IAWA Journal, Vol. 32 (1), 2011: 97–112

WOOD ANATOMY OF SPECIES CALLED “TAUARI”

Rocío A. Bernal1*, Vera Coradin2, José Camargos2, Cecília Costa3 and José Pissarra1

SUMMARY Woods from an important group of Lecythidaceae species called “tauari” can hardly be identified in the field by their gross and general features. In this study we show that, when properly delimited to the genera , and , wood anatomical characteristics can be used to identify the species known as “tauari”, even though it is not possible to separate all species. In addition to anatomical characters, wood colour is an important character to help distinguish species of Cariniana and Allantoma from species of Couratari. Detailed wood anatomical descrip- tions from “tauari” woods Allantoma, Cariniana and Couratari are given and a table with diagnostic differences is presented. Common characters of this group are axial parenchyma in narrow continuous bands, prismatic crystals in chambered axial parenchyma cells and silica bodies in ray cells. Microscopic features that help in species identification are: fibre pitting (minutely or distinctly bordered), traumatic intercellular canals, average vessel diameter, vessel element length, axial parenchyma strand length, and ray height and width. Key words: Allantoma, Cariniana, Couratari, Lecythidaceae, Tauari, wood anatomy.

INTRODUCTION

Man has always identified and classified woods according to their appearance, proper- ties and uses. Nowadays accurate wood identification depends mainly on microscopic features (Wheeler & Baas 1998), while gross features such as colour, odour, and texture can be quite variable but also provide useful information. “Tauari” (native Tupi word meaning “big forest ”) is a tropical timber very com- monly harvested in the Amazon Basin. It is widely commercialised and exported due to its good physical and mechanical properties and easy workability. However, woods from several species and different origin are grouped under the name “tauari” and com- mercial lots are usually composed of a mixture of species. Indeed, control and super-

1) BioFIG, Departamento de Biologia - Faculdade de Ciências, Universidade do Porto, Edificio FC4, Rua do Campo Alegre, s/n°, 4169-007 Porto, Portugal. 2) Laboratório de Produtos Florestais, Serviço Florestal Brasileiro, SCEN Trecho 2, Ed. Sede do Ibama, Caixa Postal 09870, 708-900 Brasília, DF, . 3) Instituto de Pesquisas Jardim Botânico do , Rua Pacheco Leão 915, Bolsista CNPq, 22460-000 Rio de Janeiro, RJ, Brazil. *) Corresponding author [E-mail: [email protected]].

Downloaded from Brill.com09/24/2021 04:06:53PM via free access 98 IAWA Journal, Vol. 32 (1), 2011 vision offices have been reporting an increasing number of “tauari” woods in the trade (Instituto Brasileiro de Desenvolvimento Florestal 1988) not all being true “tauari” spe- cies. Procópio and Secco (2008) found eight different Lecythidaceae species grouped as “tauari” in two regions of the State of Pará in Brazil. In the Brazilian Amazon the most common method for identifying species in the field is through the use of local knowledge which is based on the comparison of the tree’s vegetative parts (Lacerda & Nimmo 2010). Additionally, a species called by one name in one region might not be the same in another region; moreover, one common name might be used in multiple regions but referring to different species. Therefore, the grouping of similar species under a common name masks the actual population size of the resource species. In forest inventories it may contribute to overestimates of the timber production poten- tial of commercial trees, leading in turn to lack of trust between seller and customer (Procópio & Secco 2008; Lacerda & Nimmo 2010). Identification of “tauari” wood in the field has proved to be very difficult. Macroscopic characters like external bark morphology or presence of growth rings are insufficient for accurate identification to the species level. For the correct identification of a “tauari” timber it is necessary to collect wood samples together with the corresponding herbarium vouchers, as recom- mended by Barker (2008). Unfortunately, this is not happening in most cases. Species called “tauari” are included in the Lecythidaceae, and may belong to three genera: Allantoma, Cariniana and Couratari. According to the first anatomical survey by Diehl (1935), Lecythidaceae are a homogeneous group, characterised by bands of axial parenchyma, exclusively simple perforation plates, fibres with simple to indistinctly bordered pits, alternate intervessel pitting, and two types of vessel ray pitting. Metcalfe and Chalk (1950) in a general description of Lecythidaceae wood, concluded that there is not a clear distinction of the axial parenchyma and it cannot be used as a diagnostic tool. Wood anatomical studies of “tauari” species are scarce and the anatomical character- istics of this group have to be compiled from previous generic wood descriptions. For instance, Richter (1982) compared the wood structure of Couratari and Couroupita, and showed that anatomical characters like axial parenchyma distribution, type and distribution for inorganic inclusions, especially calcium-oxalate crystals, can be used to discriminate between the two genera. Détienne and Jacquet (1983) described Neotropical genera from the Amazon region, and included information on colour, basic density, gross and microscopic features. Generic descriptions of Corythophora, Couratari, Eschweilera, Gustavia and Lecy- this species from were presented by deZeeuw and Mori (1987). The taxonomic relationships of Neotropical Lecythidaceae with a detailed description of the characteristics of secondary xylem were studied by deZeeuw (1990). Lens et al. (2007) focused on phylogenetically informative wood characters and trends to elucidate evolutionary patterns within the family. InsideWood (2002 onwards) gives detailed wood anatomical descriptions of most tree genera of the Lecythidaceae, including Allantoma, Cariniana and Couratari. Crystals in the wood of Lecythidaceae were studied by Chattaway (1956), Richter (1982), Parameswaram and Richter (1984), deZeeuw and Mori (1987), deZeeuw

