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Wood anatomy of tribe Detarieae and comparison with tribe Caesalpinieae(Legum inosae, Caesalpinioideae) in VenezuelaJos Luis Melandri'- - & Narcisana Espinoza de Pema'1, Laboratorio de Anatomia de Maderas, Facultad de Ciencias Forestales y Am bientales. Universidad de Los Ande s.

Mrida. Venezuela, m elandriijula.ve, nepemia(iiula.ve2, Author for correspondence: melandri(uia.ve

Received 28-11-2008. Corrected 06-v n-2 008 , Accepted 04-Vin -2008 ,Abstract: We studied the wood anatomy of 29 species belonging to 10 genera of the tribe Detarieae, subfam-ily Caesalpinioideae and compare them witb tribe Caesalpinieae. Detarieae is the largest of four tribes ofCaesalpinioideae, witb 84 genera, only eleven occur in Venezuela with species of timber importance. The speci-mens were collected in Venezuela and include wood samples from thecolleclionof the Laboratorio dc Anatomade Maderas de la Facultad de Ciencias Forestales y Ambientales de la Universidad de Los Andes. Venezuela,and of Ihe Forest Products Laboratory of the liSDA Forest Service in Madison. Wisconsin. USA, Ibe Icrminol-ogy and metbodology used followed the lAWA List of Microscopic Features for Hardwood Identification of thelAWA Committee, 1989, Measurements from eacb specimen were averaged (vessel diameters, vessel elementlengtbs. intervessels pit size, fibre lengths and ray height). The species of Detarieae can be separated using acombination of diagnostic features. Wood characters that provide the most important diagnosis and may be usedin systemalics of Ueiarieae include; intercellular axial canals, rays heterocellular. rays exclusively or predomi-nantly uniseriate, prismatic crystals common in ray cells, irregular storied structure and fibre wall thickness.For comparative anatomy between Detarieae and Caesalpinieae: intercellular axial canals, heterocellular rays,rays exchisively or predominantly uniseriate. prismatic crystals common in ray cells (in Detarieae) and regularstoried structure, fibres septate, fibre wall tbick or very thick, rays honiocellular. multiscriate rays and silicabodies (in Caesalpinieae), Axial parenchyma is typically a good diagnostic feature for l.eguminosae, but not forDetarieae and Caesalpinieae comparisons. Rev. Biol, Trop, 57 (1-2): 303-319. Epub 2009 June 30,Key words: l.eguminosae, Caesalpinioideae. Detarieae, Caesalpinieae, wood anatomy, identification.

The legume tribe Detarieae is the largest offour tribes of subfamily Caesalpinioideae, com -prises 84 genera incltiding approximately halfof the genera of subfamily Caesalpinioideae.Most of these genera occur in tropical Africa,with a less diverse representation in tropi-cal America and Asia (Bameby et al. 1998.Bruneau et al. 2000, Herendeen 2000, Gassonet ai. 2003. Herendeen et al. 2003,). Only elev-en genera of the Detarieae trihe are distributedin Venezuela (B ameb y et ai 1998, Aristeguieta2003). Wood anatomy study of the tribes, gen-era or species in the subfamily Caesa lpinioideae

characters for taxonomic, systematic andproperties of commercial timbers purposesinclude Reinders-Gouwentak (1953), Baretta-Kuipers (1981), Ranjani and Krishnamurthy(1988), Dtienne and Welle (1989), Wheelerand Baas (1992). Nardi and Edimann (1992),Gasson (1994), (1996), (1999), Hhn (1999),Herendeen (2000), Miller and Dtienne (2001 ),Chauhan and Rao (2003), Gasson et ai (2003),Espinoza de Pemia and Melandri (2006a,b).

