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

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.

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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

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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-

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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. argentea Mart. & Eichl. Brazil,MG Heringer & Filho 4032, RBw 3540 = RBHw 10927 RB 93481 T. brasiliensis Eichl. Brazil,PA MGw396 T. guianensis Aubl. Brazil,PA -, Museu Goeldi 90567 MGw3233 T. cf. tanibouca Smith Brazil,MA MGw2279

• Specimens without bark.

pIes of 2, rarely up to 5, exceptionally up to X-2626), B. rabelloana, B. seriocarpa, B. 10 in B. tomentosa; tangential diameter (40-) suaveolens, B. tomentosa; with marked con• 60-220(-280) J.1m, frequency variable, (1-) fluence grading into conspicuously banded 4-22(-33) vessels/mm2; vessel element parenchyma in B. macrophylla, B.ochro• length (155-)310-630(-800) J.1m; perfora• prumna, B. oxycarpa (Fig. 6) and B. pallido• tion plates simple; intervessel pits 6-10(-11) virens; marginal or seemingly marginal pa• J.1m in diameter (axial dimension), alternate, renchyma present in B. amazonia, B. capita• circular to polygonal in outline, vestured; ta, B. grandis (INPAw X-2657, Uw 7754), vessel-ray pits similar to intervessel pits in B. ochroprumna, B. pallidovirens, B. parvi• size and shape, mostly circular, simple or folia (INPAw X-2626), B. rabelloana (Fig. with much reduced borders. Helical thicken• 2), B. seriocarpa, B. suaveo!ens, B. tomen• ings throughout body of narrow vessel ele• tosa and B. viridiflora, poorly developed in ments observed only in B. grandis (Uw B. capitata (MOw 4154) and B. congesta; 7754, Fig. 14) and B. suaveolens; thin-wall• diffuse parenchyma generally present in ed tyloses and yellowish to brown gum de• small quantities, absent in B. capitata (MOw posits in vessels of B. oxycarpa. 4154), B. grandis (INPAw X-2657), B. Fibres thin to very thick-walled, the latter ochroprumna (INPAw X-3091), B. oxycarpa only at growth increment borders (Figs. 2,4); and B. pallidovirens. length (570-)920-1630(-2180) J.1m, diam• Rays exclusively uniseriate (including lo• eter (10-)15-19(-32) J.1m; pits simple to cally biseriate rays) in most species (Fig. 9) minutely bordered, few fibres with distinct• with a small portion (7-23%) of biseri• ly bordered pits in B. amazonia (Fig. 11), B. ate rays in B. congesta and B. macrophylla; capitata (RBHw 16699), B. congesta, B. predominantly biseriate with few triseriates (z grandis (MOw 2938), B. parvifolia, B. sua• 2%) in B. tomentosa (Fig. 10); very variable veolens; septa present in some fibres of B. in height, (100-)180-350(-710) J.1m; all capitata (Uw 686), B. oxycarpa (Fig. 12) and cells procumbent or body cells procumbent B. seriocarpa. with one marginal row of square/upright Parenchyma predominantly paratracheal cells; usually with reddish-brown gum aliform to confluent with a large variation in deposits. the degree of confluence: lozenge-aliform in Inorganic inclusions: prismatic crystals B. parvifolia (INPAw X-20l7, Fig. 1) and present in the majority of Buchenavia spe• B. viridiflora; confluent well-developed in B. cies; occasionally also styloids (Fig. 18) in amazonia, B. capitata, B. congesta, B. gran• B. amazonia, B. congesta, B. kleinii, B. dis (Fig. 3), B. kleinii, B. parvifolia (INP A w ochroprumna (INPAw X-3091), B. oxycar-