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(1990) and Lens et al. (2007). The most common type of prismatic crystals occurs in the 2–more-seriate tangential parenchyma bands and is associated with unilateral thick- ening of cell walls embedding the crystal on the side toward the fibre cells, leaving a cavity on the parenchyma side. The second type of crystalliferous strands are part of uniseriate parenchyma bands with fibres contacting both tangential faces of the paren- chyma cells; in these strands the lateral thickening of the walls is more or less uni- form. Another diagnostic wood anatomical character is the presence of silica deposits, mainly as discrete bodies in the ray cells of Allantoma, Cariniana, Couratari, Esch- weilera and Lecythis. (ter Welle 1976; deZeeuw 1990; Lens et al. 2007). Indeed, ac- cording to the literature the pattern of silica bodies in ray cells can vary from a uniform distribution to a concentration in tangential bands in association with axial parenchyma bands (Richter 1982; deZeeuw 1990). Two types of traumatic structure are found in Lecythidaceae, pith flecks and inter- cellular canals. The presence of canals had been recorded in various studies, Diehl (1935), Metcalfe and Chalk (1950) and deZeeuw (1990), who called them “lysigen- ous cavities”. According to the literature there are 18 species called “tauari” within four Neo- tropical genera belonging to the Lecythidaceae: Allantoma (3 spp.), Cariniana (4 spp.),

Table 1. List of species from the family Lecythidaceae called Tauari. The first common name is the most used. Species Common name Allantoma decandra Tauari-cariri, tauari-vermelho, tauari Allantoma integrifolia (2) Jequitibá-do-amazonas, tauari Allantoma lineata Tauari-seru, seru, tauari Cariniana domestica Jequitibá-do-mato-grosso, jequitibá, tauari Cariniana micrantha Jequitibá-rosa, tauari, tauari-vermelho Cariniana pauciramosa (2) Tauari Jequitibá-vermelho, tauari-cachimbo Couratari atrovinosa (2) Tauari Tauari-claro, tauari-branco, tauari Couratari macrosperma Tauari Couratari multiflora Tauari, tauari-branco, tauari-amarelo Couratari oblongifolia Tauari-branco, tauari Couratari oligantha (2) Tauari Couratari stellata Tauari-escuro, tauari Couratari tauari (2) Tauari-murrão, tauari Couratari tenuicarpa Tauari, Tauari do igapó Eschweilera coriacea (1) Matamatá-giboia, matamatá, matamatá-preto, tauari-preto Eschweilera ovata (1) Matamatá-cascudo, matamatá, matamatá-preto, tauarisinho

(1) Not true “Tauari” (2) Non-important commercial woods

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Couratari (9 spp.) and Eschweilera (2 spp.) (Table 1). An examination of the Esch- weilera woods reveals that they do not belong to the “tauari” group because woods from Eschweilera are brownish, heavier than “tauari”, and their vessels are predominantly solitary and filled with tyloses (Coradin, unpublished results). The most appropriate com- mon name for Eschweilera species is “matamatá” (Camargos et al. 2001). From the other sixteen species five are not considered to be important wood sources for trading (Table 1). Therefore, eleven important commercial timber species called “tauari”, belonging to Allantoma, Cariniana and Couratari species, were analysed. For Allantoma and Cariniana we followed the new delimitation proposed by Huang et al. (2008). In this study a detailed description of the wood anatomy of these important “tauari” woods, using light (LM) and scanning electron microscopic (SEM) observa- tions and a table giving diagnostic characters to separate the genera and species groups are presented.