Some genera of the tribe Detarieae yieldcommercial timbers, particularly Copaifera,Eperua. Hymenaea. and Peltogyne. this last

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that is widely appreciated for its unusual colorand resistance to insects (JUNAC 1981, INIA1 9 9 6 , Bameby et al 1998, Aristeguieta 2003 ).Others genera such as Brownea and Dicymbeare important ornamental trees, in gardens,city avenues and parks (Bameby el al. 1998,Aristeguieta 2003). Medicinal plants such asCopaifera are of high industrial value for theirgum or balsam of Copaiba for medical treat-ments, manufacturing of varnishes and shel-lacs, and as a fixative of fragrances in soapsand perfiimes. The Elizahetha genus yields ahallucinogenic drug, while Hymenaea genus isused for food and forage, medical treatments.,canoe building and manufacturing of var-nishes (Mabberley 1997, Bameby et al. 1998,Aristeguieta 2003). Some Dicymbe^ Eperua,Heterostemom and Macrohbium species areendemic to the Venezuelan Guayana (Bamebyetal. 1998).

This paper provides information aboutwood anatomy of native genera of the tribeDetarieae, that has not been adequately studied,and t o compare them with t h e tribe Caesalpinieae(Espinoza de Pemia and Melandri 2006b). Themicroscopic wood anatomy of both tribes wasstudied because of its great importance inthe timber industry and the complexity of itsanatomy. The anatomical descriptions providetools for the identification of the genera andgroups within the tribe.MATERIALS AND METHODS

Microscope slides from 70 wood samplesrepresen ting 2 8 species -om 1 0 genera the tribeDetarieae were examined (only II genera ofthe tribe are distributed in Venezuela: Brownea,Copaifera, Crudia. Cynometra, Dicymbe.Elizabetha, Eperua, Heterostemon, Hymenaea.Macrohbium, Peltogyne). The majority of thespecimens were collected in Venezuela andincludes specimens from the wood collection atthe Laboratorio de Anatomia de Maderas de laFacultad de Ciencias Forestales y Ambientalesde la Universidad de Los Andes, Mrida,Venezuela (MERw) and at the USDA ForestService, Forest Products Laboratory, Madison..

We followed the List of MicroscopicFeatures for Hardwood Identification (IAWACommittee 1989) for terminology and method-ology. The following characters were recordedfor each specimen studied: presence/absenceof growth rings, porosity, vessels distribution,intervessels pit size, vestured pits, fibre wailthickness, septate fibres, axial p arenchyma pat-tems, number ofthe cells per axial parenchymastrand, ray size in height and width in cells,composition ray cell, storeyed structure, pris-matic crystals, silica bodies, and axial canal,among others characters. Generic descriptionsfollow in alphabetical order and features notlisted in the generic descriptions are eitherabsent or do not apply. For vessel diameters,vessel element lengths, fibre lengths and rayheight 25 measurements were taken from eachspecimen and averaged. The measurements areaccurate only to the 10 um level, and are report-ed accordingly. The values reported [e.g. 30(50-110) 150 (.im], are minimum value, rangeof averages, and maximum value. For otherquantitative values the most frequent range isreported. Photomicrographs were taken using afilm camera with a light microscope.

RESULTS

Generic descript ionsBrownea Jacq. - Fig. 1 & 2

Growth rings distinct, marked by mar-ginal parenchyma bands and/or thick-walledfibres. Diffuse porous. Vessels solitary and inradial multiples of 2-4, 3-17 per mm-, 40(70-128)160 um in diameter, 130(209-372)550 fimin element length. Simple perforation plates.Alternate intervessel pits circular or oval andpolygonal, minute to small, 3-6 [xm in diameter.Vessel-ray pits with distinct borders, similarto intervessel pits in size and shape, pits ves-tured. Brown gum-like deposits in vessels.Fibres non-septate, medium- to thick-walled,700(908-14 36)1680 fim in length. Axial paren-

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Fig. M . (1 & 2): Brownea species.- 1: B. coccnea, aliform, confluent parenchyma.- 2: B. grandiceps, rays heterocellularand prismatic crystals in chambered ray cells.- 3 & 4: Copaifera species.- 3: C. offidnalis. inlercelliilar axial canals (arrow)in long tangential lines, brown gum-like deposits in vessels.- 4: C, pubtflora, homoceliular rays composed of typicallyprocumbent cells. Scale bars = 250 jim.

