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. Dobbins 2

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.

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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

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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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392 IAWA Bulletin n.s., Vol. 12 (4),1991

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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 393

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394 IAWA Bulletin n.s., Vol. 12 (4),1991

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Gasson & Dobbins - Trees versus Hanas in Bignoniaceae 395

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396 IAWA Bulletin n.s., Vol. 12 (4),1991

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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 397

Trees by Tribe: See Table 1. cereifera, and one to several small rhomboi-
dal crystals in some unchambered ray cells of
C. alata (MADw 42263). Axial parenchyma
Tribe Coleeae: Kigelia, Ophiocolea, Rhodo-
and rays usually storied in Amphitecna and
colea. See Table 1.
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: Millingtonia, Oroxy1wn. See
clusters and radial multiples up to 6. .
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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398 IAWA Bulletin n.s., Vol. 12 (4),1991

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= ~eo ~
;
= :a'" li '"~ £ <: ~
'::1
'" '" .;; '"01) " "S .S:! B
~ 0;"
N
.S

l=
.~ :I
= 'S £ S
"0 .S ~
'":I 0 S
S os. £'C:
'"01) '"01) '"eo '":I
·c= ·c= ·c= ... :§ 8-
~ ~ 9 £ B

~
~ ~ S
[
01)
0 '"~
"'">
~
~ ~ £' = 0

"'"
>
£ £ £ '"
·s ...
.~
OJ
~
l e'" e'" ]'" or>e
0;
.a'" ~ '"
~ ~
~ ~ be
"> ~ '"
~
~ ~ ·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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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 399

(Table 1 continued)

~<>
g
os
e ] 01
.5 5
1 e ...
>-
-5
=
01 1:e e::>
2l
1
..e
~--.
.~
! '"

;;
~ ~ >0
~ 1 .<: e
]
= ... I
0)",

~ ~ ~ ~ ~ ~
..e
::>
0
'" 6';;'
'"e>- :3
.<:
'"~ .~
g
~ --. [ ~<> ~<> <l B.g
--.::>

I
01 0) 0
2
~ '" .... .... g 0

...
~
>- <>
e ZbO ~
1
.5 .5 'E .5 C e ..". -5 gj'~

"" a ..:::> 01 ~ ~"

I
'3'" '3'" ] ~ = 0)
"0 0) V 0 N :s!
] 8~
..El a = .5 's
bO .<: 0
0)
~ ?;- .<:

eeee
~

:e ~ s =<> ·s::> :5 =os a e e'">-

'"~ '"~ g .~ ''::: '" '"
0; 0;
:s!
.~
0)
>- >-
<> <> '"
~
>
N
~
0 ,rJ ,rJ e ~ ~
+ + + + 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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400 IAWA Bulletin n.s., Vol. 12 (4),1991

(Table 1 continued)
~ ~...
-5
.,
~...
B
~
00

u ..@ §
g '.:l= C., .,'"
'"00
., c.,
B :a ~
~
~
'.:l
:a'" '"::s .~ e 00 .Sol ~
B
'" e ~ ~
N
.~ .~ ~
-5 .Sol S
~ .5 ~

i ""
'" e
'"bO '"00 '"bO '"::s
= '1:= '1:= ~ ....~0
~
~ it ~.,>'" ~.,>'"
's. g -5
S
S
~ fr 0C:
III
'1:
~bo '9= .E'"
bO ~
.,'" ~ =
-5
~
-5 -5
~ ~
= 's.,'" ;a
'"
<l
.,>'"'"
0;
.Sol
<l
] .,= §.
.,>'" e e ~ '"
~
> '"
'"~ ]'" '"
~
~ ~ ~ .;:: .<:: .5 <§ <:::
.0
0=

