IAWA Bulletin n.s., Vol. 12 (4),1991: 389-417
WOOD ANATOMY OF THE BIGNONIACEAE, WI'IH A COMPARISON OF TREES AND LlANAS
by
Peter GBS80n 1 and David R. Dobbins2
Summary The secondary xylem anatomy of trees Hess 1943; Jain & Singh 1980; see also Greg and lianas was compared in the family Big ory 1980 and in preparation). Den Outer and noniaceae. General descriptions of the family Veenendaal (1983) have compared Bignonia and the six woody tribes are provided. Lianas ceae wood anatomy with that of Uncarina belong to the tribes Bignonieae, Tecomeae (Pedaliaceae) and Santos has recently describ and Schlegelieae, and most have ve.ssels of ed the New World Tecomeae for an M.S. two distinct diameters, many vessels per unit thesis (Santos 1990) and Flora Neotropica area, large intervascular pits, septate fibres, (Santos in press). large heterocellular rays often of two distinct The family Bignoniaceae has a wide dis sizes, scanty paratracheal and vasicentric axial tribution from about 400N to 35°S, encom parenchyma and anomalous growth. Conver passing North and South America, Africa sely, trees, which belong to the tribes Coleeae, south of the Sahara, Asia, Indonesia, New Crescentieae, Oroxyleae and Tecomeae gen Guinea and eastern Australia. It is mainly erally have narrower vessels in one diameter tropical, with most species in northern South class, fewer vessels per unit area, smaller America, and consists of lianas, trees and intervascular pits, non-septate fibres, small shrubs with very few herbs. Estimates of the homocellular rays, scanty paratracheal, ali number of genera and species vary: 650 form or confluent parenchyma, and none ex species in 120 genera (Willis 1973) or 800 hibits anomalous growth. The majority of species in 110 genera (Takhtajan 1987), al both trees and Hanas possess growth rings, though Gentry (1973) considers that the num are diffuse-porous, have non-solitary vessels ber of genera is too high. which lack helical thickenings, and few have We have examined specimens from 35 tree apotracheal parenchyma or storied structure. and 27 liana genera from six of the eight All species have alternate intervascular pitting tribes recognized by Gentry (1980), Cronquist and simple perforation plates. (1981), and Takhtajan (1987). Two tribes, Key words: Lianas, trees, wood, anomalous both neotropical and monogeneric, Tourret structure, xylem, Bignoniaceae. tieae and Eccremocarpeae, are not covered in this paper since they are herbaceous vines (Gentry 1980). Much of the world distribu Introduction tion of the family is also covered. An overall This paper has two purposes: to present comparison of trees and lianas has been made, descriptions of the secondary xylem anatomy and anatomical descriptions of the six tribes of the Bignoniaceae, and to compare the wood are presented. The tribes Coleeae, Crescen anatomy of trees and Hanas within the family. tieae and Oroxyleae consist entirely of trees Metcalfe and Chalk (1950) have surveyed the and shrubs, whereas all members of the wood anatomy of this family, but there is no tribes Bignonieae and Schlegelieae are lianas. comprehensive publication, although many The Tecomeae are mainly trees and shrubs, genera and species are covered in papers on but Campsis and Tecomanthe are climbers. particular geographical regions (Record & The trees come from a wide range of habitats
1) Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3DS, U.K. 2) Biology Department, Millersville University, Millersville, Pennsylvania 17.5.51, U.S.A.
Downloaded from Brill.com10/04/2021 01:59:21PM via free access 390 IAWA Bulletin n.s., Vol. 12 (4),1991 and climates, e.g. temperate (Catalpa), warm (lAWA Committee 1989) as a guide, with temperate and subtropical (Chilopsis), mon emphasis on those characters most pertinent soon forest (Millingtonia), secondary tropical to the comparison of trees with lianas in this forest (Oroxyiurn), arid tropics (Catophrac- family. These are: growth ring definition; tes), dry tropical forest (Tabebuia, Tecoma, vessel distribution, number, diameter, element Cybistax, Godmania), seasonal savanna (Ki- length and intervascular pit size; fibre type gelia), mangrove (Dolichandrone spathacea), (i.e. septate or non-septate) and wall thick and moist tropical forest (many genera). The ness; axial parenchyma distribution; ray size lianas tend to be mostly from moist tropical and composition; storeying of tissues; pres forest, although some do grow in drier con ence or absence of anomalous secondary ditions, e. g. Cydista and Macfadyena. thickening. Lianas are relatively rapidly growing clim In order to compare the frequency of these bers which produce very long stems and have features in trees and lianas, Tables 1 and 2 limited vascular cambial activity (Schenck (pages 398-407) were prepared. In the ten 1893; Dobbins 1971; Dobbins & Fisher 1986) tree genera in which we examined three or whereas trees and shrubs grow more slowly more species, some features were of variable and have greater cambial activity resulting in occurrence within the genus. For these genera, larger amounts of xylem and phloem. Because an index is given to denote the proportion of of their climbing habit, lianas are subject the genus with the feature (i.e., if three spe to different stresses than trees and shrubs. cies out of five have a feature, the index is Lianas form an extensive canopy, and the dis 0.6). The lianas are treated separately in tance water and solutes have to be transport Table 2. The indices give an indication of the ed is often greater than for trees (Ewers & variability of a feature within a given genus, Fisher 1989). Moreover, the control of devel and are used to calculate the overall percent opment and subsequent anatomical features age frequencies for the comparison of trees in Hanas are demonstrably different from and Hanas, which are on a genus basis. those of trees and shrubs (Schenck 1893), Information on whether the trees are de and this paper documents some of these ana ciduous or evergreen was rarely available on tomical differences in the Bignoniaceae. herbarium sheets or in the floras consulted (see the Appendix), except for Van Steenis Materials and Methods (1977). Where available this information is Apart from some Hanas which were col given in Table 1. It is assumed that most of lected fresh from the Fairchild Tropical Gar the Hanas are evergreen. den, Florida, U.S.A. (FTG), all the material At least 25 measurements were made of examined was from the following institutions: each quantitative feature per sample, and Jodrell Laboratory, Kew (KJw); museum these were bulked to obtain the means for collections, Kew (Kw); Institute of Sys species represented by more than one sample. tematic Botany, University of Utrecht, the For species with vessels of two distinct Netherlands (Uw); Rijksherbarium, Leiden, diameters, only the wider vessels were mea the Netherlands (Lw); Universidad de Los sured. Statistical analyses of mean vessel Andes, Merida, Venezuela (MERw); Forest diameter (Fig. 22), mean vessel element Products Laboratory, Madison, Wisconsin, length (Fig. 23) and intervascular pit border U.S.A. (MADw and SJRw). The slides la diameter (Fig. 24) were made using Kolmo belled FHOw are in the Jodrell Laboratory gorov-Smirnov Chi-square tests. These data, collection ( KJw), but originated from Ox plus number of vessels per square mm were ford Forestry Institute. The abbreviations for examined using an ANOVA. The relationship wood collections follow those in Stern (1988). between mean vessel diameter in J.LM and A complete list of the specimens examined is vessel number per square mm is shown in given in the Appendix (pages 415-417). Figure 25. The statistical results are given Microscopical observations of sectioned with the captions for Figures 22-25. Quan material were interpreted using the IAWA titative data are incomplete because some List of Features for Hardwood Identification samples provided too few measurements for
Downloaded from Brill.com10/04/2021 01:59:21PM via free access Gasson & Dobbins - Trees versus lianas in Bignoniaceae 391 inclusion in the tables and graphs; however, some species and genera is not uniform, i.e. a sufficient number of samples was measured some genera have more than one feature state, to compare trees and lianas in Figures 22-25 the total percentage of genera with a feature (see pages 411 & 412). may exceed 100% (e.g. in trees 48% have distinct. 55 % indistinct and 17 % absent De8criptions growth rings which totals 120%. The same The descriptions are in tribal order for applies to axial parenchyma, where particular trees: Co1eeae, Crescentieae, Oroxyleae and distribution patterns like scanty paratracheal, Tecomeae, followed by lianas: Tecom<;:ae, confluent and aliform often occur together Bignonieae and Schlegelieae. The Tecomeae and are not mutually exclusive). The final trees and lianas are described separately, one description outlines the features found in the after the other for easy reference. These tribal family and refers to the tribes to which certain descriptions provide more detail and qualify characters are restricted. some of the information given in Tables 1 (trees) and 2 (lianas). Since the anatomy of (text continued on page 397)
Legends of Figures 1-13 (trees) and 14-21 (lianas and climbers):
Figs. 1-4. Trees. - 1: Catalpa bignonioides TS. Ring-porous (scale as no.3). - 2: Jacaranda copaia TS. Diffuse-porous, winged aliform axial parenchyma (scale as no.3). - 3: Kigelia afri- cana TS. Diffuse-porous, lozenge aliform and eonfluent parenchyma (scale line = 500 111Il). - 4: Crescentia cujete TS, diffuse-porous, initial, confluent and aliform parenchyma (scale as no.3).
