Liana Biomass and Leaf Area of a "Tierra Firme" Forest in the Rio ...

Liana Biomass and Leaf Area of a "Tierra Firme" Forest in the
Rio Negro Basin, Venezuela
Francis E. Putzl
Section of Ecology and Systematics, Cornell University, Ithaca, New York 14850, U.S.A.
ABSTRACT
In evergreen tropical rain forest, growing on a nutrient-poor Oxisol (lateritic soil) near San Carlos de Rio Negro, Venezuela, the
average total above-ground dry weight of lianas was 15.7 t ha-I which was 4.5 percent of the estimated above-ground forest
biomass. The average leaf area index of lianas was 1.2 M2 m-2 and constituted 19 percent of total forest leaf area.
In comparison to trees, liana stems are small in diameter relative to the sizes of their crowns. This difference in allometry occurs
because lianas rely on other plants for support and liana stems generally have large diameter and thus efficient xylem vessels. By
maintaining the conducting capacity of their vessels for longer periods of time than do trees, lianas are able to increase further
their leaf biomass-stem diameter ratios. Increases in cross sectional areas of liana stems are associated with proportionately larger
increases in leaf biomass than comparable increases in cross sectional areas of tree stems.
LIANAS ARE WOODY CLIMBING PLANTS that typically rely on suggest an average for tropical forests of approximately
other plants for mechanical support. They are often strik- 450 trha-' (Lieth 1975).
ingly apparent in tropical forests (Spruce 1908, Richards Liana abundance and size-class distribution were es-
1952, Longman and Jenik 1974), but little quantitative timated by species in 20 randomly located 100 m2 cirinformation
exists on liana abundance. The present study cular sample plots. In each plot I measured the dbh (direports
liana biomass and leaf area in a "tierra firme" ameter at 1.3 m or above the buttresses) of every tree
(never flooded) forest in the Rio Negro Basin near San and liana >2 m tall, determined the species of each liana,
Carlos de Rio Negro, Venezuela; Jordan and Uhl (1978) and recorded the sizes of the trees upon which the lianas
gave estimates of tree biomass in the same area. Data grew. Lianas

TABLE 1. Lianas used in biomass regressions.
Stem
diam-
eter
Species Family (cm)
Abuta rufescens Menispermaceae 12.0
Abuta rufescens Menispermaceae 8.9
Apocynaceae sp. Apocynaceae 1.2
Cayaponia coriaceae Cucurbitaceae 2.4
Coccoloba sp. Polygonaceae 2.3
Derris sp. Leguminosae 0.9
Doliocarpus dentatus Dilleniaceae 8.3
Doliocarpus dentatus Dilleniaceae 3.6
Gnetum nodiflorum
G. schwackeanum
G. schwackeanum
Gnetaceae
Gnetaceae
Gnetaceae
1.2
5.2
8.2
Icacinaceae sp. Icacinaceae 2.6
Menispermaceae sp. Menispermaceae 3.4
Peritassa laevigata Hippocrateaceae 9.1
Peritassa laevigata Hippocrateaceae 4.1
Pseudoconnarus macrophyllus Connaraceae 1.8
Pseudoconnarus macrophyllus Connaraceae 2.7
High liana species richness and rarity of many liana
species made it impractical to develop allometric regrestwo,
6 carried three, 4 carried four, 0 carried five, 3
sions for each species. Consequently, I developed a mixedcarried
six, and 1 carried seven lianas; comparing the
species regression based on a total of 17 individuals repobserved
between-tree distribution of lianas with Poisson
resenting the 12 most common species (Table 1).
probabilities, it is clear that trees with at least one liana
have a higher than random probability of having more
RESULTS AND DISCUSSION
than one liana (x2 = 31.9; P < .001). Trees of some
species are particularly prone to liana infestation, and sus-
In 20 sample plots covering a total of 0.2 ha I encoun- pended lianas resemble trellises that provide ready access
tered 45 species of lianas (Fig. 1). An average of 34.5 to the canopy for other lianas.