MATERIALS AND METHODS Forty-four wood specimens from mature samples representing 11 species of the genera Allantoma (2/9), Cariniana (3/10), and Couratari (6/25) were examined: Allantoma decandra, Allantoma lineata, Cariniana domestica, Cariniana micrantha, Cariniana rubra, Couratari guianensis, Couratari macrosperma, Couratari multiflora, Couratari oblongifolia, Couratari stellata and Couratari tenuicarpa. These species, forming the “tauari” group, were selected based on the literature (Prance & Mori 1979; IBDF 1981, 1988; Fedalto et al. 1989; Mori & Prance 1990; Lorenzi 1992, 1998; Camargos et al. 2001) and on the professional experience of researchers of the Wood Anatomy section of the Forest Products Laboratory (LPF/SFB) in Brazil. Descriptions, measurements and counting of cellular elements followed the recom- mendations of the IAWA Committee (1989); at least 75 measurements of each struc- tural parameter were taken. The quantitative data of Couratari guianensis and C. stel- lata were inferred from Fedalto et al. (1989). The values reported in the anatomical descriptions correspond to the minimum, maximum and average values. For light microscopy (LM) sections between 20 to 30 µm in thickness were cut using a sliding microtome, double-stained with safranin and astrablue (Bukatsh 1972) and mounted in a synthetic resin. Fibre and vessel measurements were taken from macerated material; maceration slides were prepared according to a modified Jeffrey’s method, stained in water-safranin 1%, and mounted in glycerin-gelatin (Kraus & Arduin 1997). Measurements were done using IMAGE PRO PLUS 4.5 software. Digital images were taken with an Olympus E camera. Samples for scanning electron microscopy (SEM) were prepared as follows (personal communication by F. Lens, with minor modifications): Small wood samples were soaked in hot water (not boiling) for two or three days. Then radial (RLS), tangential (TLS) and transverse (TS) surfaces were cut with a metal knife (profile C), sharpened every 7–8 sample cuts. SEM surfaces were then bleached with household bleach, washed with water (3 times), dehydrated in ethanol (50% - 75% - 96%) and air dried. Finally, they were sputter-coated with a gold layer.

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RESULTS Anatomical descriptions follow in alphabetical order of the genera and a detailed list of quantitative data is presented in Table 2 and the principal qualitative data are shown in Table 3.

Table 2. Summary of the principal quantitative characters of the studied species. Average and standard deviation (±) values are given. DIAM = Tangential diameter of vessel lumina (µm), VEL = Vessel element length (µm), DFC = Diameter of fibre pit-chamber µ( m), FL = Fibre length (µm), RH = Ray height (µm), RW = Ray width (µm), APL = Axial parenchy- ma strand length (µm).

Species DIAM VEL DFC FL RH RW APL

Allantoma lineata 96 ± 23 472 ± 122 4.6 ± 0.8 1177 ± 201 278 ± 146 31 ± 10 498 ±127

Allantoma decandra 152 ± 32 516 ± 117 4.5 ± 0.8 1301 ± 211 264 ± 82 28 ± 5 510 ± 142

Cariniana domestica 223 ± 36 550 ± 111 4.2 ± 1.0 1540 ± 199 414 ± 95 32 ± 7 606 ± 71

Cariniana micrantha 189 ± 35 576 ± 110 3.7 ± 0.8 1652 ± 301 402 ± 90 32 ± 9 575 ± 77

Cariniana rubra 81 ± 20 471 ± 132 3.1 ± 0.4 1405 ± 266 419 ± 123 46 ± 8 470 ± 140

Couratari guianensis 170 ± 31 598 ± 110 3.7 ± 0.5 1552 ± 249 629 ± 266 47 ± 12 644 ± 106

Couratari macrosperma 210 ± 41 651 ± 126 2.8 ± 0.4 1446 ± 228 524 ± 144 52 ± 12 653 ± 111

Couratari multiflora 131 ± 24 543 ± 81 2.6 ± 0.4 1429 ± 316 413 ± 140 35 ± 9 490 ± 96