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confluent in all species, aliform parenchymaof the lozenge type. Banded parenchyma ofmore than three cells wide in B. grandicepsand B. macrophyHa. Marginal parenchymabands present. Axial parenchyma 3-6 cells perparenchyma strand. Rays heterocellular with1 to 2 rows of upright and/or square marginalcells, 9-20 per mm, exclusively ( v w B . coccnea)or mostly uniseriate (in B. grandiceps. B.macropiiyHa), 120(252-340)600 \im in height.Prismatic crystals common in ray cells (bothupright and/or square and procumbent cells)and occasionally in short chains in axial paren-chyma cells., one crystal per cell or chamber.

Material studied: 7 specimens, B . coccin-ea Jacq., MADw 32265, B. grandiceps Jacq.,MADw 4835, MADw 4836, MADw U163,MADw 14229, MADw 32266, B. macrophyHaHort, ex M ast., ME Rw 1277.

Copaifera L. - Fig. 3 & 4Growth rings distinct, marked by margin-

al parenchyma bands. Diffuse porous. Vesselssolitary and in radial multiples of 2-4, 3-8 permm -, 90(100-190)210 }im in diameter, 100(253-348)480 jxm in element length. Simple perfora-tion plates. Alternate intervesse! pits small tomedium, 6-10 fxm in diameter. Vessel-ray pitswith distinct borders, similar to intervessel pitsin size and shape, pits vestured. Brown gum-like deposits in vessels. Fibres non-septate,medium-walled, 800(908-1264)1570 um inlength. Axial parenchyma vasicentric, aliformto confluent, aliform parenchyma of the loz-enge type. Marginal parenchyma bands present.Axial parenchyma 2-4 cells per parenchymastrand. Rays mostly homocellular with typicallyprocumbent cells to heterocellular, with 1 rowof upright and/or square marginal cells, 6-8per mm , 1 to 3 occasionally up to 4 cells wide,commonly less than 1 mm, to slightly morethan 1 mm: 220(250-691)1050 ^im in height.Prismatic crystals common in chains in axialparenchyma cells, one crystal per cell or cham-ber. Intercellular canals axial in long tangential

Material studied: 13 specimens, C. offl-cinaHs (Jacq.) L., ME Rw 1533, MER w 1780,MERw 3297, MERw 5584, MERw 5585,MERw 5586, MERw 5587, MERw 5588, C.pubiora Benth. , MERw 4541, MERw 4542,MERw 4543, MERw 4544, MERw 4545.

Note: Metcalfe and Chalk ( 1950), Corothie(1967). Dtienne et al. (1982), Richter andDallwitz (2000) characterize the rays ofCopaifera as homocellular (homogeneous),but Baretta-Kuipers (1981) described them asheterocellular. Additionally, JUNAC (1981)described homocellular rays C. ofcinas andheterocellular rays in C. pubiora. Mainieriand Chimelo (1989) report for C. cf. langs-dorffti homocellular rays with tendency toheterocellular, and for C. cf. reticulate, het-erocellular rays. Conceming fibres characterJUNAC ( 1981), reports fibres partially septate,however, this feature was not observed in ourspecimens.