individuals (SD = 9.8) of 11.4 species (SD = 2.2) grew For the purpose of predicting total liana above-ground
in each plot; 39 percent of these were self-supporting dry weights I chose the logarithm (log,0) of basal area as
(upright). Self-supporting lianas constituted 20 percent the independent variable because it is the most convenient
(SD = 9) of the total number of woody plants

F 100
log y =0.12 +0.91 log x
R2 0.8185
the Oxisol site in San Carlos is an Amazon caatinga (heath)
forest on a well developed Spodosol (tropaquod). This
forest has a lower canopy and lower species diversity than
the Oxisol site and is characterized by trees with sclerophylls
and a paucity of lianas (Klinge in press). Davis
and Richards (1934) observed a similar lack of lianas in
=0 _
F-0~~~~~~~~~
_>/
_ 0
0@
heath forests in Guyana; Richards (1936) found a similar
contrast in abundance of lianas in heath forest growing
on white sand and in mixed forest growing on clay loam
in Borneo. This contrast, as Richards (1952) suggests,
may be due to xeric conditions in the understories of heath
0~~~~
forests. However, lianas are often common in forests which
suffer pronounced dry seasons (Holdridge et al. 1971).
m 0 /
Low soil fertility may also partly explain the paucity of
lianas in heath forests (Janzen 1974), but, surprisingly,
10 100 the analysis of the Oxisol and of the Spodosol from San
BASAL AREA LIANAS (cm2) Carlos showed little difference in the concentration of ma-
FIGURE 2. Total above ground dry weight of lianas as a
function of stem basal area at 1.3 m from the ground (logl0).
jor nutrients. The dependence of lianas on treefall disturbances
(Webb 1958, Putz 1982) suggests that the apparently
lower tree mortality rates on the Spodosol (pers.
obs.) may be a factor contributing to the lower abundance
of lianas. High frequency of disturbance may be the exleaf
weight ratio, weighted on the basis of species biomass planation for the super-abundance of lianas observed in
in the 20 sample plots, was 118 cm2 g-1. Jordan and Gabon (Hladik 1974) in a forest well known for its
Uhl (1978) reported a figure of 65 cm2 g- for trees in elephants (E. G. Leigh, pers. comm.).
the same forest.
Allometric relationships between stem cross sectional
Wood densities of the 45 liana species encountered area and leaf biomass (or leaf area) for trees and lianas
varied from 0.31 to 0.95 g.cm-3. Average wood density, are very different; total dry weight of leaves increased
weighted on the basis of estimated species biomass in the much more rapidly with stem basal area in lianas than in
sample plots was 0.48 g-cm-3; Jordan and Uhl (1978) trees (Fig. 3). Liana stems can be small in diameter relreported
a value of 0.96 g cm-3 for average tree wood ative to the amount of foliage supplied (Schenck 1892,
density in the same forest. Liana abundance varies from Schimper and von Faber 1938, Schnell 1970) because
forest to forest but in a "tierra firme" forest in Brazil, a lianas do not support themselves mechanically and beseasonal
evergreen forest in Thailand, and a rain forest in cause lianas generally have very efficient xylem vessels.
Ghana, lianas were similar in their abundance to the Ox- For a tree to avoid buckling under its own weight, each
isol site in Venezuela (Table 2). Immediately adjacent to height or weight increment must be balanced by a pro-
TABLE 2. Biomass and leaf area index (LAI) of lianas and trees in several tropical forests.
Total
above-ground
dry-weight
including Liana
lianas dry-weight LAI
Location Forest type (t ha-') (t ha-1) Trees Lianas Reference
Brazil Tierra Firme 687a 46^ Klinge and Rodrigues
1974
Thailand Seasonal Evergreen 403 20 7.4 3.3 Ogawa et al. 1965
Ghana Tropical Rain 308 14 Greenland and Kowal
1960
Venezuela Amazon Caatinga 309 2.1 Klinge, in press
Malaysia High Dipterocarp 475 9 7.3 0.7 Kato et al. 1978
Venezuela Tierra Firme 351 15 5.2 1.2 Jordan and Uhl 1978;
this study
a Fresh-weight.
