Charcoal anatomy of Brazilian species. I. Anacardiaceae

An Acad Bras Cienc aabc Anais da Academia Brasileira de Ciências An. Acad. Bras. Ciênc. 0001-3765 1678-2690 Academia Brasileira de Ciências RESUMO Estudos antracológicos têm tido avanço significativo nos trópicos nas últimas décadas. As bases teóricas e metodológicas da disciplina estão bem estabelecidas, entretanto, existe ainda uma forte demanda por material comparativo de referência visando melhorar a acurácia nas determinações taxonômicas, tanto em estudos paleoecológicos e paleoetnobotânicos como para contribuir com o controle da produção ilegal de carvão vegetal. Este trabalho apresenta descrições da anatomia do lenho carbonizado de 11 espécies de Anacardiaceae, em seis gêneros nativos do Brasil (Anacardium occidentale, Anacardium parvifolium, Astronium graveolens, Astronium lecointei, Lithrea molleoides, Schinus terebenthifolius, Spondias mombin, Spondias purpurea, Spondias tuberosa, Tapirira guianensis e Tapirira obtusa). Elas são caracterizadas por vasos de arranjo difuso, solitários e múltiplos, algumas vezes com tiloses e espessamentos espiralados; placas de perfuração simples, pontoações intervasculares alternas, pontoações raio-vasculares maiores que as intervasculares; raios heterocelulares, podendo apresentar canais radiais e cristais; fibras septadas com pontoações simples ou aréolas reduzidas. Os resultados obtidos são similares a descrições prévias de anatomia da madeira das mesmas espécies ou gêneros. Ainda assim, a identificação de carvões é muito mais eficiente quando as amostras desconhecidas são comparadas a equivalentes atuais carbonizados, ao invés de lâminas finas. INTRODUCTION Charcoal, frequent in archaeological or geological contexts, is an important proxy to the reconstruction of past vegetation and climate, as well as to palaeoethnobotanical interpretations. Anthracology, the identification of charcoal based on wood anatomical characters, is a discipline primarily developed for palaeovegetation studies related to archaeology, palaeoecology, geology, and others (Chabal 1997, Scheel-Ybert 2004a, Scott and Damblon 2010). It also applies for archaeobotany and for the identification of contemporary unknown charcoal, in which case it can be used for charcoal control. At present, Brazil is the world's largest producer of charcoal, but a great part of this production still comes from illegal logging of native species (FAO 2014). Almost 90% of charcoal's production goes to the iron and steel industry (Brasil 2012) - around 30-35% of the total output of this production comes from native species (IBGE 2010, ABRAF 2013). Frequently, this illegal production is associated with criminal acts against human beings, often including children and workers under slave-like conditions (Carneiro 2008, IOS 2011a, b). Besides, the production of illegal charcoal is responsible for the emission of c. 5 tons of carbon dioxide caused by improper removal of trees and released gases from carbonization (ABRAF 2009). Controlling charcoal production is therefore very important. Up to recently, it was virtually impossible for government agents to verify if charcoal loads corresponded indeed to legal extraction of reforested woods or to illegal exploitation of native vegetation. However, this can be done through anthracological analysis (Gonçalves and Scheel-Ybert 2012). Usually, species from exotic genera of distinctive wood and charcoal anatomy, such as Corymbia and Eucalyptus, are cultivated for charcoal production (Gonçalves et al. 2014). Although some agroforestry managed native species can also be legally used (Friederichs et al. 2015), most native species are still illegally exploited. An official document (DOF) containing information about the charcoal origin and the species used to produce it must accompany each charcoal load. Verifying the exactitude of this information depends on the direct verification of the load by inspection agents (Gonçalves and Scheel-Ybert 2012). Hence the importance of recognizing the charcoal anatomy of native species. In Mediterranean and temperate regions, areas of lower plant diversity, wood anatomy is better known and charcoal identification is usually based on the consultation of atlases (e.g. Greguss 1959, Schweingruber 1990, Vernet et al. 2001). However, despite numerous studies (e.g. Détienne and Jacquet 1983, Mainieri and Chimelo 1989, Barros and Callado 1997, Barros et al. 2001, 2008, Sonsin et al. 2014), wood anatomy of Brazilian species is still incompletely known, especially for extra-Amazonian and non-commercial timber species, and there are heretofore only unpublished studies on charcoal anatomy of extant species (e.g. Gonçalves 2006, 2010, Albuquerque 2012, Pinto 2013). There is, therefore, a strong demand for anthracological studies of tropical species aiming at charcoal identification, both in the academic fields of archaeobotany, palaeoecology and others, and concerning Brazilian governmental institutions such as Serviço Florestal Brasileiro, IBAMA (Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis), Polícia Militar Ambiental, and INEA (Instituto Estatual do Ambiente), due to the economic importance of charcoal commerce and especially environmental conservation. With the purpose of improving the anatomical knowledge on tropical species and assisting microscopic charcoal identification, we investigate the wood anatomy of carbonized samples from species of the rich Brazilian flora. In this paper, we present results for native species from the Anacardiaceae family. This family is widely distributed in the tropical zone, including about 700 species from 70 genera all around the