Couratari oblongifolia 122 ± 14 757 ± 91 3.2 ± 0.3 1369 ± 192 554 ± 177 35 ± 12 729 ± 106

Couratari stellata 220 ± 46 755 ± 122 3.0 ± 0.3 1620 ± 324 548 ± 225 44 ± 9 702 ± 139

Couratari tenuicarpa 211 ± 31 642 ± 134 2.7 ± 0.4 1460 ± 195 478 ± 99 40 ± 9 678 ± 119

Table 3. Selected wood anatomical features of “tauari” species. LF = Fibres with minutely bordered pits, FT = Fibres with distinctly bordered pits, UNI = Uni- seriate rays, BI = Multiseriate rays (2-seriate), MULTI = Multiseriate rays (3–6-seriate), IC = Intercellular canals, + = frequent, – = rare.

Species / Character LF FT UNI BI MULTI IC Allantoma lineata + + – Allantoma decandra + – + Cariniana domestica + – + Cariniana micrantha + + + + Cariniana rubra + – + + Couratari guianensis + – + + Couratari macrosperma + – – + Couratari multiflora + – – + Couratari oblongifolia + – + + Couratari stellata + – – + Couratari tenuicarpa + – – +

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Figure 1. General aspects of the wood structure of Allantoma, transverse and tangential longi- tudinal sections. – A: A. lineata (SEM), solitary vessels and radial multiples of 2–3, axial parenchyma narrowly banded 1 cell wide. – B: A. decandra (SEM), vessel grouping in radial multiples of 2–3, uniseriately banded axial parenchyma. Note that vessels of A. decandra are wider than those of A. lineata. – C: A. lineata, narrow and short uniseriate rays common, note the crystalliferous series of axial parenchyma (arrow). – D: A. decandra, biseriate rays com- mon, note the prismatic crystals in chambered axial parenchyma strand (arrow). – E: A. lineata (SEM), silica grain in ray cell. – F: A. lineata, pith fleck and solitary vessel with thin-walled tyloses (arrow). – G: A. lineata, prismatic crystals in chambered axial parenchyma cells (arrow). – H: A. lineata, fibres with distinctly bordered pits. — Scale bars = 200 µm in Fig. A–D & F; 50 µm in G; 10 µm in E & H.

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Generic descriptions Allantoma Miers Vessels (Fig. 1A, B) diffuse, (2–)5–7(–15)/mm2, 40% solitary, 40% in radial pairs, 20% in multiples of 3–5; rounded in transverse section; (226–)472–516(–730) µm in element length; tangential diameter (54–)96–152(–215) µm; 8–10 µm in wall thickness; exclusively simple perforation plates; intervessel pits circular to polygonal, alternate, occasionally with coalescent apertures in A. lineata, 7–10 µm in diameter; vessel-ray pits of two types: few enlarged simple pits and many smaller, distinctly bordered pits. Tyloses present, thin-walled in A. lineata. — Fibres (Fig. 1H) with distinctly bordered pits, pit borders (3.0–)4.6(–8.3) µm in diameter; pitting more common in radial than in tangential walls; (659–)1177–1301(–1787) µm in length; 25–30 µm in diameter; 14–18 µm in lumina diameter; thin- to thick-walled. — Axial parenchyma (Fig. 1A, B) in continuous tangential lines forming a network with the rays, bands 1–2 cells wide; 5–7 cells per parenchyma strand, (211–)498–510(–851) µm in strand length; crystal- liferous strands present, unilaterally thickened, adjacent to fibres, with prismatic crystals of calcium oxalate in chambered cells (Fig. 1G). Pith flecks sometimes present in A. lineata (Fig. 1F). — Rays (Fig. 1C, D) 7–10 per mm; uni- or biseriate, uniseriate rays abundant in A. lineata, biseriate rays more frequent in A. decandra; homocellular rays consisting exclusively of procumbent cells, or heterocellular rays consisting of procumbent body ray cells of variable length, or a mixture of procumbent and few square body ray cells, and a marginal row of square ray cells; (54–)264–278(–754) µm in height, rays sometimes fused in A. lineata. Silica bodies (Fig. 1E) present in body ray cells, one per cell, infrequent. Material studied: 9 specimens: A. decandra: Brazil, Pará, Procópio L. s.n. (IAN 179510); Brazil, Amazonas, Krukoff B. 8717 (Uw 16219). – A. lineata: Brazil, Pará, Texeira P. 229 (FPBw 1170) & 240 (FPBw 1172); Brazil, Pará, Bento P. 1036 (FPBw 1244); Brazil, Pará Lins A. 187 (MGw 4573); Brazil, Amazonas, Ducke A. s.n. (SJRw 22626) & s.n. (RBw 947); Brazil, Amapá, Maguire B. 51738 (Tw 37426).