Downloaded from Brill.com09/26/2021 12:15:20PM via free access Alfonso & Richter - Wood and bark anatomy of Buchenavia 127 pa (INPAw X-891), B. pallidovirens, B. ra• Sieve tubes (and companion cells) solitary bel/oana, often surrounded by a distinct or in small groups of 2 or 3, with little varia• sheath; predominantly in non-chambered ray tion throughout the genus, interspersed with cells (one crystal per cell, Fig. 19); exclu• parenchyma cell strata; cells irregular in out• sively associated with axial parenchyma line (Fig. 26), about 30-50 J.Lm in diameter (chambered cells) in B. capitata (MGw 4154, and 500-600 J.Lm in length; sieve plates hori• Fig. 17) and B. kleinii; in both radial and zontal to inclined, simple and compound in axial parenchyma in B. amazonia, B. conges• one and the same or different elements (Fig. ta, B. macrophyl/a, B. ochroprumna (INPAw 28), with 3-6 sieve areas, each with sieve X-6800), B. rabel/oana, B. tomentosa; no pores of 2-3 11m in diameter; lateral sieve crystals observed in B. capitata (RBHw areas present. 16699), B. oxycarpa (RBHw 19170), B. Phloem rays similar to xylem rays in com• seriocarpa, B. viridiflora. position and dimensions (Fig. 22); with con• Globular silica grains in axial parenchyma siderable numbers of intercellular spaces in of B. macrophyl/a, B. ochroprumna (INPAw specimens of B. capitata, B. parvijolia (Fig. X-6800, Fig. 15), B. seriocarpa; in both rays 33), B. rabel/oana and B. viridiflora; dilated (Fig. 16) and axial parenchyma of B. capitata towards the periphery due to tangential cell (RBHw 16699). division and/or inflation (Figs. 20, 21); dila• Traumatic canals observed in B. capitata, tation in some instances well developed, re• B. rabel/oana (Fig. 13) andB. tomentosa. sulting in considerable distortion of the tissue Growth increments: When present delim• arrangement in the outer bark; when in con• ited by tangential zones of thick-walled, radi• tact with 'sclerified cells' (see below) some ally flattened fibres in B. kleinii, B. parvijo• ray cells become sclerified and develop lia (INPAw X-2017; Fig. 1, see arrow); also branches (,branched ray sclereids', Figs. 39, in combination with marginal parenchyma as 40) gradually growing into intercellular in B. capitata (Fig. 4), B. rabel/oana (Fig. 2), spaces as observed in B. congesta and B. ra• B. suaveolens, B. viridijlora; in some in• bel/oana (see also Figs. 41, 42 for same fea• stances associated with a marked difference ture in Terminalia). of vessel diameter between early and late• Phloem fibres (Fig. 29, maceration) ar• wood, e.g. in B. ochroprumna (Fig. 5). ranged in regular 1- to 3-seriate continuous tangential bands in B. capitata (Fig. 20), B. congesta, B. grandis (MGw 2938), B. och• Bark description roprumna, B.oxycarpa, B. suaveolens; in General: Dark brown to reddish brown exter• small, tangentially arranged groups of less nally, locally with grey to whitish areas; sur• regular and frequent occurrence in B. grandis face smooth to fissured in a reticulate pattern; (INPAw X-2657), B. macrophyl/a, B. palli• fissures usually narrow, longitudinal and dovirens, B. parvijolia, B. rabel/oana, B. to• transverse, often cutting fairly deep into the mentosa, B. viridiflora; fibre length varying rhytidome layers; of varying thickness (0.2- from (440-)730-1910(-2340) Ilffi; diameter 1 cm) in function of bole diameter/age. variable from (12-)17-25(-35) 11m; fibres frequently gelatinous in all species (Fig. 27, Anatomy: Bark regularly stratified from cam• short arrow). bium towards periphery; strata formed by 'Sclerified cells': cells of considerable di• multi seriate tangential bands of phloem pa• mensions in transverse section (radially larger renchyma and conducting cells (sieve tubes than tangentially) with thick, polylamellate and companion cells) alternating with rows (Fig. 32), generally gelatinous walls (Fig. of sclerenchymatic cells (Figs. 20, 21, 23; 27, long arrow); axially elongated due to pro• see also Fig. 24 for same feature in Termi• nounced apical growth, varying from very nalia brasiliensis). long and thin (i.e. fibre-like, Fig. 30) to Phloem parenchyma arranged in 2- to 4- wide-bodied and short (Fig. 31); length seriate tangential bands in all species; cells (330-)540-1070(-1320) 11m and (30-)40- with large and circular simple pits. (text continued on page 135)

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Legends of Figures 1-42:

Figs. 1-6. Buchenavia, wood, transverse sections, x 44. - 1: B. parvifolia, lozenge-aliform parenchyma; growth ring boundaries indicated by thicker-walled and radially flattened fibres (arrow). - 2: B. rabelloana, aliform with some confluent and marginal parenchyma; growth ring boundaries marked by thick-walled, radially flattened fibres and marginal parenchyma. - 3: B. grandis, aliform, confluent and marginal parenchyma. - 4: B. capitata, aliform, confluent and marginal parenchyma; growth ring boundaries marked by thick-walled fibres and marginal pa• renchyma. - 5: B. ochroprumna, aliform parenchyma with pronounced confluence; growth ring boundaries associated with change in vessel diameter. - 6: B. oxycarpa, parenchyma with pro• nounced confluence forming bands.