world (Souza and Lorenzi 2008), with few representatives in temperate regions (Gupta and Agarwal 2008). It comprises trees, shrubs, and rarely woody climbers and herbs (Souza and Lorenzi 2008). Many species are of a considerable economic value, producing edible fruits, chemical compounds used in industry (gums, resins, tannins, and dyes) or medicine, and quality timbers of commercial importance (Record and Hess 1943, Paula and Alves 1997). In spite of the impossibility to cover all Brazilian species comprehensively, we expect that this initiative will provide useful information to researchers of different expertise, including forest sciences, environmental conservation, wood technology, wood anatomy, palaeoecology, and palaeoethnobotany. MATERIALS AND METHODS Twenty-three wood samples of 11 species from six genera of Anacardiaceae native to Brazil were described: Anacardium (A. occidentale L., A. parvifolium Ducke), Astronium (A. graveolens Jacq, A. lecointei Ducke), Lithrea (L. molleoides (Vell.) Engl.), Schinus (S. terebenthifolius Raddi), Spondias (S. mombin L., S. purpurea L., S. tuberosa Arruda), and Tapirira (T. guianensis Aubl., T. obtusa (Benth.) J.D.Mitch.). Wood samples were obtained from field sampling and institutional wood collections: Jardim Botânico do Rio de Janeiro, Rio de Janeiro, Brazil (RBw); Instituto de Pesquisas Tecnológicas, São Paulo, Brazil (BCTw); Instituto Florestal, São Paulo, Brazil (SPSFw). Wood samples collected by R. Scheel-Ybert (RS) are vouchered in Herbarium Alberto Castellanos, Rio de Janeiro, Brazil (GUA); duplicates are deposed in RBw and BCTw wood collections. Specialists identified the material collected by M.E. Solari (ME), although unvouchered. In all cases, samples from the trunk or large branches were analyzed, except for two samples of small branches (Astronium graveolens - ME102, ME109). Samples were wrapped in aluminum foil and carbonized in a muffle furnace at 400 ºC during 40 minutes. All samples are deposited in the charcoal collection of the National Museum at Rio de Janeiro (Antracoteca do Museu Nacional - Laboratório de Arqueobotânica e Paisagem, Universidade Federal do Rio de Janeiro, Brazil). The charring temperature chosen is the same as in many scientific studies based on charcoal analyses (e.g. Solari 1993, Prior and Gasson 1993, Scheel-Ybert 2004b, Gonçalves et al. 2012). Charcoal samples were manually split according to the fundamental wood sections (transversal, longitudinal tangential, and longitudinal radial sections) and examined under a reflected light brightfield/darkfield microscope. Descriptions followed the recommended procedures and terminology by the International Association of Wood Anatomist (IAWA Committee 1989), except for intervessel pits size, for which the internal horizontal diameter of pit apertures was measured (Scheel-Ybert and Gonçalves in press). Anthracologists conventioned to measure the pit aperture (but measuring also the pit chamber diameter when possible) because the carbonization process induces wood cell walls homogenization, frequently making pit chambers indistinguishable. The pit aperture, however, is always distinct. For quantitative measurements, arithmetic means and amplitude (minimum and maximum values) are given. Micrographs were taken using a JEOL 6300F SEM at University Montpellier II (UM-II, France), a JEOL 5310 at the Institut of Biophysics of the Federal University of Rio de Janeiro (UFRJ, Brazil), and a Leica S440 SEM at the Center of Mineral Technology (CETEM, Brazil). Some authors argued that charcoal is sufficiently conductive to be suitable for SEM studies without any special coating or preparation (Blankenhorn et al. 1972, Cutter et al. 1980). However, we have obtained better results by previously sputter-coating specimens with gold or platinum. RESULTS Wood anatomical descriptions for each genus follow. The most important anatomical features of the studied species are presented in Table I. TABLE I Number of specimens analyzes and synthesis of the charcoal anatomical features for the studied Anacardiaceae species. +: present; (+): rare; -: absent. Anacardium (L.) Rottboell (Fig. 1 a-c) Figure 1 a-c. Anacardium occidentale (BCTw 12687). Uncoated, IBCCF/UFRJ. - d-f. Astronium lecointei (RBw 6708), platinum-coated, UM-II. - a,d: Transverse section. - b,e: Tangential section - c,f: Radial section. Bars = 100 µm. Material studied: A. occidentale L. (Rio de Janeiro, Brazil - RS 19); A. parvifolium Ducke (Amazonas, Brazil - BCTw 17084). Growth rings: boundaries indistinct or absent. Vessels: wood diffuse-porous; solitary (60%), in multiples of 2 (30%) and of 3 (10%); tangential diameter 175(110-260)µm; 8(4-14) vessels/mm²; tyloses common in A. occidentale; perforation plates simple with oblique or horizontal end walls; intervessel pits alternate, 7(4-9)µm; vessel-ray pits larger than intervessel pits, with much reduced borders to apparently simple, rounded. Axial parenchyma: scanty paratracheal or vasicentric, sometimes confluent, in 2-4-celled strands. Rays: 1-2-seriate in A. occidentale, exclusively 1-seriate in A. parvifolium; 10(9-13) rays/mm; body ray cells procumbent with one row of upright and square marginal cells in A. occidentale and body cells procumbent with variable thickness and 1-3 rows of upright and square marginal cells in A. parvifolium. Fibres: septate with bordered pits and very thin walls. Secretory elements: absent. Mineral inclusions: absent. Astronium Jacq. (Fig. 1 d-f, Fig. 2) Figure 2 Astronium graveolens (RBw 3733). Uncoated, IBCCF/UFRJ. - a: Transverse section. - b: Tangential section - c: Radial section. - d: Radial canal. - e: Septate fibres. Bars = 