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Wood Anatomy of Seven Species of Tachigali (Caesalpinioideae–Leguminosae)

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Wood Anatomy of Seven Species of Tachigali (Caesalpinioideae–Leguminosae) MacedoIAWA et Journalal. – Wood 35 (1),anatomy 2014: of 19–30 Tachigali 19 WOOD ANATOMY OF SEVEN SPECIES OF TACHIGALI (CAESALPINIOIDEAE–LEGUMINOSAE) Tahysa M. Macedo 1, Claudia F. Barros 2,*, Haroldo C. Lima2 and Cecília G. Costa1, 2 1Programa de Pós-graduação Ciências Biológicas (Botânica), Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s/n, São Cristóvão, 20940-040 Rio de Janeiro-RJ, Brazil 2Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Diretoria de Pesquisa Científica, Rua Pacheco Leão 915, 22460-030 Rio de Janeiro-RJ, Brazil *Corresponding author; e-mail: [email protected] absTracT This article describes the wood anatomy of seven species of Tachigali Aublet with the aim of identifying 1) diagnostic characters at the species level and 2) anatomical features with potential for future combined morphological and molecular phylogenetic analysis. Tachigali species present fibre dimorphism and can be grouped according to the arrangement of the thin-walled fibres: tangential bands of thin-walled fibres alternating with thick-walled fibres, as in T. duckei and T. vulgaris; wavy bands , as in T. paratyensis, T. glauca and T. vulgaris; well-developed bands to describe the abundance of thin-walled fibres in contrast to thick-walled fibres, as inT. denudata and T. pilgeriana; and in islands or groups of thin-walled fibres scattered among ordinary fibres. It is recommended to explore the phylogenetic significance of the different types of fibre dimorphism in future combined molecular and morphological cladistics analyses. Keywords: Wood anatomy, systematics, Sclerolobium, Tachigali, Fabaceae. INTRODUCTION Tachigali Aublet includes about 60–70 species and is Neotropical in distribution, extending from Costa Rica to southern Brazil and Paraguay. The greatest number of species occurs in South America (Lewis et al. 2005) with about 70% of the species in Brazil’s Cerrado, Atlantic Rain Forest, and the Amazonian Rain Forest (Silva 2007). These species consist of small to large trees having alternate paripinnate leaves, terminal or lateral racemes or panicules, and fruit strongly laterally compressed and dry (Silva 2007; van der Werff 2008). These hardwood trees are known in Brazil as taxi (Camargos et al. 2001), and they are mainly used in construction (Zenid 1997), but they are also used to build rustic furniture, bridges and fences (Lorenzi 1992, 1998). The wood has also been used in restoration projects, landscaping (Lorenzi 1992, 1998) and as firewood (Oliveiraet al. 2008). © International Association of Wood Anatomists, 2014 DOI 10.1163/22941932-00000044 Published by Koninklijke Brill NV, Leiden Downloaded from Brill.com10/01/2021 05:26:14AM via free access 20 IAWA Journal 35 (1), 2014 Many authors have highlighted the similarity between Tachigali and Sclerolobium, as they differ only in a few flower characteristics, such as the attachment of pistil to stipe, the pubescence of the perianth, and the pattern of the inflorescence (Tulasne 1844; Baillon 1870; Bentham 1865, 1870; Taubert 1892, 1894; Harms 1903, 1928; Dwyer 1954, 1957). As a result, Sclerolobium is considered a synonym of Tachigali based on the similarity of flower morphology (Zarucchi & Herendeen 1993; Barneby & Heald 2002; Pennington et al. 2004; Lewis 2005; Silva & Lima 2007), as well as pollen (Graham & Barker 1981), wood anatomy (Baretta-Kuipers 1981) and molecular characteristics (Haston et al. 2005; Maia 2008). In a review of the tribe Caesalpinieae, Lewis (2005) places Tachigali in the informal “Tachigali group” that also includes Arapatiella Rizzini & A. Mattos and Jacqueshuberia Ducke, which have a cupular hypanthium and stipitate ovary. Haston et al. (2005) found that this group is well sup- ported by molecular data. The wood anatomy of some species of Tachigali has already been examined. Loureiro et al. (1983) presented macroscopic and microscopic descriptions of ten species from the Amazon. Fedalto et al. (1989) macroscopically and microscopically described T. glauca (= T. myrmecophila). Gasson et al. (2003) examined eight species, giving some diagnostic characteristics for the “Tachigali group” (as “Sclerolobium group”), and Pernía & Melandri (2006) analyzed eight species from Venezuela. The present work aims to describe the wood anatomy of seven Tachigali species in order to identify 1) diagnostic characters at the species level and 2) anatomical features with potential for future combined morphological and molecular phylogenetic analysis. MATERIALS AND METHODS This study was based on 32 samples of seven species of Tachigali (Table 1). Some samples were collected in the State of Rio de Janeiro, and others were provided from the wood collections of the Instituto de