Figs. 7 & 8. Terminalia guianensis, wood. - 7: Transverse section, aliform, confluent and mar• ginal parenchyma, traumatic canals (arrows), x 44. - 8: Tangential section, uniseriate rays with crystals, x 110. - Figs. 9-11. Buchenavia, wood, tangential section. - 9: B. grandis, uniseri• ate rays, x 110. - 10: B. tomentosa, predominantly biseriate, also triseriate rays, x 44. - 11: B. amazonia, fibres with large bordered pits, x 440. - Fig. 12: B. oxycarpa, wood, radial section, septate fibres (arrows), x 440. - Fig. 13: B. rabel/oana, wood, transverse section, traumatic canals, x 44.

Figs. 14-19. Buchenavia, wood. - 14: B. grandis, tangential section, helical thickenings in nar• row vessel element, x 220. - 15: B. ochroprumna, radial section, silica grains in axial paren• chyma, x 220. - 16: B. capitata, radial section, silica grains in rays, x 220. - 17: B. capitata, radial section, crystals in chambered cells, x 220. - 18: B. tomentosa, tangential section, styloid crystals in axial parenchyma, x 220. - 19: B. parvifolia, radial section, crystals in rays, x 110.

Figs. 20-23. Buchenavia, bark. - 20: B. capitata, transverse section, stratified, phloem paren• chyma alternating with phloem fibres (arrow); dilated rays (D); periderm (P), x 60. - 21: B. grandis, transverse section, stratified, phloem parenchyma alternating with 'sclerified cells' (ar• rows); dilated ray (D); periderms (P), x 44. - 22: B. ochroprumna, tangential section, uniseriate rays, druses in chambered cells, x 110. - 23: B. pallidovirens, transverse section, earlier formed phloem with predominantly phloem fibres (white radial line), and last formed phloem with only 'sclerified cells' (black radial line ), x 44. - Fig. 24. Terminalia brasiliensis, bark, transverse sec• tion, stratified, phloem parenchyma alternating with' sclerified cells' (arrows); periderm (P), x 44.

Figs. 25-34. Buchenavia, bark. - 25: B. capitata, transverse section, druses in seriate strands, large druse (centre), polarised light, x 110. - 26: B. rabelloana, transverse section, sieve ele• ment(s), x 440. - 27: B. ochroprumna, transverse section, gelatinous fibres (short arrow) and 'sclerified cells' (long arrow), x 220. - 28: B. macrophylla, macerated, sieve element with sieve plates - compound at one end, simple at the other, x 220. - 29: B. congesta, macerated, phloem fibre, x 44. - 30: Idem, macerated, long and thin 'sclerified cell', x 44. - 31: B. macrophylla, macerated, short and wide-bodied 'sclerified cells', x 44. - 32: Idem, transverse section, 'scleri• fied cells' with thick, polylamellate walls, x 440. - 33: B. parvifolia, radial section, ray with intercellular spaces, x 440. - 34: Idem, phellem cells with large simple pits, x 375.

Figs. 35-40. Buchenavia, bark. - 35: B. capitata, transverse section, periderm with thin-walled phelloderm cells (ph), and stratified phellem (pe) with organic contents in some cells (arrow), x 110. - 36: B. suaveolens, phellem cells with U-shaped wall thickenings (arrows), x 110. - 37: B. ochroprumna, phellem cells with evenly thickened cell walls (long arrow) alternating with one row of very thin-walled cells (short arrow), x 110. - 38: B. grandis, lenticel, x 110. - 39 & 40: B. rabelloana, macerated, branched ray sclereids, x 220. - Figs. 41 & 42. Terminalia, bark, branched ray sclereids (arrows), x 220. - 41: T. amazonia, macerated. - 42: T. brasiliensis, tan• gential section.