Figs. 5-9 Trees. - 5: Catalpa longissima TS. Diffuse-porous, confluent parenchyma (scale line = 500 J.Lm). - 6: Paratecoma peroba, the only tree with a tendency towards vessels of two dis tinct sizes (scale as no.5). -7: Oroxylum indicum TS, showing a vessel with a foraminate per foration plate and confluent parenchyma (scale line = 500 J.Lm). - 8: Markhamia sessilis TS, showing a vessel with a foraminate perforation plate (scale line = 100 111Il). - 9: Catalpa longis- sima lLS. Short vessel elements with large intervessel pits, axial parenchyma strands of two or more cells, septate fibres and short rays (scale line = 100 111Il).
Figs. 10-13 Trees. - 10: Dolichandrone spathacea TLS. Vessel elements with small intervessel pits, non-septate fibres, short uniseriate rays (scale line = 100 111Il). - 11: Tabebuia impetiginosa TLS. Tissues storied, vessel elements with large intervessel pits, short biseriate rays (scale as no. 10). - 12: Tabebuia caraiba RLS, showing rays tending towards heterocellular (scale line = 500 111Il). - 13: Deplanchea bancana RLS, showing heterocellular rays with one row of marginal square and upright cells (scale line = 100 111Il).
Figs. 14-17 Lianas. - 14: Macfadyena unguis-cati TS, showing the included phloem arrange ment typical oflianas in the tribe Bignonieae (scale line = 1 mm). -15: Cydista aequinoctialis TS. Close up of staircase type phloem (like no.14) with very wide phloem sieve cells (scale line = 500 J.Lm). - 16: Pyrostegia venusta TS. Vessels of two distinct sizes (scale as no.15). - 17. Campsis radicans TS. A climber, not usually considered a liana. Ring-porous, without phloem wedges (scale as no.15).
Figs. 18-21 Lianas. -18: Schlegelia parasitica TS. Diffuse-porous, with sparse narrow ves sels, and no phloem wedges (scale line = 500 111Il). -19: Phryganocydia corymbosa lLS. Very tall rays, septate fibres (scale as no.18). - 20: Arrabidaea corallina lLS. Short wide and narrow rays of two distinct sizes, septate fibres (scale as no.18). - 21: Schlegelia albiflora RLS. Hetero cellular rays with more than one row of square and upright cells (scale as no.18).
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Trees by Tribe: See Table 1. cereifera, and one to several small rhomboi dal crystals in some unchambered ray cells of C. (MADw 42263). Axial parenchyma Tribe Coleeae: Kigelia, Ophiocolea, Rhodo- alata and rays usually storied in and colea. See Table 1. Amphitecna Crescentia, axial parenchyma irregularly stor Growth rings indistinct or absent. Diffuse ied in Enallagma. porous. Vessels solitary, in pairs and a few small Tribe Oroxyleae: See clusters and radial multiples up to 6. . Millingtonia, Oroxy1wn. Table 1. Fibres non-septate, fibre walls thin to thick in Ophiocolea, thick in Kigelia and Rhodo- Growth rings indistinct to distinct. Diffuse colea. porous. Axial parenchyma scanty paratracheal and Vessels solitary, in pairs, radial multiples diffuse in aggregates in Ophiocolea and Rho- and a few small clusters up to 5. Perforation docolea, some confluent in Rhodocolea; in plates simple, except for occasional forami Kigelia lozenge aliform to confluent (Fig. 3), nate plates in both genera (Fig. 7). tangential band width 5 cells or more, some Fibres non-septate. times so wide that the ground tissue in some Axial parenchyma confluent and initial, areas is entirely of axial parenchyma with no also scanty paratracheal in Millingtonia and fibres. vasicentric and lozenge aliform in Oroxylum Rays uniseriate, up to 20 cells high in (Fig. 7). Rhodocolea and 40 cells high in Ophiocolea, Rays 2-3 cells wide and up to 15-30 2-4 cells wide and up to 15 cells high in Ki- cells high in Millingtonia, 1-4 cells wide and gelia. Rays homocellular, but a few square up to 25 cells high in Oroxylum, rays homo and upright cells within rays of Ophiocolea, cellular. One to several small crystals of vari many unchambered ray cells containing one ous shapes in some unchambered ray cells in or more small rhomboidal crystals in Ophio- Millingtonia (KJw 3645), small rhomboidal coleaonly. crystals in some unchambered ray cells in Oroxylum. Tribe Crescentieae: Amphitecna, Crescentia, Enallagma, Parmentiera. See Table 1. Tribe Tecomeae: 26 genera. See Table 1. Growth rings distinct in Parmentiera, oth Growth rings distinct in 18, indistinct in erwise indistinct or absent. Diffuse-porous. 