100 µm (a-c) and 20 µm (d-e). Material studied: A. graveolens Jacq (Espírito Santo, Brazil - BCTw 13545, SPSFw 436; Mato Grosso, Brazil - ME 102, ME 109); A. lecointei Ducke (Pará, Brazil - RBw 6708). Growth rings: boundaries marked by thick-walled latewood fibres in A. graveolens, and indistinct or absent in A. lecointei. Vessels: wood diffuse-porous; solitary (70%), in multiples of 2 (25%) and 3 (5%); in A. graveolens [SPSFw 436] vessels are exclusively solitary; tangential diameter 90(40-130)µm; 15(12-25) vessels/mm²; tyloses present; perforation plates simple with oblique or horizontal end walls; intervessel pits alternate, 5(4-10)µm; vessel-ray pits larger than intervascular pits, with much reduced borders to apparently simple, rounded. Axial parenchyma: scanty paratracheal and rarely vasicentric, in 1-4-celled strands. Rays 1-3-seriate in A. graveolens, 1-, rarely 2-seriate in A. lecointei; 8(7-11) rays/mm; body ray cells procumbent with 1 to 4 rows of upright and square marginal cells. Fibres: septate with simple to bordered pits and very thin-walled. Secretory elements: radial canals with tangential diameter 30(10-70)µm. Mineral inclusions: prismatic crystals in ray cells. Lithrea Miers ex Hook & Arn. (Fig. 3) Figure 3 Lithrea molleoides (USPw 520). Platinum-coated, UM-II (a-b), gold-coated, CETEM (c-e). - a: Transverse section. - b: Tangential section. - c: Radial section. - d: Vessel-ray pits. - e: Spiral thickenings in some vessels. Bars = 100 µm (a-c) and 20 µm (d-e). Material studied: L. molleoides (Vell.) Engl. (Paraná, Brazil - BCTw 8743; São Paulo, Brazil - BCTw 134, RBw 1710). Growth rings: boundaries indistinct or absent. Vessels: wood diffuse-porous; solitary (35%), in multiples of 2 (35%), 3 (15%), and 4-6 (13%); tangential diameter 45(20-80)µm; frequency vary from 40(25-60) [BCTw 134, BCTw 8743] to 95(70-110) [RBw 1710] vessels/mm²; tyloses absent; perforation plates simple and scalariform with few bars, with oblique and horizontal end walls; intervessel pits alternate, 3,5(3-5)µm; vessel-ray pits larger than intervessel pits, with much reduced borders to apparently simple, rounded; helical thickenings present in some vessels. Axial parenchyma: scanty paratracheal and vasicentric, in 1-4-celled strands. Rays: 1-3-seriate; 9(5-13) rays/mm; body ray cells procumbent with 1 to 4 rows of upright and square marginal cells. Fibres: septate with simple to bordered pits and very thin-walled. Secretory elements: absent. Mineral inclusions: prismatic crystals in fibres and ray cells [BCTw 8743, RBw 710]; crystals in radial alignment in chambered cells [BCTw 134, RBw 1710]. Schinus L. (Fig.4 a-d) Figure 4 a-d. Schinus terebinthifolius (SPSFw 479). - e-g. Tapirira obtusa (BCTw 3863). Platinum-coated, UM-II. - a,e: Transverse section. - b,f: Tangential section. - c,g: Radial section. - d: Radial canal. - Bars = 100 µm (a-c, e-g) and 20 µm (d). Material studied: S. terebinthifolius Raddi (Mato Grosso, Brazil - ME 165; Paraná - BCTw 868; Pernambuco, Brazil - BCTw 12733; Rio de Janeiro, Brazil - RS 05, RS 43; São Paulo, Brazil - SPSFw 479). Growth rings: boundaries indistinct or absent. Vessels: wood diffuse-porous; solitary (14%), in multiples of 2 (27%), 3 (19%), 4-8 (40%); tangential diameter 50(20-100)µm; 130(70-230) vessels/mm²; tyloses common in some specimens [SPSFw 479, RS 05]; perforation plates simple with oblique and horizontal end walls; intervessel pits alternate, 3,5(2-6)µm; vessel-ray pits larger than intervessel pits, with much reduced borders to apparently simple, rounded; helical thickenings present. Axial parenchyma: scanty paratracheal, fairly distinct, in 1-3-celled strands. Rays: 1-3-seriate; 10(5-15) rays/mm; body ray cells procumbent with 1 to 2 rows of upright marginal cells. Fibres: septate with simple to slightly bordered pits and very thin walls; helical thickenings sometimes present. Secretory elements: Radial canals with tangential diameter 50(30-100)µm. Mineral inclusions: prismatic crystals in ray cells. Spondias L. (Fig. 5) Figure 5 Spondias mombin (ME 201). Gold-coated, CETEM. - a: Transverse section. - b: Tangential section. - c: Radial section. - d: Prismatic crystals in ray cells. - e: Septate fibres. - f: Radial canal. Bars = 100 µm (a-c) and 20 µm (d-f). Material studied: S. mombin L. (Brazil - BCTw 13527); S. purpurea L. (Mato Grosso, Brazil - ME 179); S. tuberosa Arruda (Bahia, Brazil - RBw 6322, Pernambuco, Brazil - BCTw 18770). Growth rings: boundaries indistinct or absent. Vessels: wood diffuse-porous; solitary (55%), in multiples of 2 (35%), and 3 (10%); tangential diameter 135(60-250)µm; 15(5-30) vessels/mm²; tyloses absent in S. tuberosa, present in the other species; perforation plates simple with oblique and horizontal end walls; intervessel pits alternate, 7(5-10)µm; vessel-ray pits larger than intervessel pits, with much reduced borders to apparently simple, rounded. Axial parenchyma: scanty paratracheal, fairly distinct, in 2-4-celled strands. Rays: 2-3-seriate in S. purpurea, 3-6-seriate in S. tuberosa, and 4-7-seriate in S. mombin; 5(2-8) rays/mm; procumbent body cells with 1-4, rarely up to 6 rows of upright marginal cells. Fibres: septate with simple to slightly bordered pits and very thin-walled. Secretory elements: radial canals with tangential diameter 40(20-80)µm. Mineral inclusions: prismatic crystals in ray cells of S. purpurea and S. tuberosa and in fibres of S. tuberosa. Tapirira Aubl. (Fig. 4 e-g) Material studied: T. obtusa (Benth.) J.D.Mitch. (São Paulo, Brazil - BCTw 3863); T. guianensis Engl. (Alagoas, Brazil - RBw 5992, Santa Catarina, Brazil - SPSFw 300). Growth rings: boundaries marked by thick-walled fibres