Godmania aesculifolia + + 93 283 3.6 + +

Jacaranda cf. caucana + + 7 76 298 6.1 + +
caucana + + 21 59 336 6.1 + +
coerulea + + 11 80 316 4.7 + +
copaia + + + 4 177 346 4.4 + +
cuspidifolia + + + 80 320 7.4 + +
glabra + + 175 331 5.2 + +
hesperia + + + +
micrantha + + 19 89 324 7.2 + +
mimosifolia + + + 32 58 304 7.2 + +
obtusifolia + + + 18 89 342 6.7 + +
No. of species 10 7 6 1 10 10 10
Index 0.7 0.60.1 1
Markhamia acuminata + + 48 198 3.3 + (+) +
pierrii + + 99 250 3.3 + +
platycalyx + + 13 273 3.3 + +
sessilis + + 14 119 210 3.3 + (+) +
stipulata + + 124 292 4.4 + +
zanzibarica + + + 65 193 3.3 + +
No. of species 6 6 3 1 3 6 2 6
Index 0.5 0.2 0.5 0.3
Mayodendron igneum + + + +
Newbouldia laevis + + 50 278 4.1 + +
Pajanelia longifolia + + 4 160 356 8.8 + (+) +
Paratecoma peroba + + (+) 87 240 5.0 + +
Phyllarthron bojerianum + + + +
Radermachera gigantea + + 60 255 4.1 + +
glandulosum + + 12 88 272 4.4 + +
pinnata + + 103 302 3.6 + +
ramiflora + + 59 243 3.3 + +
sinica + + 11 59 321 3.6 + (+) +
No. of species 5 2 3 5 5 1 5
Index 0.4 0.6 1 1 0.2
Romeroa verticillata + + + (+) +
Spathodea campanu/ata + + 10 71 231 5.0 + (+) +
se"atu/a + + + 14 103 337 4.1 + (+) +
No. of species 2 2 1 2 2 2 2
Index 1 0.5 1 1 1 1
Stereospermum fimbriatum + + 5 117 258 3.6 + (+) +
neuranthum + + 7 141 240 3.6 + +
personatum + + 7 119 229 5.3 + +
xylocarpum + + + 8 73 283 3.9 + +
No. of species 4 2 2 1 4 4 1 4
Index 0.5 0.5 0.3 0.3

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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 401

(Table 1 continued) .,
~
os
u g
e>- ~ O!

1
c::
g ., ] O! .~
e
e .S e
., ! 'B
8
os ] e... ~
.c::
g
'" ~
:::l .....
.!I .!I ~e 0 .~'" .,
~O'
"'til

e1
~
.s:: .E ~
!5';'
9.,
~
~ ~ :a [ ~ ~ =
.e '" '0 ~
g .~0
g j e ""
'il ...0 0'6
.--. g
i """'-6
>- u

1
.S
., .S
., '5 .S C '"V '"A
~
-5 .<: ~ ;g .~

., .,'" . .
~ c::
'"00 "0 '" I
:::l j 8~
S., S., ] '" '" .a
.l:l
~ c
.,.,c:: .~'"00 to::0c::'" ·s ~ -!3osc:: ~
.c:: .c:: N '0.,
eee e ee
~
:::l·s
~ e-
u
~
9., .S ~
> :a fa
~ .9 u
'"
~§ .0 .0
>- >-
e>- >-
~ ~
+ + + + + + + (+) 1-3 12
+ + + + + 1(2) 20
+ + + + + (+) 1 35
+ + + + + 1 20
+ + + + (+) (1)2-3 46
+ + + + + 1-2 25
+ + + + (1)2-3 25
+ + + 1 30
+ + + + 1-3 20
+ + + + + + 1 25
+ + + + + 1 25
7 9 10 5 3 8 2 3 1
0.7 0.9 1 0.5 0.3 0.8 0.2 0.3 0.1
+ + + 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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402 IAWA Bulletin n.s., Vol. 12 (4),1991

(Table 1 continued)
~ ~
~
-5toO
.,
B
~
.. . . .~. .. .
'G .§ ]
~
'G c
.::: ., ;:.,
B :atoO ~
:a 'i5 :g -=~
. . ., ., l . ... ....,
.:::c toO
.,
S £
~
c
.~
.
.~ :I S

l
"0 .5 0
S <l 'f! .~
's. fr
toO
c
toO
c
toO
C :I
8-
~ .~
S
11., 11.,., 11.,.,
S -5
C: C: 3
.;::
[ .t;:
.;:: .;::
., toO
c
.,
0 -5 :s., .,> .,~ .,c0
~
.,.,> ~.c-=., i.,.,> .,'S"
-5 -5 -5 ~ >
.,
<l
~
~ ~ ~ '2
~ ~ 00 's.,., :a6 c c
~
S
~
.5
~
.r>
<:::=
1S<:::= ~
.r>
<:::=
~
.r>
..::