15 and absent in 9 genera. Ring-porous in Vessels solitary and in pairs in Amphitec- Catalpa bignonioides (Fig. 1) , C. bungei, C .. na and Crescentia, also in a few small clus ovata and C. speciosa, Markhamia acumina- ters in Enallagma and Parmentiera. ta, M. sessilis and M. stipulata; semi-ring~ Fibres non-septate. porous in Catophractes, Chilopsis and Mark- Axial parenchyma aliform and confluent hamia zanzibarica; all other species diffuse in all genera (Fig. 4), tangential bands 2-7 porous (Figs. 5-7). cells wide in Enallagma, initial in Crescentia Vessels: Slight tendency towards vessels and Parmentiera, also scanty paratracheal in of two distinct sizes in some parts of Para- Parmentiera. One to several small rhomboidal tecoma cross sections. Vessels solitary, in crystals in some unchambered axial parenchy pairs, radial multiples and a few clusters up ma cells in Amphitecna and Crescentia cujete. to 7 in most species, occasionally more, i.e. Rays uniseriate and up to 20 cells high in 8-17 in Dolichandrone, to 11 in Jacaranda, Amphitecna, uniseriate and biseriate and up to 20 in Paratecoma. Radial multiples most to 15 cells high in other 3 genera; homocellu common in Tabebuia chrysantha, T. chrysea, lar except in Parmentiera, which has some T. rosea and T. stenocalyx. Latewood ves rays with one row of square marginal cells. sels in clusters and tangential arrangement in One to several small crystals of various Catalpa ovata and C. speciosa and Chilopsis. shapes in some unchambered ray cells in P. Perforation plates simple, except for occasio-
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Table 1. Trees. Secondary xylem characters (98 species in 35 genera). Key: + =present Index: 1; all species in the genus have this feature; 0.1 to 0.9 is the proportion of species in the genus with this feature. E ~ E. £ 01) £ 1) ~ ] " ~ u '::1= £ <: ~ ~ ~ '::1 '"~ '"01) = '" li .;; N S eo .S:! B .~ :a :I'" .S " " £ S "0 .S ~'" 0 S 0; 'S '":I = S ~ " 9 ; '"01) 01) eo ~ ~ os. £ B £' '" '" '":I S 01) ~ ~ C: ·c= ·c= ·c = l ...0 :§ 8- '" ~ ~ 0 .~ ~ '" ~ > £' £ £ £ [ = ~ 0; OJ "> '" = ~ '" '" '" ~ be .a ~ ~ " e'" e'" '" e ·s ... '" ~ ~ <§ <§ ] or> ~ ~ l ·2 :a "> ~ ~ ~ s .S <:l '= Coleeae "'" Kigelia africana + + 11 101 196 4.4 + + Ophiocolea floribunda + + 68 263 4.7 + + R hodocolea telfairiae + + + + + Crescentieae Amphitecna lati/olia + + 18 90 245 3.6 + + Crescentia (2) + + 25 58 209 5.0 + + Enallagma (2) + + + 12 230 3.6 + + ParmenJiera (2) + + 56 244 3.9 + + Oroxyleae Millingtonia hortensis + + + 63 245 4.4 + + Oroxylum indicum + + 140 240 6.2 + + Tecomeae Catalpa bignonioides + + LWV 18 90 192 8.8 + + bungei + + + + ovata + + LWV 149 188 6.1 + + speciosa + + LWV 12 94 277 6.9 + + denticulata + + + + longissima + + + 8 87 255 6.6 + + punctata + + + + + No. of species 7 7 2 4 3 3 7 5 2 Index 0.3 0.6 0.4 0.4 0.70.3
Catophractes alexandrii + + + 68 49 139 3.6 + + Chilopsis linearis + + + + 19 113 167 4.4 + + Cotema spiralis + + + + Delostoma integrifolium + + 51 360 3.6 + + Deplanchea (2) + + + 146 408 5.2 + + Digomphia densicoma + + + + Dolichandrone atrovirens + + 66 251 10.2 + + falcata + + 20 104 352 3.9 + + heterophylla + + + 76 257 4.7 + + spathacea + + + + 14 73 274 4.4 + + stipulata + + 16 108 329 3.9 + + No. of species 5 1 3 4 5 5 5 Index 0.2 0.60.8 1 1
Ekmanianthe actinophyUa + + + + Fernandoa adenophylla + + + 17 102 247 3.7 + (+) + magnifica + + 72 208 3.6 + (+) + roxburghii + + 107 333 2.7 + + No. of species 3 1 3 3 3 3 2 Index 0.3 1 1 1 1 0.7
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(Table 1 continued)
~ <> g ose 01 >- ] .5 5 -5 01 2l 1: e ~--. = '" 1 .~ e ::> >0 e .<: ~ ~ 1 ... e 0)", ! ~ 1 ~ ~ ~ ~ 0 ] .<: .. ::> e '" 6';;' >- :3 ~ ~ ..e '"~ .~ --.::> '" g 01 ~ ~
+ + + + 2-4 15 + + + + (+) 40 + + (+) + 20
+ + + + + + + 1 20 0.5 0.5 (+) + + + + + 1(2) 15 (+) + + + + + 1-2 15 0.5 + + + + + + 1(2) 10-15
(+) + + + + 2-3 15-30 E/D + + + + + + (+) 1-4 25 SD
+ + 1-3 20 D + + 1-3 40 D + + + + + 1-5 50 D + + 1-2 15 + + 1-2 20 + + + 1-3 15 DIE + + 1-2 20 1 1 7 2 7 0.1 0.1 1 0.3 1
+ + + + (+) 1-2 40 + + + + 1-4 35 + + + + 1-2 10 + + + 1-4 15 0.5 + + + + + 1-2 15 + + 1-2 15 + + + + + + 1-3 15 + + + + + 1-3 20 + + + + + + + (+) 1-2 35 + + + + + (+) + + (+) + + 1 20 E + + + + + 1-3 30 2 5 1 2 5 3 4 3 5 2 2 1 0.4 0.20.4 0.60.80.6 0.4 0.4 0.2
+ + + + + 1-2 15 + + + + + 2-3 20 DIE + + + + + 1-2 20 D + + + 1-3 20 2 1 3 2 1 3 1 0.7 0.3 0.7 0.3 1 0.3
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(Table 1 continued) ~ ~ -5 ... 00 B., ~ ..@ § ~ u ... ~ g '.:l= C ., B., c., :a ~ ~ ., '" 00 ~ '.:l '" N 00 .Sol B :a '" ::s .~ '" e ~ ~ .5 ~ '" .~ e .~ -5 .Sol S e ~ ~ g S bO 00 '"bO '" ~ ~ 's. -5 III '" '" '"::s ~ .... ~ ~ S C:"" '1:= '1: = '1: = bO 0 ~ it ., .,'" ~ ~ fr 0 = >'" > ., ~ > '" -5 -5 -5 ~ '9