Cynometra L. - Eig. 5 & 6Growth rings indistinct marked by mar-

ginal parenchyma bands. Diffuse porous.Vessels solitary and in radial multiples of 2-4;occasionally clusters; 9-16 per mm-; 35(98-104)145 um in diameter, 150(264-331)450 \xmin element length. Simple perforation plates.Alternate intervessel pits minute to small, 3-7|im in diameter. Vessel-ray pits with distinctborders, similar to intervessel pits in size andshape; pits vestured. Brown gum-like depositsin vessels. Fibres non-septate, thick-walled,1O5O{1375-16O5)I95O jxm in length. Axialparenchyma aliform to confluent, aliformparenchyma of the lozenge type, occasion-ally unilateral. Banded parenchyma mostly inbands more than three cells wide. Marginalparenchyma bands occasionally present. Axialparenchyma 2-4 cells per parenchyma strand.Rays heterocellular with 1 to 2 rows of uprightand/or square marginal cells, rarely up to 3marginal rows, 8-16 per mm, 1 to 4 cells w ide,220(381-600)850 ^m in height. Prismaticcrystals common in upright and/or square

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Fig. 5-8.- 5 & 6: Cynometra spruceana - 5 : hands of parenchyma m ore than three cells w id e. -6 : rays heterocellular. 1-2rows of upright and/or square cells.- 7 & X: Dicymbe bernardii- 7: growth rings distinct, aliform parenchyma- 8: rayshclcroccllular, 1-2 rows of upri^t and/or square cells. Scale hars = 250 im.

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axial parenchyma cells, one crystal per cell orchamber.

Matenal studied: 3 specimens, C, spru-ceana Benth.. MADw 32048; C sp. L. MERw2 1 5 2 , MERw 2233.

Note: Metcalfe & Chalk (1950) forCynometra (except C ramiflora) and Kribs(1968) for C. aiexandri described the raysas homogeneous, but Baretta-Kuipers (1981);Richter & Dallwitz (2000) for Cynometraand Dtienne et al. (1982) for C. hostman-niana and C parvifolia described the rays asheterocellular.

Dicymbe Spruce ex Benth. - Fig. 7 & 8Growth rings distinct, marked by mar-ginal parenchyma bands and/or thick-walled

fibres. Diffuse porous. Vessels solitary andin radial multiples of 2-5; sometimes up to9; occasionally some clusters, 4-6 per mm-;80(121-150)210 ^m in diameter, 200(307-327)520 p,m in element length. Simple perfora-tion plates. Alternate intervessel pits small, 4-6[xm in diameter. Vessel-ray pits with distinctborders, similar to intervessel pits in size andshape; pits vestured. Brown gum-like depos-its in vessels. Fibres non-septate, thick- tovery thick-walled, 800(918-1600)2000 ^m inlength. Axial parencbyma mostly aliform,aliform parenchyma of the lozenge type, vasi-centric, confluent. Marginal parenchyma bandsabsent in MERw 239 and present in MADw31798. Axial parenchyma 3-4 cells per paren-chyma strand. Rays mostly homocellular withtypically procumbent cells to heterocellular,with 1-4 rows of upright and/or square margin-al cells, 8-14 per mm, mostly 1 to 2 cells wide(occasionally 3 cells wide), 100(275-706)930[xm in height. Prismatic crystals occasionallypresent in upright and/or square ray cells andin short chains in axial parenchyma cells, oneciystal per cell or chamber.

Material studied: 2 specimens, D. ber-nardii R.S Cowan, MERw 239; D. sp. MAD w31798.

Note: Metcalfe & Chalk (1950) character-

Baretta-Kuipers (1981) described the rays asheterocellular.Elizabetha Schomburgk ex Benth. - Fig. 9 & 10