Liana Biomass and Allometry 187

12
10_.
I-
YIianas= 0 109x - 0.376
0.008
Ytrees =1.368 + 0.018x
0.004
/
I
I
area of even large lianas is functional xylem. It is unlikely
that the lack of non-conducting xylem is due to the lianas
growing more rapidly in diameter than trees because the
average annual stem diameter growth rate of 50 canopy
lianas monitored for 1.5 years in San Carlos was approximately
0.5 mm. This extremely slow growth rate coupled
LL
8_/
with the lack of heartwood formation suggests that xylem
vessels in lianas maintain conducting capacity for decades.
06
IL
LLi
J4
A
A
A
To remain functional for such long periods of time,
liana vessels either do not cavitate (develop embolisms)
or somehow cavitated vessels are refilled. Considering that
most lianas grow on top of trees (i.e., in full sun) and
generally have large thin leaves, liana stems must expe-
C] - 6
02
H-
/A
A
rience high transpirational demands and consequently high
risks of vessel cavitation. Furthermore, liana stems are
S A
often physically jostled about; such mechanical distur-
0
20 40 60 80 100
BASAL AREA
120 140
bances may also result in vessel cavitation. Thus it seems
more likely that cavitated vessels in lianas can be repaired
rather than avoid cavitation in the first place.
FIGURE 3. Total dry weight of harvested trees (triangles) and Root pressure (see Stocking 1956 for a review) may
lianas (dots) plotted against stem basal area at 1.3 m from the provide the necessary force for refilling cavitated xylem
ground.
vessels. The first published account of root pressure was
a description of the phenomenon in grapevines ( Vitis sp.;
Hales 1727); xylem vessels in grapes are known to refill
in the spring with water pumped up from the roots (Schoportional
increase in stem basal area (Greenhill 1881, lander et al. 1955). Positive xylem pressures have also
McMahon 1973). When xylem loses its conducting cabeen
reported for several tropical lianas (Scholander et al.
pacity, it still serves to help mechanically support the tree; 1957, 1961). Some temperate herbs refill cavitated vesa
balance is thus maintained between requirements for
sels during the night when root pressure forces stem water
support and supply. Xylem vessels in trees retain their
potential positive (Milburn and McLaughlin 1974). The
conducting capacity for a few years at most (Zimmer- hypothesis that root pressure serves to refill cavitated vesmann
and Brown 1971); this results in a band of consels
in lianas and thus prolongs their functional life awaits
ducting wood (sapwood) surrounding a central core of
experimental examination.
nonconducting heartwood. A large portion of each growth Lianas are abundant in the tierra firme forest of San
increment in tree stems thus serves to replace noncon- Carlos de Rio Negro where they contribute substantially
ducting vessels (Preston 1958). Because lianas rely on to total forest leaf area. Per unit cross-sectional area, liana
trees for support, their stems need not be resistant to stems support more leaf weight than trees. This is because
bending. Small diameter liana stems can supply water lianas generally have large diameter and long functioning
and mineral nutrients to many more leaves than tree stems
xylem vessels.
of similar diameter partially due to the extremely wide
diameter xylem vessels found in many liana stems
(Carlquist 1975, Ayensu and Stern 1964); flow rates ACKNOWLEDGMENTS
increase with the fourth power of vessel diameter (Poi- Financial support for this study was provided by the National
seuilles's Law; Zimmermann 1978). The steepness of the Science Foundation and the Palm Society. I wish to thank P.
basal-area-leaf-weight relation in lianas (Fig. 3) suggests L. Marks, C. Uhl, C. F. Jordan, and E. G. Leigh, Jr. for their
that in contrast to trees, each diameter increment in liana
generous support and numerous helpful suggestions. Howard
and Kate Clark were both gracious hosts and helpful in the
stems allows a proportionate and large increase in the field. Contribution number 3845, Journal Series, Institute of
amount of leaf biomass supplied. Preliminary dye move- Food and Agricultural Sciences, University of Florida.
ment experiments indicate that much of the cross sectional
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