+ + + 1-2 35 D + + + + + + + 1-3 25 D + + + + (+) 1-3 40 D + + + + + 1-5 70 (+) D + + + + + + + 1-2 30 D + + + + + + 1-3 25 D 3 5 4 5 6 3 2 5 1 0.5 0.8 0.70.8 0.5 0.3 0.8 0.2 + + + + (+) 1-2 20 + + + + + + 1-3 20 + + + + + 1-3 30 + + + (+) + + + 2-3 15 + + + + + 1 10 + + + + 2 25 E + + + + 2-3 25 E + + + + + + + 2-4 20 E + + + + + 1-2 25 E + + + + 1-4 20 E 5 3 1 2 5 2 2 3 1 1 0.6 0.2 0.4 1 0.4 0.4 0.60.2 + + + + 1-4 50 + + + + + + + 1-5 30 (+) D + + + + 30 1 1 2 2 1 1 1 1 1 0.5 0.5 1 1 0.5 0.5 0.5 0.5 0.5
+ + + 2 15 D + + + + + (+) 1-4 50 + + + + 1-4 35 D + + + + + + (+) 1-3 20 2 1 4 4 3 4 2 0.5 0.3 0.8 0.5
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(Table 1 continued) ~ ~ -5 toO B., ~ .§ ~ 'Gc ~ ] 'Gc .::: .. B ;: :a .::: toO ., ., ~ -= .. 5 .. .~ .. .. S ctoO £ .~ :a .. :I .~ S ., :g ~ S "0 .5 'i 0 'f! .~ ., .. ., S ~
Tecoma aurea + + 6 97 245 5.2 + + castanifolia + + + 29 69 239 3.6 + + conspicua + + 14 102 299 6.6 + + garrocl/IJ + + + + grandis + + 19 98 287 7.0 + (+) + leucoxylon + + 21 105 308 4.4 + + pedicellata + + 224 5.8 + + serratifolia + + 73 277 4.7 + (+) + spectabilis + + 15 78 229 6.6 + + + stans + + + + 63 257 4.1 + + undulata + + 10 80 179 4.4 + (+) + No. of species 11 1 11 2 11 10 2 6 8 Index 0.1 1 0.2 0.9 0.2 0.5 0.7
Tecomaria capensis + + + + Zeyheria tuberculosa + + + + +
Total 35 genera 15 19.26.6 1.1 2.2 33.9 (1) 1.4 34.60.5 7 31: % 43 55 19 3 6 97 (3) 4 99 1 20 91
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+ + + + (+) 1(2) 20 + + + + 2 20 + + + + 2 10 + + + + + + 1-3(8) 35 + + + + + 1-3 11 + (+) + + + + + 1-2 15 + + + + + + 1-2 7 + + + + 2 21 + + + + + + 2 10 + + + + + + 1-3 45 + + + (+) 3-4 35 3 6 2 10 6 7 7 8 1 4 3 0.3 0.5 0.2 0.9 0.5 0.6 0.6 0.7 0.1 0.4 0.3
+ + 1-2 15 + + + + 1-3 10
9.8 2.5 6.2 2.4 7.2 3 22.1 8 7.5 24.60.5 17 2.8 4.1 27.1 8.6 12 3.6 1.7 28 7 18 7 21 9 63 23 21 70 1 49 8 12 77 25 34 10 5 50
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Table 2. Lianas. Secondary xylem characters (39 species in 27 genera). Key: + =presenL Index: I, all species in the genus have this feature; 0.1 to 0.9 is the proportion of species in the genus with this feature.
! !... t £.., g ! Jj u ., -c .=., C ., ~.., c.., ~ .., 1;l bI) bI) ., ., N .= :a :s .0; c S ..,S .5 i 0 ., ·2 S .~ .~ :a., ., ., :s S
Arrabidoea chica + + + 64 75 235 6.8 cora/lina + + + mollissima + + + pubescens + + + + triplinervia + + + + 28 139 303 7.7 No. of species 5 2 5 5 5 Index 0.4 1
Callichlamys lati/olia + + + 8 155 296 6.9 Chodanthus puberulus + + + 61 lOS 297 5.8 Cydista aequinoctialis + + + + 32 116 266 4.7 Macjadyena unguis-cati + + + + 48 112 285 6.1 Macrodiscus lactiflorus + + + Mansoa ve"ucifera + + + 81 68 290 4.7 Memora flaviflora + + + romeroi + + + No. of species 2 1 1 2 2 Index 0.5 0.5 1
Mussatia hyacinthina + + + Neomac/adya podopogon + + + 90 92 195 6.8 Paragonia pyramidata + + + 253 Petastoma broadwayi + + + + patelliferum + + + 25 59 246 No. of species 2 1 1 1 2 2 Index 0.5 0.5 0.5 1 1
Phryganocydia corymbosa + + + 29 133 354 Pithecoctenium crucigerum + + + 62 121 220 6.2 dolichoides + + + 21 143 263 5.2 No. of species 2 2 2 2 Index 1 1 1
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(Table 2 continued)
..,'" =u §: ~ as (;I 5 ~ .. .~ ~ '5 e::0 .., e e i) ~ B 1 ~ ~ ';( -5 23 .c ::0 ::0 'Cl ..,'" 6 '">- ..,
+ + + + + 1-4 36 + + + .+ + 1-5 40 + + + + + + 2-10 100 + + + + + + 1-3 80 + + + + + (+) + 1-16 120 + 5 5 1 5 3 2 1 5 3 0.2 0.6 0.4 0.2 1 0.6
+ + + + 1-2 90 + + + + + 1-6 340 + + + + + + 1-6 250 + + + + + + 1-6 250 + E + + + + + 1-2 50 + + + + + 1-5 300 + + ? ? + + + 1-7 110 + + + 2 1 1 1 1 0.5 0.5 0.5 0.5
+ + + + 2-5 ? + + + + + 1-3 45 + + + + 2-10 550 + + + + + + 1-8 70 + + + + + + 1-4 490 + 2 2 1 2 1 2 2 1 1 0.5 1 0.5 1 1
+ + + + (+) 1-8 430 + + + + + + 1-7 550 + + + + + + + (+) 2-16 600 + 2 2 2 2 1 1 2 2 1 1 1 1 0.5 0.5 1 1
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(Table 2 continued)
~ .s ~... 00 B 1:) ~ ] §" 1:) c c '::1 C '" !a C :a '::1 ~ 00 00 '" '0; '"c N e e C .~ :a "'" '" " " "t:I .S ~ '" 0 '8 e :g .:3 e '" ] e
Urbanolophium dusenianum + + + 15 132 174
Schlegelieae Schlegelia albiflora + + + + nicaraguensis + + + + parasitica + + 38 40 298 4.0 No. of species 3 2 3 3 2 Index 0.7 0.7
Tecomeae Campsis chinensis + + + + 16 122 237 6.1 radicans + + + 130 269 6.1 No. of species 2 2 1 1 1 2 Index 1 0.5 0.5 0.5 1
Tecomanthe venusta + + + 64 110 238 5.1
Total 27 genera 8.6 23 0.5 0.5 25.5 24.7 1 % 32 85 2 2 94 91 4
nal forarninate plates in Catophractes, Doli- signis and Tecoma aurea. Helical thickenings chandrone atrovirens andD. spathacea, Mark- in latewood vessels in Catalpa bignonioides, hamia sessilis (Fig. 8), Romeroa, Spathodea C. ovata and C. speciosa and Chilopsis line- campanuiara, Tabebuia haemantha and T. in- aris.