Growth rings distinct, marked by mar-ginal parenchyma bands and/or thick-walledfibres. Diffuse porous. Vessels solitary andin radial multiples of 2-3, sometimes up to5 , occasionally some clusters, 5-10 per mm-,70(110-130)150 ^m in diameter, 200(320-355)490 \ i .m in element length. Simple perfora-tion plates. Alternate intervesse! pits small, 4-5|xm in diameter. Vessel-ray pits with distinctborders, similar to intervessel pits in size andshape, pits vestured. Brown gum-like depos-its in vessels. Fibres non-septate, medium-to thick-walled. 950(1240-1365)1580 ^m inlength. Axial parenchyma aliform to conflu-ent, aliform parenchyma of the lozenge type,occasionally vasicentric. Banded parenchymamostly in bands 3-5 cells wide. Marginalparenchyma bands present. Axial parenchyma3-4 cells per parenchyma strand. Hays het-erocellular with I to 2 rows of upright and/or square marginal cells, 7-10 per mm, exclu-sively or mostly uniseriate. 110(220-230)420^m in height. Prismatic crystals common inprocumbent ray cells (occasionally in uprightand/or square) and occasionally in short chainsin axial parenchyma cells, one crystal per cellor chamber.

Material studied: 2 specimens, E. prin-ceps Schomburgk ex Benth., MADw 31823, E.macrostachya Benth. , MERw 5275.

Eperua Aublet. - Fig. 11 & 12Growth rings distinct marked by marginalparenchyma bands. Diffuse porous. Vesselssolitary and in radial multiples of 2-3, some-times up to 8, occasionally some clusters,3-13 per mm% 70(155-235) 300 ^m in diam-eter, 200(364-436) 600 (xm in element length.Simple perforation plates. Alternate intervesselpits circular or oval and poligonal, small to

medium, 4-8 \x\t\ in diameter. Vessel-ray pitswith distinct borders, similar to inter\essel

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Kig. y- l2 .- 9& 10: FHzabethaprinceps.-'i: growth rings distinct, marked by thick-walled fibres and aliform parenchym a.-1 0 ; rays exclusively uniscriale.- H & 12: F.perua species- II : E eucantha, intercellular axial canals (arrow), in longtangential lines and connecting or imm ersed in bands ol marginal parenchym a.- 1 2 : . purpuiva, rays mostly 3-4 wide cells.Scale bars = 250 um.

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gum-like deposits in vessels. Fibres non-sep-tate, medium-walled, 970(1104-1459)1600 ^imin length. Axial parenchyma scanty, vasi-centric thin, sometimes diffijse-in-aggregates.Marginal parenchyma bands present. Axialparenchyma 3-4 cells per parenchyma strand.Rays heterocellular with 1 to 4 rows of uprightand/or square m arginal cells, 5-8 per mm , 1 to4 cells wide. 180(200-510)750 um in height.Prismatic crystals common in long chains inaxial parenchyma cells, one crystal per cell orchamber. Intercellular axial canals in longtangential lines and coimecting or immersed inbands of marginal parenchyma, occasionallydiffiise (. purpurea), 50-150 ^im in diameter.

Material studied: 4 specimens, E. gran-diflora (Auhl.) Benth., MERw 200, E. leucan-tha Benth., MERw 194, E. purpurea Benth.,MERw 5273, MERw 5276.

Heterostemon Desf. - Fig. 13 & 14Growth rings distinct, marked by margin-

al parenchyma hands and thick-walled fibres.Diffuse porous. Vessels solitary and radial mul-tiples of 2-3, sometimes up to 5, occasionallysome clusters, 7-15 per mm-, 50(73-98)140 ^min diameter, 160(264-322)430 \im in elementlength. Simple perforation plates. Alternateintervessel pits circular or oval of them shapepolygon al, minute to small, 3-6 \sm in diameter.Vessel-ray pits with distinct borders, similarto intervessel pits in size and shape, pits ves-tured. Brown gum-like deposits in vessels.Fibres non-septate, medium- to thick-walled,700(740-949)1240 (xm in length. Axial paren-chyma vasicentric, aliform to confluent, ali-form parenchyma of the lozenge type. Bandedparenchyma occasionally more than three cellswide in H. conjugatus. Marginal parenchymabands present. Axial parenchyma 2-4 cells perparenchyma strand. Rays heterocellular with1 to 4 rows of upright and/or square marginalcells, occasionally 5 in H . conjugatus, 7-17 permm , exclusively or mostly uniseriate, 170(250-517)640 Jim in height. Prismatic crystals notcommon, mostly present in procumbent ray

Material studied: 6 specimens, H. cau-liflorus Pittier, MADw 31929, MADw 31931,H. conjugatus Spruce ex Benth., MERw 5283,MADw 31932, MADw 31933, H. mimosoidesDesf., MADW 31934.