Cariniana Casaretto Vessels (Fig. 2A–C) diffuse; (1–)2–5(–11))/mm2; 44–70% solitary; 18–49% in radial pairs, 13–23% in radial multiples of 3–5; rounded in transverse section; (273–) 471–576(–882) µm in element length; tangential diameter greater in C. domestica and C. micrantha (108–)189–223(–331) µm, and smaller in C. rubra (40–)81(–124) µm; 7–11 µm in wall thickness; exclusively simple perforation plates; intervessel pits cir- cular, polygonal (Fig. 5A, B) and rarely flattened inC. rubra, occasionally with coales- cent apertures in C. micrantha and C. rubra, 9–11µm in diameter; two types of vessel- ray pits: few enlarged simple pits and many smaller, distinctly bordered pits. Tyloses (Fig. 2B) present, thin-walled in C. domestica and C. rubra, with sclerotic walls in C. micrantha. — Fibres with distinctly bordered pits, pit borders (2.4–)3.1–4.2(–6.7) µm in diameter; pitting more common in radial walls; gelatinous fibres present in C. rubra; (752–)1405–1652(–2414) µm in length; 27–30 µm in diameter; 13–17 µm in lumina diameter; thin- to thick-walled. — Axial parenchyma (Fig. 2A–C) in bands,

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Figure 3. Traumatic intercellular canals of Cariniana micrantha. – A: Tangential lines of inter- cellular canals in cross section. – B: Traumatic canals in longitudinal view. — Scale bar = 1 cm.

in continuous tangential lines forming a network with the rays, 1–3 cells wide; 5–8 cells per parenchyma strand, (206–)470–606(–908) µm in strand length; crystal- liferous strands present (Fig. 2G, H). Pith flecks observed in C. rubra. — Rays (Fig. 2D–F) 8 per mm, 1–3(–4)-seriate; uniseriate rays frequent in C. micrantha, biseriate rays frequent in all samples, multiseriate 2–3(–4)-seriate, frequent in C. rubra; rays homocellular, consisting exclusively of procumbent cells and heterocellular rays con- sisting of procumbent body cells or a mixture of procumbent cells with few square body cells, and one marginal row of square ray cells; (126–)402–419(–745) µm in height, ray fusion present. Silica bodies present in body ray cells, mostly globular; irregularly shaped in C. domestica (Fig. 2I). Intercellular canals present in all sam- ples of C. micrantha (Fig. 3A, B). Material studied: 10 specimens: C. domestica: Brazil, , Krukoff B. 5597 (MADw 18729); Brazil, Sohst s.n. (RBHw 16898). – C. micrantha: Brazil, , Texeira P. s.n. (FPBw 832) & 301 (FPBw 2082); Brazil, Amazonas, Krukoff B. 8796 (MADw 30787) & 5095 (Tw 34206); Brazil, Amazonas, Ducke A. 71 (RBw 259). – C. rubra: Brazil, Pará, Plowman T. 8621 (MGw 3597); Brazil, Amazonas, unknown collector IAN 3825; Brazil, Roraima, Silva M. 6077 (MGw 4106).

← Figure 2. General aspects of wood structure of Carinian. – A: C. micrantha, vessels in radial multiples of 2, axial parenchyma in narrow bands 1–2 cells wide. – B: C. domestica, solitary ves- sels with tyloses, axial parenchyma in narrow bands 2 cells wide. – C: C. rubra, solitary vessels and radial multiples of 2–3, axial parenchyma narrowly banded 2–3 cells wide, gelatinous fibres present. – D: C. micrantha, narrow uniseriate and biseriate rays, fused rays present (arrow). – E: C. domestica (SEM), biseriate rays. – F: C. rubra, narrow uniseriate rays, biseriate and 3-seriate rays tallest, fused rays present (arrow). – G: C. domestica, prismatic crystals in chambered axial parenchyma strand. – H: C. micrantha (SEM), detail of the prismatic crystal, see integument that envelops the crystal. – I: C. domestica, silica bodies irregularly shaped in ray cells (arrows). — Scale bars = 300 µm in Fig. A–E; 100 µm in F; 50 µm in G.; 4 µm in H; 25 µm in I.