Downloaded from Brill.com10/04/2021 01:59:21PM via free access Gasson & Dobbins - Trees versus lianas in Bignoniaceae 407
(Table 2 continued)
.,'" =u @- 13 . .E ., .;;; 9 '" ;; '" ~ e ~ g = '" 0 § iC e i~ E .5 -5 0 ;:., e ' + + + + + 1-10 130 + + + + + + + 1-6 55 + + + + + + + 1-3 50 + + + + + + 1-2 30 3 1 3 3 2 3 3 1 1 0.3 1 0.7 0.3 + + + + + 1-4 60 D + + + + + 1-4 70 (+) D 2 2 2 2 2 1 0.5 + + + + + + 1-7 70 + 27 22.3 4.5 3 6.7 26.5 0.6 6.6 1.7 25.5 5.5 16.9 100 83 17 11 25 98 2 24 6 94 20 63 Fibres thin-walled in Tabebuia cassinoides buia barbata, T. capitata, T. cf. catinga, T. and T. rosea; thick-walled in Catophractes, chrysantha, T. guayacan and T. serrati/olia, Cotema, Ekmanianthe, Phyllarthron, Tecoma- Tecoma pedicellata, and T. spectabilis. Fibres ria and Zeyheria; very thick-walled in Tabe- entirely non-septate in 15 and in some spe- Downloaded from Brill.com10/04/2021 01:59:21PM via free access 408 IAWA Bulletin n.s., Vol. 12 (4),1991 cies of another 5 genera. Most if not all fibres genera (Fig. 12). Rays of two distinct sizes septate in Catalpa ovata, C. speciosa, C. den- in Romeroa, almost so in Markhamia sessilis, ticulata, C. longissima (Fig. 9) and C. punc- Spathodea campanulata and S. gigantea. tata, Delostoma, Fernandoa, Spathodea gi- Crystals of various shapes, several per gantea, Tecoma castanifolia, T. garrocha, T. cell in non-chambered axial parenchyma in stans and Tecomaria. At least some fibres sep Pajanelia, and in ray cells in 8 genera, a tate in Markhamia acuminata and M. sessilis, single crystal per ray cell in Markhamia Pajanelia, Radermacheragigantea andR. sini- zanzibarica. Storied in Cotema, Godmania, ca, Romeroa, Spathodea campanulata and S. Tabebuia angustata, T. barbata, T. capitata, serratula, Stereospermumfimbriatum, Teco- T. cf. catinga, T. chrysantha, T. guayacan, ma grandis, T. serratifolia and T. undulata. T. haemantha, T. impetiginosa (Fig. 11), T. Axial parenchyma paratracheal in all spe pallida, T. rosea, T. roseo-alba and T. ser- cies. Some vasicentric in Catalpa ovata, Ek- ratifolia, Tecoma conspicua, T. grandis, T. manianthe, Pajanelia, Paratecoma, Phyllar- leucoxylon, T. pedicellata and T. spectabilis thron, Radermachera, Spathodea serratuia, and Zeyheria. Some tendency towards storey Stereospermum neuranthum and S. xylo- ing of axial parenchyma in Catalpa, especially carpum, Tabebuia angustata and T. roseo- C. ovata, and of some parenchyma in alba. Diffuse in aggregates only in Ekman- Tecoma aurea, and irregularly storied in rays ianthe. Scanty paratracheal in 22 genera (not of Tabebuia caraiba and Tecoma serratifolia. in Deplanchea, Godmania, Newbouldia and Pajanelia). in 14 genera (Fig. 2). Alifonn Con- by Tribe: See Table 2. fluent in 19 genera. Unilateral paratracheal in Lianas Jacaranda copaia, J. cuspidifolia, J. glabra, Tribe Tecomeae: Campsis, Tecomanthe. See J. mimosifolia and J. obtusifolia. Initial in Table 2. 16/26 genera. Tangential bands less than 3 cells wide in Deplanchea and Dolichandrone, Growth rings indistinct in Tecomanthe, more than 3 cells wide in Dolichandrone distinct in Campsis. Ring-porous in C. radi- atrovirens, D. heterophylla and D. spathacea, cans (Fig. 17), diffuse-porous in C. chinen- Godmania and Spathodea campanulata. sis and Tecomanthe. Rays mainly 1-3 cells wide. Uniseriate in Vessels of two distinct sizes in C. chinen- Dolichandrone spathacea (Fig. 10), Jacaranda sis. Vessels solitary, in pairs, radial multiples caucana, J. coeruIea, J. hesperia, J. mimosi- and a few clusters up to 6, but longer radial folia and J. obtusifolia, Phyllanhron, Spath- multiples are abundant in smaller vessels in odea serratula, Tabebuia caraiba and T. cas- C. chinensis and latewood vessels in C. radi- sinoides; 1-2 cells wide in some species in cans. Helical thickenings in narrow vessels in 13 genera. Biseriate in Radermachera gigan- Campsis. tea, Stereospermum fimbriatum, Tabebuia Fibres septate in Campsis, some fibres capitata and T. impetiginosa (Fig. 11), Teco- septate in Tecomanthe, thin- to thick-walled. ma castanifolia, T. conspicua, T. serratifolia Axial parenchyma scanty paratracheal. and T. spectabilis. Rays up to 10 cells high Rays 1-4 cells wide and up to 70 cells in Cotema, Phyllanhron, Tabebuia angustata, high. Rays heterocellular with several rows T. barbata, T. chrysantha, T. guayacan, T. of square and upright cells at ray margins in haemantha and T. serratifolia, Tecoma con- Campsis and mixed within the body of the spicua, T. pedicel/ata, T. spectabilis and Zey- rays in Tecomanthe. Several tiny needle-like heria. Rays up to 20 cells high in some spe crystals present in some ray cells in Teco- cies of 16 genera, up to 21-50 cells high in manthe. Axial parenchyma and narrow ves some species of 12 genera and up to 70 cells sels storied in Campsis. high in Markhamia sessilis. Rays mainly Cambial variants absent, although fOUT homocellular in 22 genera, but heterocellular, slight indentations occur in cambium ofTe- with one or more rows of square or upright comanthe, which could possibly develop into marginal cells (Fig. 13), or square and up a similar variant to that in Bignonieae, but right cells within the body of the rays in 20 no wider stems available for examination. Downloaded from Brill.com10/04/2021 01:59:21PM via free access Gasson & Dobbins - Trees versus lianas in Bignoniaceae 409 Tribe Bignonieae: 24 genera. See Table 2. Pyrostegia, Saritaea, Tanaecium and Tynnan- thus; eight or more indentations in Cydista, Growth rings distinct in 9, indistinct in 23 Macfadyena (Fig. 14), Mansoa, Paragonia, genera. Phryganocydia and Pseudocalymma; number Vessels solitary, in pairs and often frequent of indentations could not be determined for clusters and radial multiples up to 12. Ves some genera because of the samples' incom sels of two distinct sizes (Fig. 16) in all spe plete cross sections. Phloem indentations step cies except Adenocalymna and Anemopaeg- ped (i. e. staircase-like) in Anemopaegma, Ar- ma robustufn. Vasicentric tracheids sOple rabidaea pubescens and A. triplinervia,' Cydis- times present in association with narrow ves ta (Fig. 15), Macjadyena, Mansoa, Memora, sel elements. Mussatia, Paragonia, Phryganocydia, Pithe- Fibres septate and thin- to thick-walled, coctenium dolichoides, Pleonotoma, Pseudo- except in Callichlamys and Tanaecium which calymma, Tynnanthus and Urbanolophium. have non-septate and thin-walled fibres. Sieve cells in indentations much larger than Axial parenchyma scanty