Hymenaea L. - Fig. 15 & 16Growth rings distinct, marked by mar-

ginal parenchyma hands and beside verythick-walled fibres in H. oblongifolia. Diffuseporous. Vessels solitary and radial multiples of2-4, occasionally some clusters, 2-3 per mm-,90(132-202)300 ^m in diameter, 160(238-354)520 ^im in element length. Simple perfora-tion plates. Alternate intervessel pits circularor oval, of them shape polygonal, small, 4-7\\m in diameter. Vessel-ray pits with distinctborders, similar to intervessel pits in size andshape, pits vestured. Reddish brown gum-likedeposits in vessels. Fibres non-septate, thick-to very thick-wa lled, 1000(134 0-1525)18 90im in length. Axial parenchyma mostly ali-form to confluent, alifonn parenchyma ofthe lozenge type. Banded parenchyma oftnore than three cells wide in H. courbariland H. oblongifolia. Marginal parenchymabands present. Axial parenchyma 2-4 cellsper parenchyma strand. Rays homoceliularwith typically procumbent cells (marginal cellssometimes slightly enlarged), 3-8 per mm, 1 to6 cells wide, 110(192-450)650 pim in height.Prismatic crystals common in long chains inchambered axial parenchyma cells, one crystalper cell or chamber.

Material studied: 11 specimens, H. cour-baril L., MERw 1507, MERw 3394, MERw4564, MERw 4565, MERw 4566, MERw4567, MERw 2489, H. oblongifoHa Hubcr,MERw 2087, MERw 2533, MERw 2648, H.pani/blia Huber, MERw 2053.

Macrolobium Schreber. - Fig. 17 & 18Growth rings distinct, marked by marginal

parenchyma bands and/or thick-walled fibres.Diffuse porous. Vessels solitary and radialmultiples of 2 - 3 , sometimes up to 8, occasion-

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Fig. 13-16.- 13 & 14: Heterostemon species- 13: H. cauliflorus, rays exclusively uniseriale.- 14: //. conjugatus. rays hel-erocellular, 2-4 rows of upri;Jit and/or square cells.- 15 & 16: Hymenaea cowbaril- 15: aliform, confluent parenchyma andmarginal banded.- 16: rays mostly 1-6 wide cells. Scale bars = 250 um.

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Fig. 17-20.- 17 & 18: Macrolobium species- 17: M. angustifoUum. growth rings distinct, marked by thick-walled fibres,vasicentric and aliform parenchyma.- 18: M molle, rays mostly uniseriate.- 19 & 20: Peltog\'ne species.- 19: P. panicuUita.alifonn to confluent, unilateral parenchyma and banded parenchym a in narrow bands or lines, up to three cells wide.- 20: P.porphyrocardia, rays homocellular, composed of ^pically procumbent cells. Scale bars = 250 xm.