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H

G

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Couratari Aublet Vessels (Fig. 4A–C) diffuse; (1–)2–4(–3)/mm2; 35–68% solitary, 25–36% in radial pairs, 7–32% in radial multiples of 3–5; rounded in transverse section; (345–)543– 757(–1050) µm in element length; tangential diameter greater in C. macrosperma, C. stellata and C. tenuicarpa (113–)210–220(–312) µm, than in C. guianensis, C. multi- flora and C. oblongifolia, (94–)122–170(–234) µm; 5–11 µm in wall thickness; ex- clusively simple perforation plates; intervessel pits circular and polygonal, alternate, coalescent, occasionally with coalescent apertures, 6–10 µm in diameter; two types of vessel-ray pits: few enlarged simple pits and many smaller, distinctly bordered pits common (Fig. 4J). Tyloses present, thin-walled in all studied species. — Fibres with distinctly bordered pits in C. guianensis and C. oblongifolia, pit borders (2.7–)3.2–3.7 (–5.3) µm in diameter; fibres with minutely bordered pits (Fig. 5C, D) in the remaining species, pit borders (1.7–)2.6–3.0(–3.9) µm in diameter; pitting more common in radial walls; gelatinous fibres present inC. multiflora;thick-walled, radially flattened fibres present in latewood of C. oblongifolia and C. tenuicarpa; (728–)1369–1620(–2504) µm in length; 22–30 µm in diameter; 11–16 µm in lumina diameter; thin- to thick-walled. — Axial parenchyma (Fig. 4A–C) in continuous tangential lines forming a network with the rays, narrow bands of 1–2(–3) cells wide, in C. stellata most frequently one cell wide; 4–8 cells per parenchyma strand, (325–)490–729(–972) µm in strand length; crystalliferous strands present, unilaterally thickened, with solitary prismatic crystals of calcium oxalate in chambered cells (Fig. 4G), frequent in all species except in C. stellata. Pith flecks observed inC. multifloraand C. oblongifolia. — Rays 6–8 per mm, 1–4(–6)-seriate; uniseriate rays infrequent, multiseriate rays 2–6-seriate common (Fig. 4D, E); homocellular rays consisting exclusively of procumbent cells, rare in C. macrosperma; heterocellular rays consisting of procumbent body cells or a mixture of procumbent cells with few square body cells (Fig. 4F), a marginal row of square ray cells; (160–)413–629(–1502) µm in height, ray fusion present (Fig. 4D, E) in all species except in C. macrosperma. Silica bodies present in body ray cells, globular, and irregularly shaped in C. macrosperma and C. stellata (Fig. 4H, I).

← Figure 4. General aspects of wood structure of Couratari. – A: C. guianensis, vessels solitary and in radial multiples of 2–4, axial parenchyma narrowly banded 1–2 cells wide. – B: C. multiflora, vessels in short radial multiples of 2–3, note that they are bigger than in C. guianensis; axial parenchyma narrowly banded 2 cells wide. – C: C. stellata, solitary and radial multiple vessels, note that in this species the vessels are wider than in C. guianensis and C. multiflora, axial parenchyma narrowly banded 1 cell wide. – D: C. multiflora, narrow multiseriate rays with some long multiseriate and uniseriate rays and fused rays (arrow). – E: C. stellata, long multiseriate rays with some narrow short multiseriate and uniseriate rays, fused rays present (arrow). – F: C. stellata, multiseriate ray showing procumbent and square body ray cells and marginal row of square ray cells. – G: C. multiflora(SEM), detail of the prismatic crystal, see integument that envelops the crystal. – H: C. stellata (SEM), silica grains in body ray cells, note the irregular form. – I: C. macrosperma, two silica bodies in a body ray cell (arrow). – J: C. oblongifolia, two types of vessel-ray pits in the same ray cell. — Scale bars = 200 µm in Fig. A–E; 50 µm in F, I; 2 µm in G; 6 µm in H; 20 µm in J.

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Figure 5. Pits found in “tauari” species; note that vessel-ray pits are similar to intervessel pits (A–D: TLS). – A: Cariniana domestica, alternate and polygonal intervessel pits. – B: Cariniana domestica, vessel-ray pits distinctly bordered. – C: Couratari tenuicarpa (SEM) & D: Couratari tenuicarpa, minutely bordered pits in libriform fibres. — Scale bar = 25µ m in Fig. A–D.