paratracheal and those in normal phloem in Arrabidaea chica, vasicentric (poorly defined) in all genera, also Cydista, Neomacjadya, Petastoma patellife- confluent in Arrabidaea chica, A. corallina rum, Pithecoctenium crucigerum, Tanaecium and A. mollissima and initial in Arrabidaea jaroba and Tynnanthus elegans, and slightly pubescens and A. triplinervia, Memora flavi- so in Saritaea. flora, Neomacjadya, Petastoma broadwayi, Pithecoctenium dolichoides, Tanaecium, Tyn- Tribe Schlegelieae: Schlegelia. See Table 2. nanthus and Urbanolophium. Rays 1-16 cells wide. Rays 1-2 cells wide Growth rings indistinct to distinct Diffuse in Callichlamys, Macrodiscus artd Saritaea, porous. up to 3 cells wide in Arrabidaea pubescens, Vessels of two distinct sizes in S. albiflora Neomacfadya, Pleonotoma and Pyrostegia, and S. nicaraguensis, but not S. parasitica up to 50 cells high in Arrabidaea chica, A. (Fig. 18). Vessels solitary and in pairs in S. corallina, Macrodiscus, Neomacfadya and albiflora, also in radial multiples and a few Pleonotoma, up to 50-100 cells high in Ar- clusters up to 6 in S. nicaraguensis and S. rabidaea mollissima and A. pubescens, Cal- parasitica. lichlamys, Petastoma broadwayi, Pyrostegia, Fibres non-septate, except for some septate Saritaea, Tanaecium and Tynnanthus micran- fibres in S. parasitica, thin-walled. thus. Other species with rays up to 101-600 Axial parenchyma scanty paratracheal and cells high (Fig. 19). Rays of two distinct vasicentric, also initial in S. albiflora and S. sizes except in Adenocalymna, Arrabidaea nicaraguensis. chica, Callichlamys, Macrodiscus, Neomac- Rays 1-2 cells wide and up to 30 cells fadya, P/eonotoma, Pyrostegia and Saritaea. high in S. parasitica; 1-3 cells wide in S. ni- Rays mainly heterocellular, often mixed or caraguensis and 1-6 cells wide in S. albiflora, with many rows of square and upright cells. up to 55 cells high in both. Rays heterocellu Axial parenchyma more or less storied in Ar- lar, with rays composed of mixed procum rabidaea triplinervia, Chodanthus, Cydista, bent, square and upright cells and several Macfadyena (with small rays), Mansoa (with rows of square and upright cells at ray mar short fibres), Pithecoctenium and Tynnanthus gins (Fig. 21). elegans. No storeying or cambial variants. One or more crystals of various shapes in unchambered ray cells in Adenocalymna and Family Description Tanaecium. Cambial variant in the form of phloem in Trees: 35 genera; 3 genera and 3 species of dentations (wedges) present in all species Coleeae; 4 genera and 7 species of Cres (Figs. 14, 15). Four indentations in Adeno- centieae; 2 genera and 2 species of Or- calymna, Arrabidaea, Chodanthus, Memora, oxyleae, 26 genera and 87 species of Neomacfadya, Petastoma, Pithecoctenium, Tecomeae. Downloaded from Brill.com10/04/2021 01:59:21PM via free access 410 IAWA Bulletin n.s., Vol. 12 (4),1991 Lianas: 27 genera; 24 genera and 33 species Discussion of Bignonieae, 1 genus and 3 species of This paper outlines some of the overall Schlegelieae, 2 genera and 3 species of wood anatomical differences between trees Tecomeae. and lianas, and for the first time provides ac counts of the wood anatomy of all the woody Growth rings distinct, indistinct or absent. tribes in the Bignoniaceae. Most species diffuse-porous, but some spe The wood anatomical features of Bigno cies of Tecomeae ring-porous or semi-ring niaceae trees and lianas fall into three groups: porous. 1) features restricted to or more common in Vessels solitary, in pairs, clusters and ra trees, 2) features restricted to or more com dial multiples of varying number. Vessel mon in lianas, and (3) features equally com clusters commoner in lianas and ring-porous mon in trees and lianas, including those trees. Vessels of two distinct sizes almost al common to the entire family. ways present in lianas but virtually absent in trees. Intervessel and vessel-ray pitting alter 1) Features restricted to or more common in - Rays exclusively uniseriate (12%), nate and of the same size, perforation plates trees and the occurrence of very infrequent fora exclusively simple, except for very occasion minate perforation plates in a scattering of al foraminate plates in some species of Oro species from Oroxyleae and Tecomeae come xyleae and Tecomeae. Helical thickenings into this category (although Handa, 1936, rare, only found in narrow vessels in ring, records foraminate plates in semi-ring-porous or diffuse-porous (with Campsis grandi- which would be treated as a liana in vessels of two distinct sizes) Tecomeae trees flora, this paper). Growth rings are absent in 19% and lianas. Vasicentric tracheids present in of trees. In lianas they are always present, some Bignonieae lianas. but are often indistinct. Aliform (50%) and Fibres either non-septate (in most trees) or initial parenchyma (49%) were recorded only septate (in most lianas), occasionally both in trees and confluent parenchyma was found types present. Simple or very minutely border in 70% of trees and only 2% oflianas. Banded ed pits usually present in the radial fibre walls. parenchyma (up to 3 cells wide in 8% and over Fibre walls usually thin to thick, occasionally 3 cells wide in 12%) was found only in trees, thin or very thick. and so were rays up to 20 cells high (72%). Axial parenchyma mainly scanty paratra cheal, vasicentric, aliform, confluent and ini 2) Features restricted to or more common in tial in trees, scanty paratracheal and vasicentric lianas - Most of the lianas have four or eight (poorly defined) in lianas. wedges of phloem resulting from the failure Rays 1-5 cells wide and up to 50 cells of the cambium to produce secondary xylem high in trees, 1-16 cells wide and usually while continuing phloem production (Dobbins over 20 and up to 600 cells high in lianas. 