Tabebuia angustata + + 64 259 3.3 + +

barbata + + 106 219 4.1 + +
capitata + + 89 258 5.2 + +
caraiba + + 24 98 254 4.4 + +
cassinoides + + 70 314 3.3 + +
cf. catinga + + 71 298 5.8 + +
chrysantha + + 47 210 6.3 + +
chrysea + + 20 48 228 4.1 + +
donnell-smithii + + 7 111 288 5.0 + +
guayacan + + + 31 220 4.1 + + +
haemantha + + 59 241 3.9 + +
impetiginosa + + 53 59 216 7.1 + +
insignis + + 140 340 4.1 + +
pallida + + 131 300 3.3 + +
rosea + + 12 125 349 + +
roseo-alba + + 57 282 4.1 + +
serratifolia + + 8 79 299 4.4 + +
stenocalyx + + 18 76 351 3.6 + +
No. of species 18 3 15 1 18 13 6 18
Index 0.2 0.80.1 0.70.3

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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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 403

(fable 1 continued)
'"
=ij
u g
·t
os .!!l
e>. <a
'8
to
..c 13
] e
1
u
.~
'" ] e
...e ....0
li! li! 1:
B ~ ~ ::I

t g ...-:a
] ..c
g e 1\)'"
] e
...'"~
~ ~
..c oS 15';;;"
=ij ~ ....
.~
'" ~ ::I '" ,......::1
e- ~
S
'">. [
.!!l
;; ~
...,u ...,u
0
e
~
,§. e,~
e ..,.I
~
.s .s
,g ~
u 0

e ~ .<:01) ~
~
i ·E .s ~ c -5 [g ••
.... as
s'" '"! ""
"'-
""~
I\)
V II ·0
5 ~ I\)

] 1! <'I :s! ::1'9

'3
·s~
O/) ::I
C;- ~ 0
·c~::I :5
c
'"i!- i!- .~
u u
S
~
.5
.l:l .E
'"~ ~ a~ I\)
N
..9
.s
~
..:::C
0
~
a .r>a
.r>
~
>. >.
0;

e e- e- e e e e .g
'">. '" >. '"
>.
0; ~
:s!
'" U

+ + + + + 1-2 10
+ + + + + 1-2 8
+ + + 2 11
+ + + + + 1 13
+ + + 1 11
+ + + + (1)2-3 12
+ + + + (1)2 9
+ + + 2-4 15
+ + + + (1)2-5 21
+ + + + + + 1-2 9
+ + + + + + 1-2 10
+ + + + + 2 12
+ + + + + 1-2 14
+ + + + + + 1-2 13
+ + + + + + 1-3 15
+ + + + + + + 1-2 15
+ + + + + + (1)2 10
+ + + + + 1-2 20
1 12 1 2 11 6 11 16 10 17 1
0.1 0.7 0.1 0.1 0.6 0.3 0.6 0.9 0.6 0.90.1

+ + + + (+) 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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404 IAWA Bulletin n.s., Vol. 12 (4),1991

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
! ..,
£

-u
g
.=c :a.=., C.,.., .,:s
.,
1;l
., ..,
~
Jj
c..,
..,S
~
:a .5 i
bI) bI)
c S
.,:s
.0; N
·2 S .~
., ., 0 .~
<l
i"".,
., .,:s .s,
bI) bI) bI)
~ e ~ ~ S
i.,..,
·cc ·cc ·cc
[
bI)
8. '-
0
., ~ 8- .,
<l
.,..,
·cc
i.,..,
£ £ £ .,b ~ >
<l
.,., Ol t
~ ~ ~ 00 .~ 6 .., .>l ..,
cto
~ £
~ ~ ~ ·8 ., :a >
<l
.c > S S .5
Bignonieae
Adenocalymna alliaceum + + 85 55 232 4.7
Anemopaegmo. carrarense + + + 77 69 295
robustum + + 49 39 366 6.1
No. of species 2 1 1 2 1
Index 0.5 0.5 0.5

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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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 405

(Table 2 continued)

=..,u'" §:
~ as.. (;I 5
~
'5 .~
e
e::0
..,~
~ B 1.c ~::0 ~::0 e ..,'" i)

i
-5 23 'Cl ';(
6
'"e>- =..,
arg
<II
.~
B
E ~ ~
g g '"~ ,§,
= s 0
§ ..,
~ fr .S e0 J:
c:: !a.., "<t
-5
on ~
'"
'"'" '"
!'l '3'"e;- '".~ ~
::0 -; .c ..c:: A
:s!
.c 'Cl 8
e e
.;:: ~ :s!
§ .~ '">- '" '" '">-
~
<:=
~
<:=
.c
<;:: u
B ~
0
u :5 e e>- >-
e e
u
.g
'"
+ + + + + 1-4 180
+ + + + 1-10 ISO + E
+ + + + 1-6 230 (+)
2 2 2 2 2
1 1 1