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29 30

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90(-140) Jlm, respectively; arranged in regu• Terminalia (Figs. 7, 8,24,41,42) lar, 1-3-seriate tangential bands and accom• panied by few phloem fibres in B. capitata The secondary xylem and phloem of neo• (RBHw 16699), B. grandis (INPAw X- tropical Terminalia (5 species, 8 samples) 2657, Fig. 21), B. macrophylla, B. pallido• examined is essentially identical to that of virens (Fig. 23), B. parvifolia, B. rabello• Buchenavia. Also, the features observed ana, B. suaveolens, B. tomentosa, B. viridi• agree very well with those reported by Van lora; in small groups and of more sporadic Vliet (1979) and thus do not warrant a sepa• occurrence in B. congesta, B. grandis, B. rate description. Differences are few and ochroprumna, B. oxycarpa (INPAw X-89l). largely restricted to quantitative features such Inorganic inclusions: Druses located in as vessel dimensions. Slight differences were chambered parenchyma cells (Fig. 22) which also observed in parenchyma distribution, are arranged in short tangential lines (cross frequency, size and location of crystals, pres• section, Fig. 25), present in all species ex• ence of 'sclerified cells' in bark, etc.; when• cept B. rabelloana; druse-bearing parenchy• ever appropriate, these differences will be ma strands decreasing in frequency with dis• considered in the following discussion. tance from cambium (in B. rabelloana cham• bered parenchyma cells were also observed Discussion but without druses); in the outer, dilated por• The wood anatomy of the species of tion of the phloem very large druses in iso• Buchenavia and neotropical Terminalia ex• lated, inflated parenchyma cells occur in most amined in this study is remarkably homo• species (Fig. 25), absent in B. congesta, B. geneous. However, there are variations in grandis (MGw 2938), B. parvifolia, B. viri• type and frequency of axial parenchyma, lo• diflora; other crystal types and silica not ob• cation of crystals, presence/absence of silica served. and septate fibres, and growth increment bor• Organic, presumably phenolic contents ders. These features may prove to be useful present in parenchymateous tissue of all spe• for identifying species or species groups. cies (Fig. 35). Detienne & Jacquet (1983) describe con• Periderm very consistent in arrangement siderable variation of axial parenchyma for and composition throughout the genus. Radial neotropical Terminalia and use it in their depth varies from 110-830 Jlm; the number species identification key. Axial parenchyma of periderms per radial distance was impos• arrangement is also useful identifying spe• sible to quantify because of interspersed col• cies or species groups in Buchenavia (Table lapsed phloem tissue obscuring the bound• 3). Only B. oxycarpa lacks both marginal and aries. diffuse parenchyma; B. macrophylla, B. Phellem stratified, composed of one to ochroprumna, B. oxycarpa, B. pallidovirens several rows of rectangular cells with either show confluent to banded, only B. viridiflora all walls evenly thickened (Fig. 37) or u• a typically aliform-lozenge parenchyma. shaped thickenings (Fig. 36); transverse Considerable within-species variation in just walls with very large simple pits in some parenchyma is observed in specimens of B. species such as B. capitata (MOw 4154), B. parvifolia and Terminalia amazonia (Table 4). grandis, B. ochroprumna, B. parvifolia (Fig. Thus, although some Buchenavia species ap• 34), B. tomentosa, B. viridiflora; alternating pear to have characteristic parenchyma distri• with usually one, occasionally several rows bution, the same individual pattern(s) also of very thin-walled cells (Fig. 37). Phello• occur(s) in Terminalia, a fact which renders derm poorly developed, composed of few this feature somewhat ambiguous diagnosti• rows (1-6) of thin-walled cells (Fig. 35). cally for species identification within and at• Lenticels observed only in B. capitata, B. tribution to these two taxa. grandis (INPAw X-2657, Fig. 38) and B. Detienne and Jacquet (1983) found locali• parvifolia. Rhytidome persistent in all spe• sation and type of crystals to be of diagnostic cies, encompassing 1-2 periderms and in• significance in Terminalia. Judging from the cluding parts of the collapsed phloem. present study only the occurrence of pris-

Downloaded from Brill.com09/26/2021 12:15:20PM via free access 136 IAWA Bulletin n.s., Vol. 12 (2),1991 I I I I I I I I I I I I I I I I I I 1 1 I 70 90 60 100 120 140 110190 2 160 130 160 160 100 1 140 190 170 140 100 160 210 200 200 220 width vessel ray + + diam. + + + + + + + fibres vessel helical + + + (+) bordo Buchenavia. in +;p +;p +;p +; r, p septate characters +; r +; r +; r +; r +; r +; r +; r +; r +;r +; cp +;cp +; r, p +; r, p +; r, p +; r, p +; r, p wood + + + + + + + + + + + + (+) (+) +; r (+) (+) (+) +; r, p diffuse crystals silica fibres chambered strands. = quantitative and + + + + + + + + + + + + + + + (+) (+) qualitative axial parenchyma; cp = + + + + + + bands selected ray; p of = Parenchyma distribution + + + + + + + + + + + + + + (+) (+) (+) (+) (+) (+) Variation 3. + + + Table lozenge confl. aliform aliform confl. marginal character poorly developed; r = X-2657 nr. INPAw X-2626 Uw 25612 Uw 8044 INPAw X-2017 lNPAw X-3278 lNPAw X-3571 SPW 43 Uw 5801 RBHw 19170 RBHw 16683 lNPAw X-1612 lNPAw X-3091 lNPAw X-6800 lNPAwX-891 Uw 13694 Uw7754 MGw2938 INPAw X-2381 lNPAw Uw686 MGw4154 Uw 32810 RBHw 16699 Coil. absent; ( ) = ? present; - tomentosa suaveolens viridiflora macrophylla grandis sericocarpa rabel/oana amazonia pallidovirens ochroprumna = + B. B. B. B. parvifolia B. B. B.oxycarpa B. B. kleinii B. Bi Species B. congesta B. B. B. capitata