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Material studied: 25 specimens: C. guianensis: Brazil, Barbosa M. 171 (FPBw 480); Brazil, Roraima, Silva N. 5152 (MGw 5221); , Polak 115 (MADw 47019). – C. macrosperma: Brazil, Pará, Barbosa M. 150 (FPBw 493); Brazil, Roraima, Rosa N. 483 (IAN 1764); Brazil, Acre, Krukoff B. 5687 (MADw 18758) & (Amazonas) 6451 (MERw 2158); Brazil, Rondônia, Ferreira C. 7431 (Tw 50236). – C. multiflora: Brazil, Maranhão, Frões R. 34473 (IAN 1765); Brazil, Pará, Maciel U. 633 (MGw 3257); Brazil, Roraima, Silva M. 6196 (MGw 3813); Surinam, Stahel 44 (MADw 19572); , Karstedt 34 (RBHw 16837); Brazil, Amazonas, INPA 148451 (MERw 4862). – C. oblongifolia: Brazil, Pará, Barbosa M. 1022 (FPBw 778) & 237 (RBw 6678); Brazil, Pará, Silva N. 5152 (MGw 5399); Brazil, Black 74-953 (SJRw 45540); Brazil, Pará, Cia. Florestal Monte Dourado, 535 (RBw 7096). – C. stellata: Brazil, Pará, Barbosa M. 238 (FPBw 1132) & 406 (FPBw 352); Surinam, Collector unknown (RBHw 16840); origin unknown, Coelho s.n. (RBw 6838). – C. tenuicarpa: Brazil, , Silva M. 4316 (MGw 2085); Brazil, Amazonas, Krukoff B. 7254 (MADw 30802). DISCUSSION AND CONCLUSION

According to various systematic treatments (Huang et al. 2008; Mori et al. 2007; Morton & Prance 1998) and wood anatomical studies (Lens et al. 2007; deZeeuw 1990) Al- lantoma, Cariniana and Couratari genera are closely related. The distinction between “tauari” group species can usually be done using significant differences in quantitative features in combination with qualitative characters. The principal structural characters of the “tauari” group can be summarised as follows. Vessels – Diffuse, mostly solitary or in radial pairs, multiples of 3 to 5 are present but less frequent, rounded in outline. Vessels are grouped in three categories accord- ing to average tangential diameter of vessel lumina: 1) < 100 µm in Allantoma lineata and Cariniana rubra; 2) 100–200 µm in Allantoma decandra, Cariniana micrantha, Couratari guianensis, Couratari multiflora and Couratari oblongifolia; 3) > 200 µm in Cariniana domestica, Couratari macrosperma, Couratari stellata and Couratari tenuicarpa. Element length all ranged within one IAWA Hardwood List category of 350–800 µm; vessel elements of Couratari are generally longer than in Allantoma and Cariniana. All species show simple perforation plates in end walls. Intervessel pits alternate, circular to polygonal (Fig. 5A), 6–11 µm in diameter, sometimes with coalescent apertures. Vessel-ray pits are of two types: smaller, distinctly bordered and similar in size and shape to intervessel pits (Fig. 5B) or larger simple pits with ex- tremely reduced borders (Fig. 4J). Tyloses present, normally thin-walled (Fig. 2B), but in Cariniana micrantha with sclerotic walls. Fibres are of two types: 1) fibres with minutely bordered pits, with the pit chamber less than 3 µm in diameter present in Couratari macrosperma, C. multiflora, C. stellata and C. tenuicarpa (see Fig. 5C, D); 2) fibres with distinctly bordered pits with chambers over 3 µm in diameter are present in Couratari guianensis, Couratari oblongifolia and all Allantoma and Cariniana species. Allantoma species show distinctly bordered pits, with the largest diameters of the group (Fig. 1H). Fibres thin- to thick-walled. Gelatinous fibres observed inCariniana rubra (Fig. 2C) and Couratari multiflora.