in late wood in T. obtusa, indistinct or absent in T. guianensis. Vessels: wood diffuse-porous; solitary (50%) and in multiples of 2 (27%), 3 (18%), and 4 (5%); tangential diameter 90(60-140)µm; 20(10-30) vessels/mm²; tyloses usually present; perforation plates simple with oblique or horizontal end walls; intervessel pits alternate, 5(3-7)µm; vessel-ray pits larger than intervessel pits, with much reduced borders to apparently simple, rounded; helical thickenings present. Axial parenchyma: scanty paratracheal, fairly distinct, in 2-3-celled strands. Rays: 1-2(3)-seriate in T. obtusa, 2-3-seriate in T. guianensis; 7(3-10) rays/mm; body ray cells procumbent with 1 to 5 rows of upright marginal cells. Fibres: septate fibres with simple to slightly bordered pits, very thin-walled. Secretory elements: radial canals with tangential diameter 30(10-60)µm. Mineral inclusions: prismatic crystals in ray cells. Even if vessels tangential diameter and frequency may sometimes differ, it is not possible to distinguish the studied species through charcoal anatomy. Genera, however, or groups of species, can be distinguished by considering a combination of features such as radial canals, helical thickenings, rays width, vessels diameter, and vessels frequency, as proposed in the dichotomous key below. Features such as diameter of intervessel pit apertures and rays frequency do not vary significantly, while the presence of tyloses may depend on the part of the stem analyzed, and that of crystals on environmental aspects. Those are not reliable diagnostic characters, but can be useful to ascertain the identification. Dichotomous key Radial canals absent 2 Radial canals present 3 Mean vessel tangential diameter less than 50µm, more than 40 vessels/mm² Lithrea Mean vessel tangential diameter more than 100µm, less than 20 vessels/mm² Anacardium Helical thickenings present 4 Helical thickenings absent 5 Mean vessel tangential diameter less than 50µm, more than 100 vessels/mm² Schinus Mean vessel tangential diameter more than 50µm, less than 40 vessels/mm² Tapirira Rays 1-3-seriate 6 Rays more than 3-seriate Spondias mombin, Spondias tuberosa Rays 1-3-seriate Astronium Rays 2-3-seriate Spondias purpurea DISCUSSION Anatomical description of charcoal samples intends to improve taxonomical identification and to provide better comparative material to charcoal identification in tropical regions. The decision of describing charcoal is due to methodological requirements. Charcoal analysis in reflected-light microscopy is technically easier and much less time-consuming than traditional techniques of wood analysis involving slides preparation. Besides, we highlight the unequivocal advantage of comparing similar materials. Charcoal identification is much more effective when unknown samples are compared to charred extant equivalents, instead of to wood slides. This happens not only because of easiness of observation, but also in reason of minor anatomical changes due to carbonization. As a rule, wood structure is very well preserved after carbonization, but some modifications may occur, such as: homogenization of the cell walls; tangential and radial shrinkage (which may induce small changes in vessels diameter and in the size of axial and radial parenchyma cells); and disappearance or modification in cell contents (Prior and Gasson 1993, Kim and Hanna 2006, Gonçalves et al. 2012). Anatomical characteristics of Brazilian Anacardiaceae charcoal are, as in wood, relatively homogeneous, but the different genera studied can be distinguished through wood anatomy. The present results closely agree with general descriptions of Anacardiaceae wood (Record and Hess 1943, Metcalfe and Chalk 1950) and with generic and specific wood descriptions (e.g. Tortorelli 1956, Mainieri 1958, Kribs 1968, Pérez Olvera et al. 1980, Martijena 1987, Barajas-Morales and Gómez 1989, Fedalto et al. 1989, Mainieri and Chimelo 1989, Terrazas and Wendt 1995, Richter and Dallwitz 2000, Barros et al. 2001, Miller 2007, Gupta and Agarwal 2008). In most cases, qualitative and quantitative data from the carbonized samples are very similar to previously published studies in wood anatomy, e.g. for Anacardium occidentale (Hess 1946, Dechamps 1979, Détienne and Jacquet 1983, Dong and Baas 1993), Astronium graveolens (Détienne and Jacquet 1983, Nardi Berti and Edlmann Abbate 1992), Lithrea spp (Hess 1946, Terrazas 1994), Lithrea molleoides (Pinho 1969), Schinus spp (Hess 1946, Jofre 1988, Terrazas 1994), Schinus terebinthifolius (Tavares 1970), Spondias spp (Williams 1936, Hess 1946, Kryn 1952, Détienne and Jacquet 1983, Dong and Baas 1993, Terrazas 1994, Sosef et al. 1998), Spondias mombin (León 2003), Tapirira spp (Hess 1946, Terrazas 1994), Tapirira guianensis (Williams 1936, Dechamps 1979). This is particularly interesting for, in spite of morphometrical changes due to the carbonization process (Mc Ginnes et al. 1971, Prior and Alvin 1986, Prior and Gasson 1993, Gonçalves et al. 2012), wood anatomical features seem directly comparable between carbonized and non-carbonized samples. In other cases, however, a few features differed from these previous descriptions, especially regarding vessels tangential diameter and frequency - e.g., in Astronium graveolens vessels are 140% smaller and 80% more frequent, and in Astronium lecontei 75% smaller and 60% more frequent than it was reported by Dechamps (1979); in Schinus terebinthifolius vessels are 60% more frequent than reported by Tavares (1970); in Spondias mombin vessels are 10% smaller than reported by León (2003); in Spondias purpurea vessels are 100% smaller and more frequent than reported by Barajas-Morales