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Table 4. Variation of selected qualitative wood characters in Terminalia L.

Parenchyma distribution Species alifonn alifonn confl. marginal diffuse crystals silica fibres lozenge confl. bands septate

T. amazonia MGw779 + + +; r + BCTw 13254 + +; r + RBHw 12861 + +; r, p + RBHw 18688 + +; r, p + T. argentea RBHw 3540 (+) + + +; cp T. brasiliensis MGw 396 + + +; r, p + T. guianensis MGw3233 + (+) + +; r, p + T. tanibouca MGw 2279 + +;p + =present; - =absent; ( ) =character poorly developed; r =ray; p =axial parenchyma; cp =chambered parenchyma strands.

matic crystals in chambered parenchyma Clearly, the small sample size of only one strands in B. capitata and B. kleinii* appear specimen for most species and considerable to constitute a diagnostic feature. The re• within species variation does not allow defi• maining data on crystals (presence/absence, nite conclusions about the diagnostic signifi• type, location) assembled here for Buche• cance of crystalline inclusions in Buchenavia. navia (Table 3) and Terminalia (Table 4) are Four Buchenavia species (B. capitata ex• ambiguous in that cept Uw 686, B. klein ii, B. oxycarpa, B. a) crystals are present in all but one (B. se• suaveolens) have average diameters of ~ 100 riocarpa, only one sample) of the species J.lm (see Table 3). Van Vliet (1979) and De• examined. This observation agrees with tienne and Jacquet (1983) have found similar Van Vliet's (1979) description of the same values for the latter two species. However, as species, cited there as B. acuminata but vessel diameter is also a function of sample since reduced to synonymy according to diameter (age) no taxonomic significance is Stace (pers. comm., 1989); attributed to this quantitative character. The b) in several species (B. capitata, B. oxycar• same holds true for ray width. Only the pre• pa, B. viridiflora) crystals are present in dominantly biseriate rays in B. tomentosa some, absent in other specimens; might warrant a clear separation from the re• c) crystal size is a continuum from isodiamet• maining species with predominantly uniseri• ric to intermediate (prismatic) and elongate ate rays. (styloids) within and between specimens/ Silica was observed in some Buchenavia species. species, not in Terminalia. The absence of

*) The material of B. kleinii examined by Van Vliet (Lindeman & De Haas 2355 = Uw 13699) originates, according to the collectors, from the same group of trees as the material used here (Lindeman & De Haas 2350 = Uw 13694). As the vouchers of both specimens are sterile, doubts have since been raised as to the correct generic attribution, and its relegation to Ter• minalia was proposed informally (Lindeman, pers. comm.). Despite this uncertainty it was decided to maintain the original designation (B. kleinii) for the benefit of both consistency and comparison with Van Vliet's account.