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Axial parenchyma – Reticulate, in narrow bands 1–3 cells wide forming a typical network with rays; Couratari stellata shows narrow bands of 1 cell wide; 4– 8 cells per parenchyma strand. Rays – Uniseriate rays are more common in Allantoma lineata and biseriate rays in Allantoma decandra and Cariniana species. Multiseriate rays are more frequent in Couratari. They can be homocellular or heterocellular. The homocellular rays composed exclusively of procumbent cells. The heterocellular rays have usually procumbent body ray cells and a marginal row of square cells, as in Allantoma lineata; in some cases, in the body of the rays there are procumbent cells or a mixture of procumbent and few square cells, as in Cariniana micrantha and Allantoma decandra. In the “tauari” group, procumbent and square ray cells could be mixed throughout the rays as in Couratari guianensis and, usually, more than one condition may be present in one sample. Pith flecks occur in Allantoma lineata (Fig. 1F), Cariniana rubra, Couratari mul- tifloraand Couratari oblongifolia. Intercellular canals occur in all samples of Cariniana micrantha (Fig. 3). Crystals are abundant in almost all species, but in Couratari stellata they are scarce and small. Silica bodies occur in ray cells of all studied samples, in contrast to deZeeuw (1990) who did not observe them in Allantoma decandra and Couratari guianensis. They are mostly globular with a rough nodular surface (see Fig. 1E & 4H), although oblong silica bodies (cf. ter Welle 1976 and deZeeuw 1990) were observed in Cariniana domestica, Couratari macrosperma and Couratari stellata (see Fig. 2 I). Usually one silica body per cell is present, but in Couratari macrosperma two grains can be observed (Fig. 4 I). The silica bodies were uniformly distributed in the body ray cells.

Microscopic wood characters of Allantoma tauaris as vessel size, fibre length and ray width, relate them more closely with Cariniana tauaris. Furthermore, gross characteris- tics as wood colour are similar for tauaris of both genera; in the Brazilian forest they are called “tauari-vermelho” meaning “red-tauari” (see Table 1). Interestingly, Huang et al. (2008), studying morphological and anatomical floral characters of the generaAllantoma and Cariniana, proposed Cariniana decandra to be named Allantoma decandra. Our results show great similarity between Allantoma decandra (Cariniana decandra) and Allantoma lineata in size of vessel lumina, fibre length and ray width. Couratari species have vessels with a tangential diameter higher than 100 µm, and the wood colour can be yellow to white-yellow, as can be seen in Table 1, where some Couratari species are named “tauari-amarelo” or “tauari-branco” meaning “yellow- tauari” and “white-tauari” in English, respectively. Therefore, these characteristics separate Couratari species from Allantoma and Cariniana. The grouping of several species under a common name results in an overestimation of the volume of this “individual pseudospecies” inventories. Therefore, if low density species are logged based on inaccurately estimated population sizes, a logging company could harvest a species to a level that exceeds its capacity to regenerate. No harvesting plan will succeed in providing human needs while preserving viable populations unless the components of such populations are precisely known (Lacerda & Nimmo 2010). Therefore, botanical identification of the species under management is the first and

Downloaded from Brill.com09/24/2021 04:06:53PM via free access Bernal et al. — Wood anatomy of “Tauari” 111 basic requirement for the conservation of tropical forests. Here we show that regardless of the wood structure being fairly uniform, individual species called “tauari” can be identified and separated by a combination of several characters, such as the occurrence of fibres with distinctly or minutely bordered pits, the presence of intercellular canals (in Cariniana micrantha), differences in ray width (Table 3), and differences in average vessel diameter, vessel element length and axial parenchyma strand length (Table 2). Wood colour is very useful and helps to separate Couratari (white, yellowish or gray- ish) from Allantoma and Cariniana (pink, reddish or brownish).

ACKNOWLEDGEMENTS

We are very grateful to Professor Pieter Baas for critical comments and improvement of the manuscript. Thanks are due to the Directors and Curators of the wood collections in Brasilia (FPBw), Pará (MGw, IAN), Rio de Janeiro (RBw), Merida (MERw), Madison (MADw, SRJw), Tervuren (Tw), Utrecht (Uw) and Hamburg (RBHw) for sending wood samples. Dr. F. Lens supplied us with information on SEM methodology, Dr. Carlos Sá and Daniela Silva assisted us during the SEM observations at the Materials Centre of the University of Porto - CEMUP. Rocío A. Bernal acknowledges the support of a PhD scholarship from the Alβan Program (reference E05D052643BR) and a grant from the Portuguese Foundation for Science and Technology (FCT) (reference SFRH/BD/43847/2008).

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