and Gómez (1989); in Tapirira guianensis vessels are 60% more frequent than reported by Dechamps (1979). Intervessel pits are generally smaller than reported in the literature, but the data reported for non-carbonized wood refer to the diameter of chamber, while in this work we have measured pit apertures. Quantitative variation, particularly lower vessels tangential diameter, and higher vessels frequency, can be explained by the carbonization process, but intraspecific variability and/or ecological factors can also be implied. CONCLUSIONS Anatomical description of charcoal aims to improve taxonomical identification and to provide better means to anthracological studies in the tropics. Descriptions of tropical charcoal anatomy are of utmost importance in different fields of knowledge, such as archaeobotany, palaeoecology, wood technology, and environmental conservation. They are most important as a comparative material for the identification of unknown charcoal pieces from archaeological or sedimentary contexts, which may lead to palaeoenvironmental and palaeoethnobotanical interpretations. The use of anthracology to control illegal charcoal commerce also shows a strong potential in Brazil. There are as yet few studies with charcoal intended to reduce the indiscriminate use of native species, but they are generally focused in Eucalyptus, which is the most frequently species legally used for charcoal production (e.g. Botrel et al. 2007, Gonçalves et al. 2014). This is the first attempt to directly describe the anatomy of charcoal samples aiming to contribute to the construction of a database that might eventually contribute to prevent the continuity of deforestation and help the sustainability of the charcoal supply chain. The use of anthracology to control illegal charcoal commerce is, therefore, a particularly important application of the present work. Additionally, this work may contribute to a better knowledge of wood anatomy of native tropical species, which is still incompletely known. Indeed, while this is the first time that charcoal anatomy descriptions are presented for all the studied species, two species are here described for the first time both in charcoal as in conventional wood anatomy: Anacardium parvifolium Ducke and Spondias tuberosa Arruda. REFERENCES ABRAF. 2009. Anuário estatístico da ABRAF 2009: ano base 2008. Brasília: ABRAF - Associação Brasileira de Produtores de Florestas Plantadas. In: http://www.bibliotecaflorestal.ufv.br/handle/123456789/3905. ABRAF 2009 Anuário estatístico da ABRAF 2009: ano base 2008 Brasília ABRAF - Associação Brasileira de Produtores de Florestas Plantadas http://www.bibliotecaflorestal.ufv.br/handle/123456789/3905 ABRAF. 2013. Anuário estatístico da ABRAF 2013: ano base 2012. Brasília: ABRAF - Associação Brasileira de Produtores de Florestas Plantadas . In: http://www.bibliotecaflorestal.ufv.br/handle/123456789/3910. ABRAF 2013 Anuário estatístico da ABRAF 2013: ano base 2012 Brasília ABRAF - Associação Brasileira de Produtores de Florestas Plantadas http://www.bibliotecaflorestal.ufv.br/handle/123456789/3910 ALBUQUERQUE AR. 2012. Anatomia comparada do lenho e do carvão aplicada na identificação de 75 espécies da floresta Amazônica, no estado do Pará, Brasil. Unpublished Dissertation. Escola Superior de Agricultura "Luiz de Queiroz", Universidade de São Paulo, Brasil. ALBUQUERQUE AR 2012 Anatomia comparada do lenho e do carvão aplicada na identificação de 75 espécies da floresta Amazônica, no estado do Pará, Brasil Escola Superior de Agricultura "Luiz de Queiroz", Universidade de São Paulo Brasil BARAJAS-MORALES J and GÓMEZ CL. 1989. Anatomia de Maderas de Mexico: Especies de una selva baja caducifolia. Universidad Nacional Autónoma de México. BARAJAS-MORALES J GÓMEZ CL 1989 Anatomia de Maderas de Mexico: Especies de una selva baja caducifolia Universidad Nacional Autónoma de México BARROS CF and CALLADO CH. 1997. Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do Estado do Rio de Janeiro, Brasil. Vol. I. Rio de Janeiro: Jardim Botânico do Rio de Janeiro. BARROS CF CALLADO CH 1997 Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do Estado do Rio de Janeiro, Brasil I Rio de Janeiro Jardim Botânico do Rio de Janeiro BARROS CF, CALLADO CH, CUNHA M, MARCON-FERREIRA ML, TAMAIO N, MARQUETE O and COSTA CG. 2008. Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do estado do Rio de Janeiro, Brasil. Vol. III. Rio de Janeiro: Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. BARROS CF CALLADO CH CUNHA M MARCON-FERREIRA ML TAMAIO N MARQUETE O COSTA CG 2008 Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do estado do Rio de Janeiro, Brasil III Rio de Janeiro Instituto de Pesquisas Jardim Botânico do Rio de Janeiro BARROS CF, CALLADO CH, MARCON ML, COSTA CG, CUNHA M, LIMA HRP and MARQUETE O. 2001. Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do Estado do Rio de Janeiro, Brasil. Vol. II. Rio de Janeiro: Instituto de Pesquisas Jardim Botânico do Rio de Janeiro . BARROS CF CALLADO CH MARCON ML COSTA CG CUNHA M LIMA HRP MARQUETE O 2001 Madeiras da Mata Atlântica: Anatomia do lenho de espécies ocorrentes nos remanescentes florestais do Estado do Rio de Janeiro, Brasil II Rio de Janeiro Instituto de Pesquisas Jardim Botânico do Rio de Janeiro BLANKENHORN PR, JENKINS GM and KLINE D. 1972. Dynamic mechanical properties and microstructure of carbonized hardwoods. Wood Fiber 4(3): 212-224. BLANKENHORN PR JENKINS GM KLINE D 1972 Dynamic mechanical properties and microstructure of carbonized hardwoods Wood Fiber 4 3 212 224 BOTREL MCG, TRUGILHO PF, ROSADO SCS and SILVA RM. 2007. Melhoramento genético das propriedades do carvão vegetal de Eucalyptus. Rev Árvore 31(3): 391-398. BOTREL MCG TRUGILHO PF ROSADO SCS SILVA RM 2007 Melhoramento genético das propriedades do carvão vegetal de Eucalyptus Rev Árvore 31 3 391 398 BRASIL. 