Pesquisas Tecnológicas de São Paulo (BCTw) Table 1. Collection sites and RBw of Trachigali species sampled. Species Collection sites Wood collection (RBw) T. denudata Floresta da Tijuca (Rio de Janeiro) 3274, 8763, 8764, 8765 Parque Nacional das Fontes do Ipiranga (São Paulo) 9115 T. duckei Parque Nacional de Itatiaia (Rio de Janeiro) 9056, 9057, 9058, 9059, 9060, 9061 T. glauca Floresta Nacional do Tapajós (Pará) 6658, 9116, 9117 T. paratyensis Arboreto do Jardim Botânico do Rio de Janeiro (Rio de Janeiro) 36, 3003, 5205, 9051 T. pilgeriana Arboretum of Jardim Botânico do Rio de Janeiro (Rio de Janeiro) 9052, 9123 Serra de Paranapiacaba (São Paulo) 9118 T. rugosa Parque Nacional de Itatiaia (Rio de Janeiro) 9053, 9054, 9055 T. vulgaris Paracatu (Minas Gerais) 4996 Santarém (Pará) 374, 2374, 9120, 9121, 9122 Caxias (Maranhão) 6107 Tocantins 9119 Downloaded from Brill.com10/01/2021 05:26:14AM via free access Macedo et al. – Wood anatomy of Tachigali 21 Table 2. Summary of the qualitative, differential anatomical features of the species studied. These features were used in PCA (data not shown) and cluster analysis. T. = Trachigali, A. = Arapatiella. — 0 = absent; 1 = present; min /mean /max. T. denudata T. duckei T. glauca T. paratyensis T. pilgeriana T. rugosa T. vulgaris T. A. trepocarpa Growth rings distinct, marked by a distinct fibre zone 1 0 1 0 1 0 0 0 Growth rings indistinct 1 1 0 0 0 1 0 0 Shape of alternate pits polygonal 1 1 1 1 1 1 1 0 Thin-walled fibres in wavy bands 0 0 1 1 0 0 1 0 Thin-walled fibres in tangential bands 0 1 0 0 0 1 0 0 Thin-walled fibres in islands 0 1 1 1 0 0 0 0 Well-developed thin-walled fibre bands 1 0 0 0 1 0 0 0 Thin- to thick-walled fibres 1 1 1 1 1 0 1 0 Thick-walled fibres 0 0 0 0 0 1 0 1 Scanty paratracheal parenchyma 0 1 0 0 0 1 1 0 Vasicentric parenchyma 1 1 1 1 1 1 1 0 Aliform parenchyma 1 0 1 1 1 0 1 0 Banded parenchyma 0 0 0 0 0 0 0 1 Two cells per parenchyma strand 1 0 0 1 1 0 0 0 3–4 cells per parenchyma strand 1 1 1 1 0 1 1 1 5–8 cells per parenchyma strand 0 1 1 0 1 0 1 0 Biseriate rays 1 1 0 1 0 1 0 1 Heterocellular rays 1 1 1 1 1 1 1 0 Silica bodies 0 0 0 0 0 0 1 0 and Instituto de Pesquisas Jardim Botânico do Rio de Janeiro (RBw) (Table 1). The scientific nomenclature followed the “Lista de Espécies da Flora do Brasil 2012” (Lima 2012) and “W³ Tropicos” (http://www.tropicos.org/). Wood sections were prepared in three planes, and their thickness ranged from 15 to 20 µm. All samples were bleached, stained with safranin and astra blue (Bukatsch 1972), dehydrated (Johansen 1940; Sass 1958) and mounted in synthetic resin. The hardest samples were softened by boiling in water or glycerin 50%. When this method failed, polyethylene glycol (PEG) 1500 was used instead (Rupp 1964). Macerations were prepared with Franklin’s solution (Jane 1956), stained with aqueous safranin 1% and mounted on semi-permanent slides with glycerin 50% (Strasburger 1924). The descriptions, measurements and images were obtained with an Olympus BX50 light microscope with a digital image processing system (Image Pro Plus, version 3.0 for Windows) fitted with a video camera (Media Cybernetics CoolSNAP-Pro). The termi- nology followed the “IAWA List of Microscopic Features for Hardwood Identification” (IAWA Committee 1989). Downloaded from Brill.com10/01/2021 05:26:14AM via free access 22 IAWA Journal 35 (1), 2014 The fibre dimorphism was defined according to Ter Welle and Koek-Noorman (1978, 1981) and Van Vliet (1981). It should be noted that the two articles used different ter- minology: pseudoparenchyma and fibre dimorphism, respectively. The different arrange- ments of thinner and thick-walled fibres was classified based mainly on Ter Welle & Koek-Noorman (1981), although other important papers were examined to understand these features (Ter Welle & Koek-Noorman 1978; Baas & Zweypfenning 1979; Ter Welle & Koek-Noorman 1981; Van Vliet 1981; Van Vliet & Baas 1984; Baas 1986; Archer & Van Wyk 1993; Graham et al. 1993; Mennega 1997; Olson & Carlquist 2001; Graham et al. 2005; Marcon-Ferreira 2008). As a result, in this study, fibre dimorphism is described as: i) tangential bands of thin-walled fibres alternating with bands of thick- walled fibres (Ter Welle & Koek-Noorman 1981; Van Vliet 1981; Archer & Van Wyk 1993; Olson & Carlquist 2001); ii) well-developed bands with thin-walled fibres more abundant than the thicker-walled fibres (Van Vliet 1981); iii) wavy bands of thin-walled fibres (Ter Welle & Koek-Noorman 1981); and iv) islands or groups of thin-walled fibres scattered among ordinary fibres (Ter Welle & Koek-Noorman 1981). To test the validity of the wood anatomical features in separating the species ana- lyzed, unweighted cluster analysis and principal components analysis (Manly 1994) were performed, and Table 2 summarizes the anatomical features used. All the analyses were carried out in vegan package (Oksanen et al.
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