1971). This cambial variant was restricted to Rays mainly homocellular in trees, hetero Bignonieae in our samples, and Gentry (1980) cellular in lianas. Rays of two distinct sizes believes it is restricted to this tribe. The only common in lianas but virtually absent in sample of Tecomanthe (Tecomeae) available trees. Storied or irregularly storied tissues was very narrow, only 13 mm in diameter in equally uncommon in trees and lianas, but cluding bark, and had four very slight phloem characteristic of certain genera of Crescen indentations. Wider stems of Tecomanthe tieae and Tecomeae trees. would be needed to see whether phloem Crystals uncommon, usually small and of wedges occur, which would extend their oc varying shape, several per non-chambered ray currence outside the Bignonieae. Internal or less commonly non-chambered axial paren phloem has been recorded in Campsis (Teco chyma cell. Prismatic crystals even rarer. meae) by Handa (1936), but this is an entire Cambial variant in the form of phloem in ly different variant from the phloem wedges dentations restricted to Bignonieae lianas. in Bignonieae. Downloaded from Brill.com10/04/2021 01:59:21PM via free access Gasson & Dobbins - Trees versus lianas in Bignoniaceae 411 Vessels of two distinct sizes were found Close examination of narrow tracheary ele in 91 % of lianas whereas only one tree spe ments usually resulted in the majority being cies with diffuse porous wood has any ten identified as vessels. In addition to having dency towards this feature (Paratecoma pero- vessel of two distinct sizes, the mean vessel ba, Fig. 6). Some vasicentric tracheids are diameters of lianas are significantly wider found in association with narrow vessels in than in trees, and in 63% of lianas are above Macfadyena (first reported by Carlquist 100 11m. The presence of wide vessels in 1985a, who called the genus Doxantha). They lianas is well known, and in taxa with both occur in this and other species of Bignonieae tree and liana representatives the vessels in in association with narrow vessel elements. the lianas are almost invariably wider (Abuta 40 r/J G) '<3 G)c. Vl 30 ..... o Trees & Shrubs 0 G) • Lianas OJ) .:s 20 i3 8 & 10 40 60 80 100 120 140 160 180 200 Tangential vessel diameter (~) Fig. 22. Histogram comparing the frequency distribution of mean tangential vessel diameter in trees and shrubs versus lianas (vessels are significantly wider in lianas than in trees, 63% over 100~: Kolmogorov-Smirnov test, Chi-square = 12.14, P < 0.01; ANOVA F = 7.1, P < 0.008). r/J 40 .~ uG) C...... r/J 30 0 o Trees & Shrubs G) OJ) 00 • Lianas E 20 G) ~ c.. 10 140 180 220 260 300 340 380 420 460 500 540 Vessel element length (~) Fig. 23. Histogram comparing the frequency distribution of mean vessel element length in trees and shrubs versus lianas (similar in trees and lianas: Kolmogorov-Smirnov test, Chi-square = 2.56, P < 0.05, not significant; ANOVA F = 0.5, P < 0.05, not significant). Downloaded from Brill.com10/04/2021 01:59:21PM via free access 412 IAWA Bulletin n.s., Vol. 12 (4),1991 Percentage of species both wide and narrow vessels means that even if some wide vessels become blocked 40 by embolism the narrower ones would re o. main functional. Vessel element length is j similar in both groups regardless of vessel 30 CIl diameter and would be expected to have less influence than vessel diameter on water flow. Septate fibres are much more common in 20 lianas (83%) than trees (20%) and this seems to be linked with a lower frequency of axial parenchyma and the almost total absence of 10 bands of parenchyma in lianas. Spackman Trees versus Lianas 2345678910 Intervessel pit diameter (J.1ffi) Mean vessel diameter in J.1ffi 200,------, Fig. 24. Histogram comparing the frequency ~ distribution of mean intervessel pit border 180- 0 A 0 diameter in trees and shrubs versus lianas Trees A (intervessel pits are significantly larger in ~ 160- A lianas than in trees, 50% over 6 j.Lm: Kolmo 0 0 gorov-Smirnov test, Chi-square = 10.18, A 0 P < 0.01). 140 0 0 0 A 0 120- J!' AD 0 DO A 0 grandijlora, Menispermaceae, Mennega 1982; A A Dr) A... • Zanthoxylum, Rutaceae, Caesalpinia and 100 A ~ AI. 0 Bauhinia, Leguminosae, Bamber 1984; Carl AI. .. quist 1985b; Ewers 1990). Lianas also et al. 80 have significantly more vessels per unit area (tA 0 than trees (P < 0.0001). The relationship be A DO 0 0 tween vessel density and mean vessel diam 60- A Ali! A A 0 eter in trees and lianas is shown in Fig. 25. A A The intervascular pits of lianas are also sig 40 0 0 nificantly larger than in trees. Half of the liana species have pit borders exceeding 6 11m. 20 0'------:'20,------4.LO------:6LO-----='80=------'1 00 All these vessel features suggest that Big Vessels per square ffiffi noniaceae lianas are much more efficient at water conduction than trees. Water conduc Fig. 25. Scatter diagram showing the relation tivity is directly proportional to the fourth ship between mean tangential vessel diameter power of the radius of a vessel or capillary, (11m) and number of vessels per square mm so in identical conditions one vessel twice the in trees and lianas. Trees: closed triangles, y = diameter of another will transport sixteen 117.7 - 2.279 x, r = -0.6184 (p <0.0001). times as much water (Zimmermann 1983; Lianas: open squares, y = 158.49 - 1.997x, r Ewers & Fisher 1989). The greater density = 0.4557 (p < 0.005). There are significant of vessels per unit area and larger intervascu ly more vessels per unit area in lianas, usual lar pits would further enhance the ability of a ly more than 20 per square mm: ANOV A liana to transport water, and the presence of F = 51.6, P < 0.0001. Downloaded from Brill.com10/04/2021 01:59:21PM via free access Gasson & Dobbins - Trees versus lianas in Bignoniaceae 413 and Swamy (1949) observed that there is little wide (Wheeler et al. 1986). All Bignoniaceae axial parenchyma in species with many sep have alternate intervascular pitting and simple tate fibres, but this is not a simple or univer perforation plates in the vessels, as do 87% sal relationship (Chalk 1983). Septate fibres and 89% of taxa worldwide. The presence of are found in many taxonomic groups and in distinct growth rings is higher for Bignonia- plants growing in a wide range of temperate ceae than taxa worldwide (48% trees, 32% and tropical habitats. Baas and Schweingruber lianas versus 31 % worldwide). Likewise, ves (1987) consider that septate fibres are largely sels not solitary (all trees and lianas versus restricted to tropical families (using observa 83% worldwide), paratracheal parenchyma tions and data from Wheeler et aI. 1986; Met (100% trees and lianas versus 50% world calfe & Chalk 1950). Fahn et aI. (1986) show wide), storied tissues (18% trees, 25% lianas ed that in Israel, septate fibres occur most versus 12% worldwide), and rays three or frequently in climbers. Wolkinger (1970) and more cells wide (60% trees, 86% lianas ver Fahn (1982) also found living protoplasts in sus 39% worldwide) are greater for Bigno both non-septate and septate fibres, which niaceae than world woods. are probably longer-lived in the septate fibres allowing sufficient time