2012. Balanço energético nacional 2012: ano base 2011. Rio de Janeiro: Empresa de Pesquisa Energética. Ministério de Minas e Energia. In: https://ben.epe.gov.br/downloads/Relatorio_Final_BEN_2012.pdf. BRASIL 2012 Balanço energético nacional 2012: ano base 2011 Rio de Janeiro Empresa de Pesquisa Energética. Ministério de Minas e Energia https://ben.epe.gov.br/downloads/Relatorio_Final_BEN_2012.pdf CARNEIRO MS. 2008. Crítica social e responsabilização empresarial: análise das estratégias para a legitimação da produção siderúrgica na Amazônia Oriental. Cad CRH 21(53): 323-336. CARNEIRO MS 2008 Crítica social e responsabilização empresarial: análise das estratégias para a legitimação da produção siderúrgica na Amazônia Oriental Cad CRH 21 53 323 336 CHABAL L. 1997. Forêts et Sociétés en Languedoc (Néolithique Final, Antiquité Tardive). L'Anthracologie, Méthode et Paleoécologie. Doc archéol franç 63: 1-188. CHABAL L 1997 Forêts et Sociétés en Languedoc (Néolithique Final, Antiquité Tardive). L'Anthracologie, Méthode et Paleoécologie Doc archéol franç 63 1 188 CUTTER BE, CUMBIE BG and MCGINNES A. 1980. SEM and shrinkage analyses of Southern Pine wood following pyrolysis. Wood Sci Techn 14: 115-130. CUTTER BE CUMBIE BG MCGINNES A 1980 SEM and shrinkage analyses of Southern Pine wood following pyrolysis Wood Sci Techn 14 115 130 DECHAMPS R. 1979. Étude anatomique de bois d'Amérique du Sud. Volume I. Acanthaceae à Lecythidaceae. Tervuren: Musée Royal de l'Afrique Centrale. DECHAMPS R 1979 Étude anatomique de bois d'Amérique du Sud. Volume I. Acanthaceae à Lecythidaceae Tervuren Musée Royal de l'Afrique Centrale DÉTIENNE P and JACQUET P. 1983. Atlas d'Identification des bois de l'Amazonie et des régions voisines. Montpellier: Centre Technique Forestier Tropical. DÉTIENNE P JACQUET P. 1983 Atlas d'Identification des bois de l'Amazonie et des régions voisines Montpellier Centre Technique Forestier Tropical DONG Z and BAAS P. 1993. Wood Anatomy of Trees and Shrubs from China. V. Anacardiaceae. IAWA J 14(1): 87-102. DONG Z BAAS P. 1993 Wood Anatomy of Trees and Shrubs from China. V. Anacardiaceae IAWA J 14 1 87 102 FAO. 2014. FAOSTAT - Forestry Data. FAO - Food and Agriculture Organization of the United Nations. In: http://faostat3.fao.org/faostat-gateway/go/to/home/E. FAO 2014 FAOSTAT - Forestry Data FAO - Food and Agriculture Organization of the United Nations http://faostat3.fao.org/faostat-gateway/go/to/home/E FEDALTO LC, MENDES IC and Coradin VTR. 1989. Madeiras da Amazonia: Descrição do lenho de 40 espécies ocorrentes na Floresta Nacional do Tapajós. Brasília: Inst. Bras. Meio Ambiente e Rec. Nat. Renov. IBAMA. FEDALTO LC MENDES IC Coradin VTR. 1989 Madeiras da Amazonia: Descrição do lenho de 40 espécies ocorrentes na Floresta Nacional do Tapajós Brasília Inst. Bras. Meio Ambiente e Rec. Nat. Renov. IBAMA FRIEDERICHS G, BRAND MA, CARVALHO AF and KÜSTER LC. 2015. Qualidade da madeira e do carvão vegetal de bracatinga (Mimosa scabrella Benth.). Ciênc Madeira 6: 79-87. FRIEDERICHS G BRAND MA CARVALHO AF KÜSTER LC 2015 Qualidade da madeira e do carvão vegetal de bracatinga (Mimosa scabrella Benth.). Ciênc Madeira 6 79 87 GONÇALVES TAP. 2006. Contribuição ao conhecimento da anatomia da lenho das famílias Anacardiaceae, Annonaceae, Aquifoliaceae, Apocynaceae e Araliaceae, através de amostras carbonizadas do lenho de espécies brasileiras. Unpublished Monography. Graduação em Engenharia Florestal, Universidade Federal Rural do Rio de Janeiro, Brasil. GONÇALVES TAP. 2006 Contribuição ao conhecimento da anatomia da lenho das famílias Anacardiaceae, Annonaceae, Aquifoliaceae, Apocynaceae e Araliaceae, através de amostras carbonizadas do lenho de espécies brasileiras Graduação Universidade Federal Rural do Rio de Janeiro Brasil GONÇALVES TAP. 2010. Modificações estruturais decorrentes da carbonização no lenho de espécies arbóreas do cerrado no estado de São Paulo, Brasil. Unpublished Dissertation. Faculdade de Ciências Agronômicas, Universidade Estadual Paulista, Brazil. GONÇALVES TAP 2010 Modificações estruturais decorrentes da carbonização no lenho de espécies arbóreas do cerrado no estado de São Paulo, Brasil Faculdade de Ciências Agronômicas, Universidade Estadual Paulista Brazil GONÇALVES TAP, BALLARIN AW, NISGOSKI S and MUÑIZ GIB. 2014. A contribution to the identification of charcoal origin in Brazil. I - Anatomical characterization of Corymbia and Eucalyptus. Maderas, Cienc Tecnol 16(3): 323-336. GONÇALVES TAP BALLARIN AW NISGOSKI S MUÑIZ GIB 2014 A contribution to the identification of charcoal origin in Brazil. I - Anatomical characterization of Corymbia and Eucalyptus Maderas, Cienc Tecnol 16 3 323 336 GONÇALVES TAP and SCHEEL-YBERT R. 2012. Contra o carvão ilegal: estudo da anatomia da madeira pode ajudar a salvar florestas nativas. Ciência Hoje 292: 74-76. GONÇALVES TAP SCHEEL-YBERT R 2012 Contra o carvão ilegal: estudo da anatomia da madeira pode ajudar a salvar florestas nativas Ciência Hoje 292 74 76 GONÇALVES TAP, MARCATI CR and SCHEEL-YBERT R. 2012. The effect of carbonization on wood structure of Dalbergia violacea, Stryphnodendron polyphyllum, Tapirira guianensis, Vochysia tucanorum, and Pouteria torta from the Brazilian cerrado. IAWA J 33: 73-90. GONÇALVES TAP MARCATI CR SCHEEL-YBERT R 2012 The effect of carbonization on wood structure of Dalbergia violacea, Stryphnodendron polyphyllum, Tapirira guianensis, Vochysia tucanorum, and Pouteria torta from the Brazilian cerrado IAWA 33 73 90 GREGUSS P. 1959. Holzanatomie der Europäischen Laubhölzer und Sträucher. Budapest: Akad. Kiadó, 330 p. GREGUSS P 1959 Holzanatomie der Europäischen Laubhölzer und Sträucher Budapest Akad. Kiadó 330 GUPTA S and AGARWAL M. 2008. Wood anatomy of Anacardiaceae from India with special reference to the systematic position of Rhus. IAWA J 29: 79-106. GUPTA S AGARWAL M. 2008 Wood anatomy of Anacardiaceae from India with special reference to the systematic position of Rhus IAWA J 29 79 106 HESS RW. 1946. Identification of New World Timbers. Part II - Anacardiaceae. Trop Woods 87: 11-34. HESS RW 1946 Identification of New World Timbers. Part II - Anacardiaceae Trop Woods 87 11 34 IAWA COMMITTEE. 