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silica in Terminalia was previously noted by ture appears impossible. The most remark• Van Vliet (1979, 43 species), Ter Welle able feature of both Buchenavia and Termina• (1976) and Detienne and Jacquet (1983). Sili• lia bark is the regular occurrence (and fre• ca was observed in one sample of B. capitata quency) of the so-called 'sclerified cells' (see (RBHw 16699) and B. ochroprumna (INPAw bark description). This neutral term was cho• X-6800), in B. macrophylla, and B. serio• sen provisionally because of the unknown carpa (Table 3). Van Vliet (1979) observed ontogeny of these cells. Though often but by silica in B. seriocarpa, Ter Welle (1976) and no means always resembling fibres in general Detienne and Jacquet (1983) report silica ab• appearance (shape, dimensions), these ele• sent in Buchenavia. Some authors consider ments differ from normal phloem fibres by silica to be an important and consistent diag• the sclerified, i.e. polylamellate, pitted and nostic character at the various taxonomic lev• very thick walls, and the ovoid transverse els (Richter 1981a, 1985; Ter Welle 1976; section (larger diameter radially). This cell Ter Welle & Detienne 1986). The inconsis• type, already observed by several authors in tency of its occurrence in Buchenavia species the bark of various taxa, is known under an (vide B. capitata, B. ochroprumna) poses array of different denominations: 'fibrous serious difficulties of interpretation: In B. sclerotic cells' (Shimakura 1936), 'fibre• capitata the vouchered specimen RBHw sclereids' (Holdheide 1951; Santos 1960), 16699 is the only one with silica while, ac• 'fibres' (Chattaway 1955c, 1959), 'sclereids' cording to Benoist (1931, cited by VanVliet (Srivastava 1963), 'sclerotic phloem fibres' 1979), Detienne and Jacquet (1983) and Ter (Parameswaran 1980). Only a thorough on• Welle (pers. comm., cited by Van Vliet 1979), togenetic study will reveal whether they are silica is absent in this particular species. Con• of parenchymatous origin or derived directly sequently, these observations raise some from fusiform cambial initials, and will give doubt as to the correct identification of the rise to an appropriate terminology. Perhaps specimens labeled B. capitata and, for that even both forms develop in different taxa. matter, B. ochroprumna. 'Sclerified cells' are present in all Buche• Septate fibres were observed in one sam• navia and Terminalia species studied with ple of B. capitata (Uw 686), in B. oxycarpa varying frequency between and within indi• and B. seriocarpa, reported for the latter also vidual species. They may already occur at the by Van Vliet (1979). Detienne and Jacquet age of two years (Holdheide 1951), and their (1983) report non-septate fibres in B. capitata proportion increases with age (Chattaway and B. oxycarpa. Yet, the within species in• 1955c). In B. pallidovirens, the scleren• consistency (e.g. B. capitata) does not at chymatic tissue of the younger phloem con• present make septate fibres a viable taxon• sists predominantly of phloem fibres, while omic character. Within Terminalia amazonia in the older phloem strata only 'sclerified (Table 4) septate fibres are of constant occur• cells' occur. Hence, frequency variation can• rence. not be used as a diagnostic feature for species Helical thickenings were found by Van differentiation, even when the exact age of Vliet (1979) in vessel elements and axial pa• the material studied is known. renchyma of B. grandis (syn. B. huberi); an Phloem fibres are arranged in continuous additional species (B. suaveolens, narrow tangential bands in some species, and in tan• vessel elements only) is reported here (Table gentially arranged groups in others. Richter 3). Distinctly bordered pits in fibre walls, not (1981a) observed similar differences in scle• previously described for Buchenavia, were reid arrangement in Cinnamomum (Laura• observed occasionally in B. amazonia, B. ceae) and used this as one of several argu• capitata (RBHw 16699), B. congesta, B. par• ments for separating different species groups vifalia and B. suavealens. in this genus. However, the little material of Buchenavia studied here makes attaching The bark anatomy of the taxa studied is as diagnostic value to this character premature. homogeneous as the wood anatomy. Separat• Many of the observations made by Roth ing Buchenavia and Terminalia by bark struc- (1969, 1981) on the bark structure of one

