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Downloaded from Brill.Com10/04/2021 04:40:51AM Via Free Access 386 IAWA Journal, Vol IAWA Journal, Vol. 18 (4), 1997: 385-399 BARK ANATOMY OF ARBORESCENT LEGUMINOSAE OF CERRADO AND GALLERY FOREST OF CENTRAL BRAZIL by Cecilia G. Costa 1, Vera T. Rauber Coradin 2, Chiudia M. Czarneski2 & Benedito A. da S. Pereira 3 SUMMARY The bark anatomy of28 species of arborescent Leguminosae of 'cerrado' and gallery forest in the Brazilian Federal District was examined. The most significant characteristics for taxonomic purposes were determined to be: delimitation between collapsed and non-collapsed phloem; phloem stratification; type and position of sieve plates; dilatation patterns; ar­ rangement and contents of sc1ereids; and presence of secretory cells. The bark data support the idea that Papilionoideae is the most advanced group of the Leguminosae. Key-words: Bark anatomy, cerrado, Leguminosae, phloem. INTRODUCTION Morphological and anatomical bark characteristics have been used for identification and evaluation of the taxonomic position of plants, e. g. Whitmore (1962), Teixeira et al. (1978), Roth (1981), Trockenbrodt & Parameswaran (1986), Richter (1990), Archer & Van Wyk (1993). There have been more anatomical studies of xylem than of bark because of the economic importance of xylem, its higher decay-resistance and its rela­ tive ease of preparation for anatomical observations. Most wood anatomists use the same terrninology (IAWA 1989), wh ich is not the case for the bark, in spite of the efforts of Esau (1969) and others. Roth (1981) suggested some terms, but her defini­ tions sometimes conflict with already existing ones. Trockenbrodt (1990) critically reviewed the terrninology used in bark anatomy. More recently, Junikka (1994) re­ vised terms used in macroscopical analysis of the bark. The outer bark functions as a thermal insulant that protects the cambium and con­ ducting tissue against high temperatures in forest fires, which is an important role in ecosystems such as the cerrado that are subjected to periodic burnings (MacieI1993). This paper presents an anatomical study of the bark of arborescent Leguminosae occurring in cerrado and in gallery forest of Brazil and aims to deterrnine which fea- 1) Area de Botänica Estrutural, Jardim Botänico do Rio de Janeiro, Rua Jardim Botänico, 1008, 22.460-000 Rio de Janeiro, RJ, Brazil. 2) Laborat6rio de Produtos Florestais, LPF / IBAMA, SAIN Av. L4 Lote 04, 70.818-900 Brasflia, DF, Brazil. 3) Instituto Brasileiro de Geografia e Estatfstica, IBGE, Rod. BR 251 Km 1, 70.000 Brasflia, DF, Brazil. Downloaded from Brill.com10/04/2021 04:40:51AM via free access 386 IAWA Journal, Vol. 18 (4), 1997 tures are useful for identification. This family was chosen because of its variety in bark structure (Roth 1981) and for its economical importance. Leguminosae are conspicu­ ous in the cerrado and gallery forest, both in terms of species diversity as weIl as abundance of individuals (Kirkbride 1984). MATERIALS AND METHODS This investigation is based on material from the central region of Brazil. The Leguminosae species studied are listed below, arranged alphabeticaIly. The samples were obtained from the trunk of adult trees of 10 cm or more in diam­ eter, at a height of 1.0-1.3 m. Two sampIes from different individua1s were used when­ ever possib1e. After impregnation with glycol polyethylene (PEG, DP 1500) (Rupp 1964), the material was cut at a thickness of 10-12 11m and double stained with crysoi­ din/ acridin red and astra blue. Maceration of the phloem elements was obtained by a modification ofFranklin's method (Fedalto 1982). Histochemistry tests for lignin and phenolic compounds were made according to Johansen (1940) and Jensen (1962), re­ spectively. Crystals were observed in polarized light and specific tests were used to determine their chemical composition (Strasburger 1924). The G-layer in the gelati­ nous fibres was detected by chlorozinc iodine (Strasburger 1924) and polarized light. Periderm was observed in only some species studied, because it easily falls out in others. The terminology used in this work follows Trockenbrodt's (1990) recommenda­ tions. List of studied species Each name is followed by subfamily, habitat, geographical origin, collection date and herbarium number. Acacia polyphylla A.DC.: Mimosoideae, cerrado, Brasilia (DF) / Brazil, 01109/94, IBGE 32283 - gallery forest, Palmital (MG) / Brazil, 16/11194, IBGE 32474. - Acosmium dasycarpum (Vog.) Yakovl.: Papilionoideae, cerrado, Brasilia (DF) / Brazil, 16/11194, IBGE 32477. - Ana­ denanthera peregrina (L.) Speg.: Mimosoideae, cerrado, Brasilia (DF) / Brazil, 07/93, IBGE 32295 - gallery forest, Palmital (MG) / Brazil, 16/11194, IBGE 32469. -Apuleia leiocarpa (Vog.) Macbr.: Caesalpinioideae, gallery forest, Palmital (MG) / Brazil, 25/03/92, IBGE 29032 - gallery forest, Brasilia (DF) / Brazil, 01/09/94, IBGE 32478. - Bowdichia virgilioides Kunth: Papilionoideae, cerrado, Brasilia (DF) / Brazil, 07/93, IBGE 32296. - Centrolobium to­ mentosum Guilles ex Benth.: Papilionoideae, gallery forest, Brasilia (DF) / Brazil, 01109/94, IBGE Carp. 287. - Copaijera langsdorffii Desf.: Caesalpinioideae, gallery forest, Brasilia (DF) / Brazil, 07/93, IBGE 32293. - Dalbergia miscolobium Benth.: Papilionoideae, cerrado, Brasilia (DF) / Brazil, 23/06/94, IBGE 32287. - Dimorphandra gardneriana Tu!.: Caesalpinioideae, gallery forest, Dian6polis (TO) / Brazil, 12/92, IBGE 29049. - Dimorphandra mollis Benth.: Caesalpinioideae, cerrado, Brasilia (DF) / Brazil, 07/91, IBGE 30218 - cerrado, Brasilia (DF) / Brazil, 16/11194, IBGE 32475. - Dipteryx alata Vog.: Papilionoideae, cerrado, Dian6polis (TO) / Brazil, 06/12/91, IBGE 28783 - cerrada, Sao Joao da Alian<;a (GO) / Brazil, 10/09/94, IBGE 32352. - Enterolobium contortisiliquum (VelI.) Morong.: Mimosoideae, cer­ rada / gallery forest, Brasilia (DF) / Brazil, 01/09/94, IBGE 32297. - Enterolobium gummijerum Downloaded from Brill.com10/04/2021 04:40:51AM via free access Costa, Rauber Coradin, Czarneski & Pereira - Bark of Leguminosae 387 (Mart.) Macbr.: Mimosoideae, cerrado, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32470 - cerra­ do, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32461. - Erythrina verna Vell.: Papilionoideae, cerrado, Brasilia (DF) 1 Brazil, 01/91, IBGE 30101 - gallery forest, Brasilia (DF) 1 Brazil, 01/ 09/94, IBGE 32288. - Hymenaea courbaril L.: Caesalpinioideae, gallery forest, Brasilia (DF) 1 Brazil, 07/93, IBGE 32289. - Hymenaea stigonocarpa Mart. ex Hayne: Caesalpinioideae, cerrado, Brasilia (DF) 1 Brazil, 07/93, IBGE 32294. - Machaerium opacum Vog.: Papilionoideae, cerrado, Brasilia (DF) 1 Brazil, 12/91, IBGE 27153 - cerrado, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32473 - cerrado, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32471. -Mi­ mosa laticifera Rizzini & Mattos: Mimosoideae, cerrado, Dian6polis (TO) 1 Brazil, 01/92, w 1n. - Parkia platycephala Benth.: Mimosoideae, cerrado, Dian6polis (TO) 1Brazil, 03/12/91, IBGE 29079. - Plathymenia reticulata Benth.: Mimosoideae, cerrado, Brasilia (DF) 1 Brazil, 01/091 94, IBGE 32282. - Platymisciumfloribundum Vog.: Papilionoideae, gallery forest, Brasilia (DF) 1 Brazil, 01/09/94, IBGE 32284. - Platypodium elegans Vog.: Papilionoideae, gallery forest, Brasilia (DF) 1Brazil, 07/93, IBGE 32290. - Pterodon emarginatus Vog.: Papilionoideae, cerrado, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32472. - Sclerolobium aureum (Tu!.) Benth.: Caesalpinioideae, cerrado, Barreiros (DF) 1 Brazil, 12/91, IBGE 28985 - cerrado, Barreiros (DF) 1 Brazil, 02/92, IBGE 31323 - cerrado, Barreiros (DF) 1 Brazil, 07/93, IBGE 32291.­ Sclerolobium paniculatum var. rubiginosum (Mart. ex Tu!.) Benth.: Caesalpinioideae, gallery forest, Brasilia (DF) 1 Brazil, 12/1 0/92, IBGE 31134 - gallery forest, Brasilia (DF) 1 Brazil, 071 93, IBGE 32292. - Sclerolobium paniculatum var. subvelutinum Benth.: Caesalpinioideae, cerrado, Brasilia (DF) 1 Brazil, 06/94, IBGE 32285. - Slryphnodendron adstringens (Mart.) Covile: Mimosoideae, cerrado, Brasilia (DF) 1 Brazil, 23/06/94, IBGE 26999 - cerrado, Brasilia (DF) 1 Brazil, 23/06/94, IBGE 26107. - Vatairea macrocarpa (Benth.) Ducke: Papilionoideae, cerrado, Brasilia (DF) 1 Brazil, 16/11/94, IBGE 32476. RESULTS Usually the boundary between collapsed and non-collapsed phloem is gradual. A few species have a sharp dei imitation between the two regions, e. g. Acacia polyphylla, Co­ paifera langsdorffii, Dimorphandra mollis, Hymenaea courbaril and Mimosa laticifera. The phloem is usually storied in Papilionoideae (Fig. 1), and non-storied in Caesal­ pinioideae and in Mimosoideae. Among the Caesalpinioideae observed, only Apuleia leiocarpa has storied phloem (Fig. 2). In the majority of Papilionoideae, the conduc­ tive elements, parenchyma cells, fibres and secretory cells are organized in short tan­ gential bands and these bands present a radial pattern, demarcated laterally by phloem rays. Only one member ofPapilionoideae studied, Erythrina verna, presents very large bands ofthose elements (Fig. 3). Gelatinous fibres usually occur in the phloem (Fig. 3, 13, 14); generally they form larger bands in the collapsed phloem, whose walls often are flattened and impregnated by phenolic compounds. The sieve elements can occur isolated or in groups of 2-4 or more elements, in tangential, radial or crowded arrangement. The sieve elements in trans verse section have an irregular shape; in the Caesalpinioideae, they are 268-527 f.lIll in length and 20-37 /lm in diameter, in the Mimosoideae 228-420 f.lIll in length and 20-51 f.lIll in diameter, and in the Papilionoideae 153-304/lm in length and 18-42/lm in diameter (Table I). The Papilionoideae
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