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230 um in diameter, 133 (236-439) 590 \imin element length. Simple perforation plates.Altemate intervessel pits circular or oval andpolygonal, small, 5-8 |,im in diameter. Vessel-raypits wiih distinct borders, similar to intervesselpits in size and shape, pits vestured. Brown gum-like deposits in vessels. Fibres non-septate,thin-walled to thick-walled, 670(817-1765)2350|im in length. Axial parenchyma vasicentric.aliform to confluent, aliform parenchyma ofthe lozenge type. Banded parenchyma of 2-4cells wide. Marginal parenchyma bands present.Axial parenchyma 2-5 cells per parenehymastrand. Rays heterocellular with 1 to 2 rows ofupright and/or square marginal cells (sometimesto 3), 9-20 per mm,, exclusively or mostly uni-seriate in M gracile, M. iimbatum, M. molleand M. rubrum, uniseriate and biseriate in M.acaciifolium. M. angustifbiium. M. multijugimiand M. punctatum. 100(220-390)650 \km inheight. Storied structure not observed,, only raysirregularly storied (rays in echelon) in M. molle.Prismatic crystals common in ray cells (bothupright and/or square ray cells and procumbentcells) and occasionally in short chains in axialparenchyma cells in AY. angustifolium. M. grac-ile. M. iimhatum, M. molie. M. multijugum andM. punctatum. Prismatic crystals common inaxial parenchyma cells and sporadic in ray cellsin M acaciifolium and M. rubrum. one to occa-sionally two crystals per cell or chamber.

Material studied: 15 specimens, M.acaciifolium (Benth.) Benth., MERw 2084,MERw 3374, MADw 14917, M. angustifo-iium (Benth.) Cowan, MADw 31604, MADw31617, MADw 31620, M. gracile Spruce exBenth., MADw 31611, M Iimbatum Spruceex Benth., MERw 2050, M. moile (Benth.)Cowan, MADw 14912. MADw 31613, MADw31614, M. multijugum (DC.) Benth.. MADw31605. MADw 31612. M. punctaium Spruceex Benth., MERw 2021, M. rubrum R.SCowan. MADw 22398.

Peitogyne Vogel. - Fig. 19 & 20Growth rings distinct, marked by marginal

parenchyma bands in P. panicuiata but absentin P. floribunda. Diffuse porous. Vessels soli-tary and in radial multiples of 2-3, sometimesup to 5, occasionally some clusters, 20-46 permm-, 50(60-120) 140 ^m in diameter, 200(260-390) 420 ^im in element length. Simpleperforation plates . Alternate in tervessel pitscircular or oval, of them shape polygonal, sm allto medium, 5-8 \im in diameter. Vessel-ray pitswith distinct borders, similar to intervesselpits in size and shape, pits vestured. Reddishbrown gum-like deposits in vessels. Fibresnon-septate, very thick-walled, 860(1208-1700)1820 um in length. Axial parenchymamostly aliform to confluent, aliform paren-chyma of the winged type, unilateral. Bandedparenchyma in narrow bands or lines, up tothree cells wide. Marginal parenchyma bandspresent in P. panicuiata. Axial parenchyma 3-5cells per parencbyma strand. Rays homocel-lular with typically procumbent cells. 5-10 permm, 2 to 4 cells wide, 140(160-480)830 [xm inheight. Storied structure in axial parenchymain P. panicuiata and irregularly storied inaxial parenchyma in P. floribunda. Prismaticcrystals common in chains in chamhered axialparenchyma cells, one crystal per cell or cham-ber. Intercellular canals of traumatic originpresent in P. floribunda.

Material studied: 7 specimens. P. panicu-iata Benth., MERw 1760. MERw 2463, P flo-ribunda (Kunth) Pittier, MERw 1524. MERw1771, MERw 4568, MERw 4569. MERw4570.

Note: Metcalfe and Chalk (1950) mentiontendencies to storied axial parenchyma in somespecies of Peltogyne. This feature was alsofound in P. floribunda (^ P. porphyrocardia)by JUNAC (1981), but is not mentioned byCorothie (1967) for the same specie and by

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Dtienne et al. (1982) and Nardi and Edlmann(1992) for P . paniculata. However, other spe-cies of the genus were described by Kribs(1968) for /cw/7ora. Dtienne ei fl/. (1982)and Miller and Dtienne (2001) for P . venosa,all with storied structure not observed.DISCUSSION

