files (Fig. 2A, B). Secondary phloem up to the last-formed periderm 2 mm in depth. Sieve cells square to rectangular with rounded corners in TS, rather radially com- pressed by axial parenchyma cells, in the conducting phloem (Fig. 2C). In RLS, sieve cells are 1453 (658–1954) μm long (measured in maceration), and have blunt ends. Abundant sieve areas are present along the radial walls, distributed in a single file pattern. Axial parenchyma cells circular to subcircular in TS (Fig. 2C), with variable length and diameter in RLS (130 μm long and 54 μm wide, in average), elongated in the con- ducting phloem (232 μm long) to barrel-like in the non-conducting phloem (70 μm long). Elongated nuclei and flattened to rather convex extremes. Phloem rays in TS, straight in the conducting phloem, sinuous and tangentially ex- panded towards non-conducting phloem. In RLS, homocellular, with procumbent ray parenchyma cells, 63 μm in length and 16–21 μm in height, with distinct nuclei. In TLS, rays with elliptic-oblong cells, uniseriate to partially biseriate, 1–35 cells in height. Towards the non-conducting phloem, uniseriate rays turn bi- and triseriate with dilated cells. Ray sclereids with small lumina almost always present in contact with fibres. Strasburger cells absent. Fibres in TS radially flattened, arranged in continuous tangential rows, square to rectangular in section, of two types: 1) lignified thick-walled fibres, with small lu- mina, and 2) non-lignified, thin-walled immature fibres, notably collapsed. Each thick- walled fibre row is separated by 4 (3–5) thin-walled immature fibre rows. Occasion- ally, two consecutive fibre rows present thick walls (Fig. 2C). The fibre wall shows abundant weakly bordered pits with an extended and oblique inner aperture. Fibres 2236 (874–3213) μm in length, 27.5 (16.2–43.2) μm in width. Tips pointed, some- times with small protuberances. Wall 8.1 μm thick, in average. The fibre rows be- come disorganised and discontinuous in the non-conducting phloem. Downloaded from Brill.com10/07/2021 07:42:43AM via free access 370 IAWA Journal, Vol. 22 (4), 2001 De Magistris & Castro
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