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Taxonomy, Buchenavia, Terminalia, Com• in Their 1963 Revision Exell & Stace Recog• Bretaceae

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Taxonomy, Buchenavia, Terminalia, Com• in Their 1963 Revision Exell & Stace Recog• Bretaceae IAWA Bulletin n.s., Vol. 12 (2),1991: 123-141 WOOD AND BARK ANATOMY OF BUCHENAVIA EICHL. (COMBRETACEAE)l by Veronica Angyallossy Alfonso2 and Hans Georg Richter3 Summary Karstedt & Parameswaran 1976; Richter Wood and bark structure of Buchenavia 1981a, b, 1985, 1990; Trockenbrodt 1989; (Combretaceae) was studied for infrageneric Trockenbrodt & Parameswaran 1986). variation and taxonomic utility. Despite its Wood and bark structure of 15 species (23 wide neotropical distribution Buchenavia has samples) of Buchenavia and 5 species a homogeneous wood anatomy, axial paren­ (8 samples) of neotropical Terminalia were chyma distribution being the most variable examined to see if they - either singly or feature. The presence or absence of septate in combination - provided useful features fibres and presence and location of silica for distinguishing between Buchenavia and grains in ray and/or axial parenchyma might Terminalia. Buchenavia and Terminalia are eventually allow recognition of individual known to have similar wood anatomy. Bark species or species groups. Bark anatomy of­ anatomy might provide a means of distin­ fers no diagnostic features for. species differ­ guishing between the two genera, and be­ entiation. Wood and bark structure of Buche­ tween Buchenavia species. navia (15 species studied) intergrades fully The genus Buchenavia Eichl. was estab­ with that of neotropical Terminalia (5 species lished in 1866 (Exell & Stace 1963) by the studied), thus corroborating results of studies German botanist A.W. Eichler (1839-1887) on morphology (Exell & Stace 1963) and leaf who was co-editor of Flora Brasiliensis. His anatomy (Stace, personal communication, original treatise of Buchenavia included eight 1989) which postulate a very close relation­ species. More recently Buchenavia has been ship between the two genera. studied by Exell & Stace (1963, 1966) and Key words: Wood anatomy, bark anatomy, Alwan AI-Mayah & Stace (1985). taxonomy, Buchenavia, Terminalia, Com­ In their 1963 revision Exell & Stace recog­ bretaceae. nised 24 validly described species. In a forthcoming monograph of neotropical Com­ bretaceae to be published in Flora Neotropica Introduction (Stace, in print) the number of valid species The present study was undertaken to con­ is reduced to 21. tribute to the general taxonomy of Buchenavia, Buchenavia is part of the tribe Terminalieae particularly under the aspect of combined of the family Combretaceae and most closely wood an bark anatomy. Increasingly, the lat­ related to Terminalia (Exell & Stace 1963). ter is employed in differentiating taxa of simi­ Stace (pers. comm., 1989) reports that there lar or nearly identical wood structure, and are no absolute vegetative differences be­ has proven a useful complementary element tween Buchenavia and Terminalia while dif­ in taxonomic studies (e.g. Alfonso 1983; ferences in flower and fruit are always clear­ Chattaway 1955a-c; Esau & Cheadle 1984; cut. 1) Dedicated to Professor Dr. Drs. mult. h. c. Walter Liese on the occasion of his 65th birthday (31 January 1991). 2) Instituto de Pesquisas Tecnologicas, Divisao de Madeiras, Cidade Universitana, 05508 Sao Paulo, SP, Brazil. 3) Bundesforschungsanstalt fUr Forst- und Holzwirtschaft, Institut fUr Holzbiologie und Holzschutz, Leuschnerstr. 91, 2050 Hamburg 80, Germany. Downloaded from Brill.com09/26/2021 12:15:20PM via free access 124 IAWA Bulletin n.s., Vol. 12 (2),1991 Buchenavia is confined to but fairly wide­ cently, taxonomic aspects were highlighted spread in tropical America from the Carib­ by Den Outer and Fundter (1976) who dis­ bean to southern Brazil, with the largest cussed the bark structure of Strephonema number of species concentrated in northern pseudocola in relation to other taxa of Com­ continental South America (Guyanas, Ama­ bretaceae of African origin (Buchenavia not zon valley, see map). The genus is repre­ included). Zahur (1959) described bark of sented by small shrubs to large trees from Terminalia catappa and T. myriocarpa from about three to fifty metres tall, the former Hawaii; Roth (1981) described bark of Buch­ growing mostly in frequently inundated low­ enavia capitata, Terminalia guyanensis and T. land forests, the latter more in upland rain amazonia, all from Venezuela. forests (Exell & Stace 1963). Material and Methods Tables 1 (Buchenavia) and 2 (Terminalia) list the species studied and gives information on origin, collectors', herbarium and wood collection numbers. For most (9 of the 15) Buchenavia and (4 of the 5) Terminalia spe­ cies only one sample was available for exam­ ination. Microscopic sections 10-15 Jlm in thick­ ness were prepared from all wood specimens following common laboratory procedures, stained with malachite green (transverse and tangential) and safranin (radial), and subse­ quently mounted permanently in a synthetic medium ('Permount'). Bark material was sectioned