1989. IAWA list of microscopic features for hardwood identification. IAWA Bull 10(3): 219-332. IAWA COMMITTEE 1989 IAWA list of microscopic features for hardwood identification IAWA Bull 10 3 219 332 IBGE. 2010. Produção da extração vegetal e da silvicultura. Rio de Janeiro: IBGE - Instituto Brasileiro de Geografia e Estatística. 25: 50. IBGE 2010 Produção da extração vegetal e da silvicultura Rio de Janeiro IBGE - Instituto Brasileiro de Geografia e Estatística 25 50 50 IOS. 2011a. A floresta que virou cinza. 31 p. Instituto Observatório Social. In: <http://uc.socioambiental.org/sites/uc.socioambiental.org/files/revista_obs1.pdf> IOS 2011a A floresta que virou cinza 31 Instituto Observatório Social. http://uc.socioambiental.org/sites/uc.socioambiental.org/files/revista_obs1.pdf IOS. 2011b. O aço da devastação. 33 p. Instituto Observatório Social. In: <http://www1.ethos.org.br/EthosWeb/arquivo/0-A-620Pesquisa_A%20Devasta%C3%A7%C3%A3o%20do%20A%C3%A7o.pdf> IOS 2011b O aço da devastação 33 Instituto Observatório Social http://www1.ethos.org.br/EthosWeb/arquivo/0-A-620Pesquisa_A%20Devasta%C3%A7%C3%A3o%20do%20A%C3%A7o.pdf JOFRE MM. 1988. Estudio anatomico de las especies: Schinus latifolius (Molle) & Schinus molle (Pimiento). Depto. Tecnologia de la Madera, Universidad de Chile. JOFRE MM 1988 Estudio anatomico de las especies: Schinus latifolius (Molle) & Schinus molle (Pimiento) Depto. Tecnologia de la Madera, Universidad de Chile KIM NH and HANNA RB. 2006. Morphological characteristics of Quercus variabilis charcoal prepared at different temperatures. Wood Sci Techn 40: 392-401. KIM NH HANNA RB 2006 Morphological characteristics of Quercus variabilis charcoal prepared at different temperatures Wood Sci Techn 40 392 401 KRIBS DA. 1968. Commercial foreign woods on the American market. New York: Dover Publications, 241 p. KRIBS DA 1968 Commercial foreign woods on the American market New York Dover Publications 241 KRYN JM. 1952. The anatomy of the wood of the Anacardiacae and its bearing on the phylogeny and relationships of the family. Unpublished PhD thesis, University of Michigan, United States. KRYN JM 1952 The anatomy of the wood of the Anacardiacae and its bearing on the phylogeny and relationships of the family PhD University of Michigan United States LEÓN WJH. 2003. Estudio anatómico del xilema secundario de 17 especies de la familia Anacardiaceae en Venezuela. Acta Bot Venez 26: 1-30. LEÓN WJH 2003 Estudio anatómico del xilema secundario de 17 especies de la familia Anacardiaceae en Venezuela Acta Bot Venez 26 1 30 MC GINNES EA, KANDEL SA and SZOPA PS. 1971. Some structural changes observed in the transformation of wood into charcoal. Wood Fiber 3(2): 77-83. MC GINNES EA KANDEL SA SZOPA PS 1971 Some structural changes observed in the transformation of wood into charcoal Wood Fiber 3 2 77 83 MAINIERI C. 1958. Madeiras denominadas Caixeta. Inst. Pesq. Tec., São Paulo, n. 572. MAINIERI C 1958 Madeiras denominadas Caixeta Inst. Pesq. Tec. São Paulo 572 MAINIERI C and CHIMELO JP. 1989. Fichas de características das madeiras brasileiras. 2ª ed., IF, São Paulo. MAINIERI C CHIMELO JP 1989 Fichas de características das madeiras brasileiras 2ª IF São Paulo MARTIJENA N. 1987. Wood anatomy of Lithrea ternifolia (Gill.) Barkley & Rom. (Anacardiaceae). IAWA Bull 8: 47-52. MARTIJENA N. 1987 Wood anatomy of Lithrea ternifolia (Gill.) Barkley & Rom. (Anacardiaceae). IAWA Bull 8 47 52 METCALFE CR and CHALK L. 1950. Anatomy of dicotyledons. Oxford: Clarendon Press. METCALFE CR CHALK L. 1950 Anatomy of dicotyledons Oxford Clarendon Press MILLER RB. 2007. Fluorescent woods of the world. In: Flynn JH (Ed), A Guide to the More Useful Woods of the World. Madison: Forest Products Society, p. 271-305. MILLER RB 2007 Fluorescent woods of the world Flynn JH A Guide to the More Useful Woods of the World Madison Forest Products Society 271 305 NARDI BERTI R and ELDMANN ABBATE ML. 1992. Legnami tropicali importati in Italia: anatomia e identificazione. Vol. II. America Latina. Firenze: CNR. NARDI BERTI R ELDMANN ABBATE ML 1992 Legnami tropicali importati in Italia: anatomia e identificazione II Firenze CNR PAULA JE and ALVES JL. 1997. Madeiras Nativas: Anatomia, dendrologia, dendrometria, produção e uso. Fundação MOA, Brasília. PAULA JE ALVES JL 1997 Madeiras Nativas: Anatomia, dendrologia, dendrometria, produção e uso Fundação MOA Brasília PÉREZ OLVERA CP, VALDOVINOS CTF and GÓMEZ MAR. 1980. Estudio anatomico de la Madera de 43 especies tropicales. Bol Téc Inst Nac Invest For, n.63. PÉREZ OLVERA CP VALDOVINOS CTF GÓMEZ MAR 1980 Estudio anatomico de la Madera de 43 especies tropicales Bol Téc Inst Nac Invest For 63 PINHO RA. 1969. Contribuição ao Estudo Anatômico do Lenho Secundário de Árvores da Flora dos Cerrados - II. Arq Bot Estado de São Paulo 4: 4-6. PINHO RA 1969 Contribuição ao Estudo Anatômico do Lenho Secundário de Árvores da Flora dos Cerrados - II Arq Bot Estado de São Paulo 4 4 6 PINTO AA. 2013. Anatomia do Lenho Carbonizado