Downloaded from Brill.com09/26/2021 12:15:20PM via free access Alfonso & Richter - Wood and bark anatomy of Buchenavia 139 species of Buchenavia and two species of surprising as Exell & Stace (1963, 1966) Terminalia cannot be corroborated by the found the morphology of vegetative organs present study: Not only Terminalia (Roth and leaf anatomy (Stace, pers. comm., 1989) 1969) but also Buchenavia bark features to be similar and indicative of a close rela• small dtuses in crystalliferous parenchyma tionship between the two genera. strands in the non-collapsed phloem, and There appears to be considerable intra• large druses in inflated cells in the collapsed specific variation and overlap in wood ana• phloem; in both genera wall thickenings of tomical characters within Buchenavia. Con• phellem cells are of the same type (Le. 'U'• sequently, different species cannot always be shaped) rather than 'u' -shaped in Terminalia reliably distinguished using wood and/or and inverted in Buchenavia as reported by bark anatomy at the present state of knowl• Roth (1969); ray dilatation and ray sc1ereids edge. This contrasts Van Vliet's (1979) con• were observed in Terminalia and Buchenavia tention that species of Buchenavia are distin• bark while Roth (1969,1981) lists these fea• guishable using wood anatomical characters. tures as absent in both taxa. Studies of more vouchered material per spe• Moeller (1882) described axial parenchy• cies are necessary to determine which, if any, ma cells with large and simple pits for Termi• wood and bark features are diagnostic of nalia; the same type of wall pitting can be ob• Buchenavia species. Silica and septate fibres served in parenchymatous phellem cells of (wood) and fibre arrangement (bark) appear Buchenavia. to be the most promising features for further Branched ray sc1ereids observed in B. study. congesta, B. rabelloana, T. amazonia and T. brasiliensis constitute an interesting phenom• Acknowledgements enon. The rather bizarre looking sc1ereids The authors are indebted to Antonio C.F. differentiate from ray cells in direct contact Barbosa and Claudia Muche for the skilfull with the axially oriented 'sc1erified cells'. preparation of wood and bark slides, and to They differ from regular ray sclereids by Christina Waitkus for her valuable assistance forming slender and pointed branches that with the illustrations. The study was spon• grow into and gradually fill intercellular sored by the Fundar;:ao de Amparo it Pesquisa spaces. The sclerification of ray tissue ele• do Estado de Sao Paulo (FAPESP, Brazil) ments in such contact areas has been reported and the Deutsche Akademische Austausch• by many authors (Moeller 1882; Holdheide dienst (DAAD, Federal Republic of Ger• 1951; Chattaway 1955c, 1959; Zahur 1959; many) through grants to the first author. Parameswaran & Liese 1970; Richter 1981, Their support is gratefully acknowledged. 1990; Trockenbrodt 1989). Branched scle• reids differentiating apparently from axial Rererences parenchyma are described and illustrated Alfonso, V.A. 1983. Caracterizar;:ao anato• in Esau (1969). However, a reference to mica da madeira e da casca das principais branched ray sc1ereids has not been found in especies de Eucalyptus do Estado de Sao the literature consulted. Paulo. Silvicultura 8: 720-725. Finally, there is little variation within and Alwan Al-Mayah, A.R.A. & C.A. Stace. between Buchenavia species or between 1985. Resurrection of two Eighteenth Buchenavia and Terminalia in stratification of Century names in American Combreta• non-collapsed and collapsed phloem, type of ceae. Studies on the Flora of the Guianas sieve element and sieve plate, ray dilatation, 7. Nord. J. Bot. 5: 447-449. structure of periderm and rhytidome. Chattaway, M.M. 1955a. The anatomy of bark. II. Oil glands in Eucalyptus species. Conclusions Austr. J. Bot. 3: 23-27. Wood and bark anatomy alone do not Chattaway, M.M. 1955b. The anatomy of allow distinguishing Buchenavia from neo• bark. III. Enlarged fibres in the blood• tropical Terminalia. The similarity in wood woods (Eucalyptus spp.). Austr. J. Bot. and bark structure of these two genera is not 3: 28-38.