after penetration with polyethylene glycol (DP 15(0); individual sections of 10 to 25 Jlm thickness, protected by scotch tape or a hardened layer of commercial nail polish. The 'scotch tape' method follows the procedures outlined by Rupp (1964), modi­ Distribution of Buchenavia. fied by Wilhelm (1975) and Kruse (1977). The staining of the sections is done while still As the wood of Buchenavia species is adhering to the tape. In the other case, micro­ very similar to that of Terminalia in appear­ tome sections from the PG-impregnated block ance and gross structure they cannot be easily are glued to a cover glass by means of a thin separated and are traded jointly as one group layer of colourless nail polish. Further pro­ of commercial timbers. They are used mainly cessing (staining, dehydration, mounting) is in internal and external construction, as rail­ done with the sections firmly adhering to the way sleepers, cross arms, posts, and for fur­ cover glass. The sections were double-stained niture (Mainieri et at. 1983). with malachite green and hematoxylin or, in There are various descriptions of the few cases, with astra blue and chrysoidine/ wood anatomy of Buchenavia (e.g. Detienne acridine red. Macerations were prepared with & Jacquet 1983; Lindeman & Mennega 1963; Jeffrey's solution (equal parts of 10% ae­ Mainieri et al. 1983); however, only Van quous solution of nitric and chromic acid) Vliet (1979) has discussed the wood struc­ and stained with safranin (wood) or double­ ture of Buchenavia as part of a comparative stained with chrysoidine/acridine red and study of the whole family. astra blue (bark). The bark structure of some combretaceous Quantitative data are based on 10 (vessel taxa (Bucida, Combretum, Terminalia) was pit diameter), 25 (diameter and length of ves­ first described by Moeller (1882). More re- sel elements, fibre diameter wood/bark), 30 Downloaded from Brill.com09/26/2021 12:15:20PM via free access Alfonso & Richter - Wood and bark anatomy of Buchenavia 125 Table 1. Specimens examined of the genus Buchenavia Eiehl. Species Origin ColI. Nr., Herb. Nr. Wood ColI. Nr. B. amazonia Alwan & Stace* Guyana Sebatier & Prevost 2179 Uw 32810 B. capilala (Vahl) Eichl. Brazil,PA MGw4154 idem Brazil,AM Kubitzki 75-87 RBHw 16699 idem Surinam, Rio Para B.B.S. 91 (sterile) Uw686 B. congesla Ducke Brazil,AM W. Rodrigues,lNPA 15008 lNPAw X-2581 B. grandis Ducke Brazil,AM -, Museu Goeldi 55715 MGw2938 idem Brazil,AM Loureiro, Coelho & Mello lNPAw X-2657 (received as B. huberi) lNPA 14480 idem Brazil,AM Krukoff 6472 Uw7754 B. kleinii Exell ? Brazil,PR Lindeman & De Haas 2350 Uw 13694 (sterile) B. macrophyl/a Eichl. Brazil,AM Lima & Chagas,lNPA 13306 lNPAw X-1612 B. ochroprumna Eichl. Brazil,AM W. Rodrigues & Coelho 6770, lNPAw X-3091 lNPA 15234 idem Brazil,AM Coelho & Lima 380, lNPAw X-6800 lNPA 81230 B. oxycarpa (Mart.) Eichl. Brazil,AM W. Rodrigues 2254, lNPAwX-891 lNPA8665 idem Brazil,PA Kubitzki 82 -4 7 RBHw 19170 B. pallidovirens Cuatrec. Columbia Rooden et al. 545 Uw25612 B. parvifolia Ducke Brazil,AM Loureiro, Coelho & Mello, lNPAw X-2626 lNPA 14449 idem Brazil,AM W. Rodrigues 5466, lNPAw X-2017 lNPA 14135 B. rabel/oana Mattos Brazil, SP Kuhlmann s.n., SP 157742 SPW 43 B. sericocarpa Ducke* Brazil,AM Krukoff 6916 (type) Uw8044 B. suaveolens Eichl. Brazil,AM Kubitzki 75-87a RBHw 16683 B. lomen/osa Eichl. Brazil, ill -,lNPA 15839 lNPAw X-3278 B. viridiflora Ducke Guyana BAFOG 1313 Uw 5801 idem Brazil,AM W. Rodrigues & Loureiro 65 lNPAw X-3571 * Specimens without bark. (rays/mm, ray height in mm and number of The terms used in wood and bark descrip­ cells) and 50 (number of vessels/mm) and tions adhere to the character lists proposed by fibre length wood/bark) individual counts, IAWA (1989) and Trockenbrodt (1990), re­ respectively. In all cases but vessel pit diam­ spectively. eter the statistical requirements of minimum number of measurements (N = t2 x s2 / E2; Wood description where t = student t, s = standard deviation, E 2 = (0.1 x X)2, according to Freese 1967) General: Heartwood light brown to yellow­ were fulfilled. The numerical values given ish, not well differentiated from the lighter in the description are the ranges of species coloured sapwood; grain straight to slightly means accompanied by overall minimum and wavy; texture fine to medium; surface non­ maximum values between brackets. Specific fluorescent; without specific odour or taste. collection numbers are cited for species with Growth increments more or less distinct. multiple samples to indicate that this is the Anatomy: Vessels arranged in a diffuse-por­ only sample(s) that showed this particular ous pattern, approaching semi-ring-porous in feature. B. ochroprumna; solitary and in radial multi- Downloaded from Brill.com09/26/2021 12:15:20PM via free access 126 IAWA Bulletin n.s., Vol. 12 (2),1991 Table 2. Specimens examined of the genus Terminalia L. Species Origin ColI. Nr., Herb. Nr. Wood Coil. Nr. T. amazonia (Gmel.) Exell Venezuela -, Museu Goeldi 31909 MGw779 idem Brazil,PA N.T. Silva 3412 BCTw 13254 idem· Guatemala Inv.For.EI Petn 9512 RBHw 12861 idem· Surinam Stahel93a RBHw 18688 T.
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