de 10 Espécies Nativas da Planície Costeira do Rio Grande do Sul como Subsídio a Pesquisas Arqueobotânicas e Paleoecológicas. Unpublished Monography. Graduação em Arqueologia, Instituto de Ciências Humanas e da Informação, Universidade Federal do Rio Grande, Brasil. PINTO AA 2013 Anatomia do Lenho Carbonizado de 10 Espécies Nativas da Planície Costeira do Rio Grande do Sul como Subsídio a Pesquisas Arqueobotânicas e Paleoecológicas Graduação Instituto de Ciências Humanas e da Informação, Universidade Federal do Rio Grande Brasil PRIOR J and ALVIN KL. 1986. Structural changes on charring woods of Dichrostachys and Salix from Southern Africa: The effect os moisture content. IAWA Bull 7(3): 243-262. PRIOR J ALVIN KL. 1986 Structural changes on charring woods of Dichrostachys and Salix from Southern Africa: The effect os moisture content IAWA Bull 7 3 243 262 PRIOR J and GASSON P. 1993. Anatomical changes on six African hardwoods. IAWA J 14(1): 77-86. PRIOR J GASSON P 1993 Anatomical changes on six African hardwoods IAWA J 14 1 77 86 RECORD SJ and HESS RW. 1943. Timbers of the New World. Yale: Univ. Press, N. Haven, 640 p. RECORD SJ HESS RW 1943 Timbers of the New World Yale Univ. Press, N. Haven 640 RICHTER HG and DALLWITZ MJ. 2000 onwards. Commercial timbers: descriptions, illustrations, identification, and information retrieval. In English, French, German, Portuguese, and Spanish. Version: June 2009. In: http://delta-intkey.com RICHTER HG DALLWITZ MJ 2000 Commercial timbers: descriptions, illustrations, identification, and information retrieval http://delta-intkey.com SCHEEL-YBERT R. 2004a.Teoria e métodos em Antracologia. 1. Considerações teóricas e perspectivas. Arq Mus Nacional 62(1): 3-14. SCHEEL-YBERT R 2004a Teoria e métodos em Antracologia. 1. Considerações teóricas e perspectivas Arq Mus Nacional 62 1 3 14 SCHEEL-YBERT R. 2004b. Teoria e métodos em Antracologia. 2. Técnicas de campo e de laboratório. Arq Mus Nacional 62(4): 343-356. SCHEEL-YBERT R 2004b Teoria e métodos em Antracologia. 2. Técnicas de campo e de laboratório Arq Mus Nacional 62 4 343 356 SCHEEL-YBERT R and GONÇALVES TAP. In press. Primeiro Atlas Antracológico de Espécies Brasileiras / First Anthracological Atlas of Brazilian Species. Rio de Janeiro: Museu Nacional. SCHEEL-YBERT R GONÇALVES TAP Primeiro Atlas Antracológico de Espécies Brasileiras / First Anthracological Atlas of Brazilian Species Rio de Janeiro Museu Nacional SCHWEINGRUBER FH. 1990. Anatomy of European woods. Bern/Stuttgart: P. Haupt. SCHWEINGRUBER FH 1990 Anatomy of European woods Bern/Stuttgart P. Haupt. SCOTT AC and DAMBLON F. 2010. Charcoal: Taphonomy and significance in geology, botany and archaeology. Palaeogeogr Palaeoclimat Palaeoecol 291: 1-10. SCOTT AC DAMBLON F 2010 Charcoal: Taphonomy and significance in geology, botany and archaeology Palaeogeogr Palaeoclimat Palaeoecol 291 1 10 SOLARI ME. 1993. L'Homme et le bois en Patagonie et Terre de Feu au cours des six derniers millénaires: recherches anthracologiques au Chili et en Argentine. Unpublished PhD thesis, Université Montpellier-II, France. SOLARI ME 1993 L'Homme et le bois en Patagonie et Terre de Feu au cours des six derniers millénaires: recherches anthracologiques au Chili et en Argentine PhD Université Montpellier-II France SONSIN JO, GASSON P, MACHADO SR, CAUM C and MARCATI CR. 2014. Atlas da Diversidade de Madeiras do Cerrado Paulista / Atlas of Wood Diversity in the Cerrado of São Paulo. Botucatu: FEPAF. SONSIN JO GASSON P MACHADO SR CAUM C MARCATI CR. 2014 Atlas da Diversidade de Madeiras do Cerrado Paulista / Atlas of Wood Diversity in the Cerrado of São Paulo Botucatu FEPAF SOSEF MSM, HONG LT and PRAWIROHATMODJO S (Eds). 1998. Plant Resources of South-East Asia. Leiden: Backhuys Publishers. SOSEF MSM HONG LT PRAWIROHATMODJO S 1998 Plant Resources of South-East Asia Leiden Backhuys Publishers SOUZA VC and LORENZI H. 2008. Botânica Sistemática. Nova Odessa: Inst Plantarum. SOUZA VC LORENZI H 2008 Botânica Sistemática Nova Odessa Inst Plantarum TAVARES EJS. 1970. Anatomia do lenho de Schinus terebinthifolius Raddi. Bol Rec Nat SUDENE/DRN 8(1-2): 195-199. TAVARES EJS 1970 Anatomia do lenho de Schinus terebinthifolius Raddi Bol Rec Nat SUDENE/DRN 8 1-2 195 199 TERRAZAS TS. 1994. Wood anatomy of the Anacardiaceae: ecological and phylogenetic interpretation. Unpublished dissertation, University of North Carolina, United States. TERRAZAS TS 1994 Wood anatomy of the Anacardiaceae: ecological and phylogenetic interpretation University of North Carolina United States TERRAZAS TS and WENDT T. 1995. Systematic wood anatomy of the genus Tapirira Aublet (Anacardiaceae): a numerical approach. Brittonia 47(2): 109-129. TERRAZAS TS WENDT T. 1995 Systematic wood anatomy of the genus Tapirira Aublet (Anacardiaceae): a numerical approach Brittonia 47 2 109 129 TORTORELLI LA. 1956. Maderas y bosques argentinos. Buenos Aires: Editorial Acme, 910 p. TORTORELLI LA 1956 Maderas y bosques argentinos Buenos Aires Editorial Acme 910 VERNET JL, OGEREAU P, FIGUEIRAL I, MACHADO YANES C and UZQUIANO P. 2001. Guide d'identification des charbons de bois préhistoriques et récents: Sud-Ouest de l'Europe : France, Péninsule Ibérique et Îles Canaries. Paris: CNRS ed., 395 p. VERNET JL OGEREAU P FIGUEIRAL I MACHADO YANES C UZQUIANO P 2001 Guide d'identification des charbons de bois préhistoriques et récents: Sud-Ouest de l'Europe : France, Péninsule Ibérique et Îles Canaries Paris CNRS 395 WILLIAMS L. 1936. Woods of Northeastern Peru. Botanical Series v 15. Chicago: Field Museum of Natural History, 608 p. WILLIAMS L 1936 Woods of Northeastern Peru. Botanical Series v 15 Chicago Field Museum of Natural History 608