All wood characters have been recorded,but only the most systematically and diagnosti-cally important ones are displays in Table 1and the following discussion. These charactersemphasized the anatomical information to helpin the identification ofthe species and genera ofthe Venezuelan Detarieae. Diagnostic featuresfor reliable identification and potentially phy-logenetically valuable information within thetribe Detarieae include: fibre wall thickness,ray composition, ray width, intercellular axialcanals and storied structure, parenchyma typeand prismatic crystals in ray cells. Qu antitativefeatures also vary (see Table !), but most varytoo much to be useil in identifications orcomparisons. The exceptions are vessels permm- { e . g . species o Peltogyne) ray width {e.g.species of Hymenaea) and intervessel pit size.Vessels per mm - ^ and ray width are a good diag-nostic quantitative character in these group s.

Comparison of Detarieae w ithCaesalpinieae

1

All legume woods have simple perforationplates, ahemate intervessel pitting, vessel-raypits similar to intervessel pits in size and shape,fibres w ith simple pits, vestured pits (with someexceptions) and in general axial parenchymawith m ostly 2-4 cells per strand. In general., thespecies ofthe subfamily Caesalpinioideae havemedium to thick fibre walls, aliform, confluentand marginal parenchyma, homoceilular raysor with a row of square or upright marginalscells, biseriate rays non-storeyed and prismaticcrystals in chambered axial parenchyma cells(Baretta-Kuipers 1981, Hhn 1999, Herendeen

wood characters provide the most system-atically important characters between Detarieaeand Caesalpinieae:Fibres: septate only in Sehizohbium(Caesalpinieae) and are absent in Detarieaegenera. Usually thin to thick walled inDetarieae and thick and very thick in the mostof Caesalpinieae.Rays : there is considerable variation inray cell composition and ray width betweentaxa. Homocellular rays are most common inCaesalpinieae. while in Detarieae, most com-mon rays are heterocellular. Concerning widthray, in agreement with Metcalfe and Chalk(1950), Baretta and Kuipers (1981), Dtienne

and Welle (1989), Mainieri and Peres (1989).Miller and Dtienne (2001) and Gasson et al.(2003), we observed in both tribes two groups:1) Uniseriate (exclusively or predominant-ly) to biseriate rays in Brownea. Eliiabetha,Heterostemom, Macrolobium^ Sclerolobiumsiibbullatum and Tachigali. 2) Multiseriaterays in Caesalpinia, Campsiandra. Copaifera,Cynometra, Delonix regia. Dimorphandra,Dicymbe, Eperua, Hymenaea, Mora, Peltogyneand Schizolobium.Intercellular axial canals: the presenceand distribution of axial canals is a good diag-nostic and systematic character in Detarieae.They are found in long tangential lines andimmersed in bands of marginal parenchyma inCopaifera and Eperua (Detarieae tribe), alsoreported and discussed by Barettta-Kuipers(1981), Dtienne et al. (1982), Dtienne andWelle (1989), Miller and Dtienne (2001 ) andGasson et al. (2003), occasionally diffuse inEperua purpurea, absent in ail Caesalpinieaegenera.

Axial parenchyma in narrow bandsuncommon in both tribes, however, com-mon in Caesalpinia coriaria, C. ebano ofCaesalpinieae tribe and Macrohbium gracile,M. limbatum, M. molle. Peltogyne floribunda.P . paniculata and occasionally in Hymenaeaoblongifolia and Macrohbium acaciifolium ofDetarieae tribe. Bands more than three cells

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S|33 EJ Dl (0) 5 ench. (iermanand Spanish. Version: liith October 2002. Available:http://biodiversity.uno.edii/delta/ [24 May 2006J,

Wheeler, E. & Baas, P. 1992. Fossil wood of theLeguminosae: a case study in xylem evolution andecological anatomy. In P.S Herendeen & D.I. Dilcher(eds). Advances in Legume Systematics, Part 4. TheFossil Record, pp. 281-301, Royal Botanic Gardens,Kew, England.

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