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Chattaway, M.M. 1955c. The anatomy of Moeller, J. 1882. Anatomie der Baumrinden. bark. IV. Radially elongated cells in the J. Springer, Berlin. phelloderm of species of Eucalyptus. Outer, R.W. den & J.M.Fundter. 1976. The Austr. J. Bot. 3: 39-47. secondary phloem of some Combretaceae Chattaway, M.M. 1959. The anatomy of and the systematic position of Strepho• bark. VII. Species of Eugenia. Trop. nema pseudocola A. Chev .. Acta Bot. Woods. 111: 1-14. Neerl. 25: 481-493. Detienne, P. & P. Jacquet. 1983. Atlas Parameswaran, N. 1980. Some remarks on d'identification des bois de l' Amazonie et the nomenclature of fibres, sclereids and des regions voisines. Centre Technique fibre-sclereids in the secondary phloem of Forestier Tropical, Nogent-sur-Marne. trees. IAWA Bull. n.s. 1: 130-132. Esau, K. 1969. The phloem. Handbuch der Parameswaran, N. & W. Liese. 1970. Mi• Pflanzenanatomie. V /2. Gebruder Born• kroskopie der Rinde tropischer Holzarten. traeger, Berlin, Stuttgart. In: Handbuch der Mikroskopie in der Esau, K. & V.I. Cheadle. 1984. Anatomy of Technik (H. Freund, ed.) V /1, 2nd edi• the secondary phloem in Winteraceae. tion: 227-306. Umschau Verlag, Frank• IAWA Bull. n.s. 5: 13-43. furt/Main. Exell, A.W. & C.A.Stace. 1963. A revision Richter, H.G. 1981a. Anatomie des Sekun• of the genera Buchenavia and Ramatuela. diiren Xylems und der Rinde der Laura• Bull. Brit. Mus. (Nat. Hist.), Bot. 3: 3-46. ceae. Sonderbande NatuTW. Ver. Ham• Exell, A.W. & C.A.Stace. 1966. Revision burg 5. Paul Parey Verlag, Hamburg, of the Combretaceae. Bull. Soc. Brot. 2 Berlin. (40): 5-25. Richter, H.G. 1981b. Wood and bark anat• Freese, F. 1967. Elementary statistical meth• omy of Lauraceae. I. Aniba Aublet. ods for foresters. USDA Forest Service. IAWA Bull. n.s. 2: 79-87. Agriculture Handbook 317. Washington. Richter, H.G. 1985. Wood and bark anat• Holdheide, W. 1951. Anatomie mitteleuropa• omy of Lauraceae. II. Licaria Aublet. ischer Geholzrinden. In: Handbuch der IAWA Bull. n.s. 6: 187-199. Mikroskopie in der Technik (H. Freund, Richter, H.G. 1990. Wood and bark anat• ed.) V/l: 193-367. Umschau Verlag, omy of Lauraceae. III. Aspidostemon Frankfurt/Main. Rower & Richter. IAWA Bull. n.s. 11: IAWA 1989. IAWA list of microscopic fea• 47-56. tures for hardwood identification. (E.A. Roth, 1. 1969. Estructura anat6mica de la Wheeler, P. Baas & P.E. Gasson, eds.). corteza de algunas especies arb6reas de IAWA Bull. n.s. 10: 219-332. Combretaceae. Acta Bot. Venez. 4: 227- Karstedt, P. & N. Parameswaran. 1976. Bei• 239. trage zur Anatomie und Systematik der at· Roth, I. 1981. Structural patterns of tropical lantischen Rhizophora-Arten. Bot. Jahrb. barks. Handbuch der Pflanzenanatomie. Syst. 9: 317-338. IX/3. Gebr. Borntraeger, Berlin, Stutt• Kruse, J. 1977. Mikrotechnologische und gart. biologische Untersuchungen an Rinden• Rupp, P. 1964. Polyglykol als Einbettungs• platten. Dissertation, Fachbereich Biolo• medium zum Schneiden botanischer Pra• gie, Universitat Hamburg. Unpublished. parate. Mikrokosmos 53: 123-128. Lindeman, J.J. & A.M.W.Mennega. 1963. Santos, C.F.O. 1960. Estudo do escleren• Bomenboek voor Suriname. Kemink & quima no floema secundarlo de algumas Zoon, Utrecht. dicotiled6neas lenhosas. Rev. Agricult. Mainieri, C., J.P. Chimelo & V.A. Alfonso. 35: 199-206. 1983. Manual de identifica9ao das prin• Shimakura, M. 1936. On the sclerenchyma cipais madeiras comerciais brasileiras. of bast in conifers. Bot. Mag. (Tokyo) 50: Ed. Promocet. Publica9lio IPT 1226. Slio 206-213. Paulo.

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Srivastava, L.M. 1963. Secondary phloem Welle, B.J.H. ter. 1976. Silica grains in in the Pinaceae. Univ. Calif. Publ. Bot. woody plants of the neotropics, especially 36: 1-142. Surinam. In: Wood structure in biologi• Trockenbrodt, M. 1989. Entwicklung, Varia• cal and technological research. (P. Baas, bilitat und taxonomische Bedeutung der A.J. Bolton & D.M.Cading, eds.). Lei• Rindenstrukturen von Quercus robur L., den Botanical Series.3: 107-142. Ulmus glabra Mill., Populus tremula L. Welle, B.J.H. ter & P. Detienne. 1986. und Betula pendula Roth. Dissertation, Wood and Timber. In: Flora of the Gui• Fachbereich Biologie, Universitiit Ham• anas. 85. Chrysobalanaceae (A.R.A. burg. Unpublished. Gorts-van Rijn, ed.). Koeltz Scientific Trockenbrodt, M. 1990. Survey and discus• Books, Germany. sion of the terminology used in bark anat• Wilhelm, G. 1975. Uber den mikrobiellen omy. IAWA Bull. n.s. 11: 141-166. Abbau der Rinde von Fagus sylvatica und Trockenbrodt, M. & N. Parameswaran. Picea abies. Dissertation, Fachbereich 1986. A contribution to the taxonomy of Biologie, Universitat Hamburg. Unpub• the genus Inga Scop. (Mimosaceae) based lished. on the anatomy of the secondary phloem. Zahur, M.S. 1959. Comparative study of IAWA Bull. n.s. 7: 62-71. secondary phloem of 423 species of Vliet, G.J.C.M. van. 1979. Wood anatomy woody dicotyledons belonging to 85 fam• of the Combretaceae. Blumea 25: 141- ilies. Cornell Univ. Agr. Expt. Station 223. Memoir 358.

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