for septa to develop. Rays of two distinct sizes were found in Acknowledgements 63% of lianas and only 5% of trees. Rays are D.F. Cutler, P.J. Rudall, E.A. Wheeler wider, taller (usually over 20 cells high, and and G.D. Wallace suggested numerous im often much taller) and more heterocellular in provements to the manuscript. Angela Pear lianas than in trees. Large heterocellular rays son and Alison Hughes sectioned much of are considered by Carlquist (1962, 1988) to the material. M. Simmonds provided the be a juvenile characteristic. All the lianas we statistical analyses. R. Brummitt, D. Hunt, examined were from relatively small stems M. Wilmot-Dear, S. Bidgood and A. Gentry up to about 5 cm in diameter, so there is a advised on the many problems associated strong possibility that this influenced in part with the taxonomy and nomenclature of the the high incidence of heterocellular rays in family. the lianas. It seems likely that some lianas This work was supported in part by a have large rays because they receive support Faculty Grants Award from Millersville Uni from other plants, and do not need a high versity and a sabbatical leave to David R. proportion of fibres to support them. Another Dobbins. possibility is that rays may be important in accommodating the physical torque and stress placed on climbers, as shown by Fisher and Ewers (1989). Baas, P. & F.H. Schweingruber. 1987. Eco logical trends in the wood anatomy of 3) Features equally common in trees and trees, shrubs and climbers from Europe. lianas - There are several features in this IAWA Bull. n.s. 8: 245-274. category, i.e. distinct growth rings (43% v. Bamber, R.K. 1984. Wood anatomy of 32%), diffuse-porous (97% v. 94%), vessels some Australian rainforest vines. Pro not solitary, simple perforation plates (all spe ceedings of Pacific Regional Wood Anat cies), alternate intervascular pitting and ves omy Conference October 1-7, 1984, sel-ray pitting of the same size (all species), Tsukuba, Japan: 58-60. similar vessel element length and paratracheal Carlquist, S. 1962. A theory ofpaedomorpho parenchyma. Fibre wall thickness is similar sis in dicotyledonous woods. Phytomor in the two groups, although none of the lianas phology 12: 30-45. have very thick-walled fibres whereas 1% of Carlquist, S. 1985a. Vasicentric tracheids as a the trees do. drought survival mechanism in the woody The frequency of some of these characters flora of southern California and similar has been compared with the same characters regions; review of vasicentric tracheids. in 4,080 woods from many families world Aliso 11: 37-68. Downloaded from Brill.com10/04/2021 01:59:21PM via free access 414 IAWA Bulletin n.s., Vol. 12 (4),1991 Carlquist, S. 1985b. Observations on func Gentry, A.H. 1979. Distribution patterns of tional wood histology of vines and lianas: neotropical Bignoniaceae: some phytogeo vessel dimorphism, tracheids, vasicentric graphical implications. In Tropical Botany tracheids, narrow vessels, and parenchy (eds. K. Larsen & L.B. Holm-Nielsen): ma. Aliso 11: 139-157. 339-354. Academic Press, London, New Carlquist, S. 1988. Comparative wood anat York, San Francisco. omy: Systematic, ecological and evolution Gentry, A.H. 1980. Bignoniaceae. Part 1 ary aspects of dicotyledon wood. Springer (Crescentieae and Tourrettieae). Flora Verlag, New York. Neotropica Monograph 25. Chalk, L. 1983. Wood structure. In: Anatomy Gregory, M. 1980. Wood identification: an of the Dicotyledons, 2nd Ed. Vol. 2, Ox- annotated bibliography. IAWA Bull. n. s. ford University Press. 1: 3-41. Cronquist, A. 1981. An integrated system of Handa, T. 1936. Abnormal vascular bundle in classification of flowering plants. Colum the stem of Campsis grandiflora K Schum. bia University Press, New York. Japan. J. Bot. 8: 47-58. Diniz, M.A. 1988. Family 124: Bignonia IAWA Committee. 1989. IAWA list of mi ceae. In Flora Zambesiaca. Vol. 8 Part 3: croscopic features for hardwood identifica 61-85. tion with an appendix on non-anatomical Dobbins, D.R. 1971. Studies on the anoma information.IAWA Bull. n.s. 10: 219- lous cambial activity in Doxantha unguis 332. cati (Bignoniaceae). ll. A case of differ Jain, D.K. & V. Singh. 1980. Studies in ential production of secondary tissues. Bignoniaceae. VIT. Wood anatomy. Pro Amer. J. Bot. 58: 697-705. ceedings of the Indian Academy of Scien Dobbins, D.R. & J.B. Fisher. 1986. Wound ces. Plant Science 89: 443-456. responses in girdled stems of lianas. Bo Liben, L. 1977. Bignoniaceae. In: Flora tanical Gazette 147: 278-289. d' Afrique Centrale (Zaire, Rwanda, Bu Ewers, F. & J.B. Fisher. 1989. 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Springer Verlag, IAWA Bull. n.s. 9: 203-252. Berlin, Heidelberg, New York, Tokyo. APPENDIX The species examined are listed below. The country or geographical region of origin is that from which the specimen came, not necessarily its native habitat. If the exact source of the specimen is not known, but the native region is, this is in parentheses. Synonymy is taken from Diniz (1988), Gentry (1973, 1977, 1979, 1980), Perrier de la B§.thie (1938), Sprague (1906), Liben (1977), Van Steenis (1977), and herbarium specimens at Kew. The tribal affinity for each genus is taken from Gentry (1980). TREES: Coleeae: Kigelia africana, Lw (Lewalle 5327, Bu 38355; E. latifolia, (New World), MADw rundi), KJw Nigeria, Kw 13640, K. africana 16510. - Parmentiera cereifera, (New World), (labelledK.pinnata) Mozambique, Kw 13643, Kw 13675; P. macrophylla, Malaysia, SJRw K. africana (labelled K. pinnata) S Africa, 38465. Uw 22020, K. africana (labelled K. aethio- N.B.: Enallagma may be considered part of pum) Ethiopia (Eritrea), Kw 13639. - Ophio- Amphitecna. coleafloribunda (labelled Coleafloribunda), Madagascar, Kw 13578. - Rhodocolea tel- Oroxyleae: fairiae (labelled Colea teljairiae), Madagas Millingtonia hortensis, Burma, KJw 3645, car, SJR w10766. Thailand, Lw, Geesink 3427. - Oroxylum indicum, Philippines Kw 13669, Singapore Crescentieae: Kw 13668. Amphitecna latifolia (labelled Crescentia cucurbitina), USA (Florida), Kw 13585. - Tecomeae: Crescentia alata, Mexico, SJRw 9599; C. cu- Catalpa bignonioides, N America, KJw, jete, Guyana, Kw 13580; C. cujete (labelled no data; C. bungei, China, SJRw 29857; C. C. acwninata), Jamaica, Kw 13579. - Enal- denticulata, Cuba, SJRw 19997; C. longis- lagma donnell-smithii, Costa Rica, SJRw sima, Dominica, Kw 13570; C. longissima Downloaded from Brill.com10/04/2021 01:59:21PM via free access