+ + + + + 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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406 IAWA Bulletin n.s., Vol. 12 (4), 1991

(Table 2 continued)

~ ~...
.s00
~
B
1:) ] §"
1:)
c
c
'::1 C '"~ !a C :a00
'::1
:a.S'" " '" '"c00 "e "e
'" '0; N
e :g
.~
"t:I ~ e'" '" 0 '8
e... "
<l
.:3C
's..
l
'" ]
'"00 '"00 '"
00
~
'"'" 00 ~
....0 i <l
c
.1::
c
'1::
c
.1:: e .1::c :§ 8- '"~ '"'" <l
.s~ .s~ .s~ 8. .§ '"
<l O! ~ "c> '"
"'"1::
be ..s
'"
.... '" ~
!;l c
"e'"
i'j B
~ ~ ~ ..:c "'" :a "'"> ..c ">'" e .S

Pleonotoma clematis + + +
Potamoganos microcalyx + + + 21 157 355
Pseudocalymma alliaceum + + + 28 127 363 6.6
Pyrostegia lIenusta + + + 85 62 304 5.6
Saritaea magnifica + + + 72 63 243 4.0
Tanaecium jaroba + + + + 28 114 265 6.1
Tynnanthus elegans + + + 48 177 247 4.7
micranthus + + + +
No. of species 2 2 1 2 2
Index 1 0.5 1 1

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.

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Gasson & Dobbins - Trees versus lianas in Bignoniaceae 407

(Table 2 continued)

=.,'" @-

I
u
13 <a
:a 13
.~
.5
~ e

i
~
..:. £ ] S S ....e0 '" .,'" i)

i
.~
-5
9
E
'"e
>.
..c:
~
.E
;;
g
=
.,'" '"~
e
N
.;;;
§
~

[ ~.,
0
C .5 ;:., e
~
~
~ 0 '<t
-5
:9 .~ ....~0 ~
g
e '3'"e;- "".~9
<:

e'" e'"
::;.
§ '"
0:;: <a
.':::
~
'">.
..c:
'"~
A
'"
~ ..c:
'">. :9
..c:
<C
..c:
<C
..c:
'I: U
'"
~
<:
0
U :5 e .... e- e- e>. e .8
+ ? ? + + 1-3 40
+ + + + 1-5 400 +
+ + + + 1-5 290 (+)
+ + + + + 1-3 100
+ + + + + 1-2 65
+ + + + + + 1-7 90 +
+ + + + + + + 1-6 180 +
+ + + + + 1-4 55 +
2 2 1 2 2 2 1 2
1 0.5 1 0.5 1

+ + + + + 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-

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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). Alifonn in 14 genera (Fig. 2). Con-
Lianas by Tribe: See Table 2.
fluent in 19 genera. Unilateral paratracheal in
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.

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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.

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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
1) Features restricted to or more common in
trees. Intervessel and vessel-ray pitting alter-
trees - Rays exclusively uniseriate (12%),
nate and of the same size, perforation plates
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 Campsis grandi-
semi-ring-porous or diffuse-porous (with
flora, which would be treated as a liana in
vessels of two distinct sizes) Tecomeae trees
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.

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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)
'<G)3
c.
Vl
.....0
30 o Trees & Shrubs

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
.~
u
G)
C.
r/J
.....0 30
G)
o Trees & Shrubs
OJ)
• Lianas
E 20
00

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).

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412 IAWA Bulletin n.s., Vol. 12 (4),1991

Percentage of species

40
o.
j
both wide and narrow vessels means that
even if some wide vessels become blocked
by embolism the narrower ones would re-
main functional. Vessel element length is
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- A 0
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,

0
P < 0.01). 140 A
0
0 0
A 0
120- J!' AD 0
A DO
0
grandijlora, Menispermaceae, Mennega 1982; A A Dr)

Zanthoxylum, Rutaceae, Caesalpinia and 100 A...•

A ~ AI.
Bauhinia, Leguminosae, Bamber 1984; Carl- AI. ..
0

quist 1985b; Ewers et al. 1990). Lianas also

have significantly more vessels per unit area
80
(t A
A
0

than trees (P < 0.0001). The relationship be- 0

DO
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=------'100
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: ANOVA
liana to transport water, and the presence of F = 51.6, P < 0.0001.

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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.
Acknowledgements
Rays of two distinct sizes were found in
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.

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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-
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Ewers, F. & J.B. Fisher. 1989. Variation in rundi), Belgium.
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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

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