Lianas and Trees in a Liana Forest of Amazonian Bolivia

Author(s): Diego R. Pérez-Salicrup, Victoria L. Sork, Francis E. Putz

Source: BIOTROPICA, 33(1):34-47.
Published By: The Association for Tropical Biology & Conservation
DOI: http://dx.doi.org/10.1646/0006-3606(2001)033[0034:LATIAL]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.1646/0006-3606%282001%29033%5B0034%3ALATIAL
%5D2.0.CO%3B2

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BIOTROPICA 33(1): 34–47 2001

Lianas and Trees in a Liana Forest of Amazonian Bolivia1

Diego R. Pérez-Salicrup2, Victoria L. Sork

Department of Biology, University of Missouri–St. Louis, 8001 Natural Bridge Road, St. Louis, Missouri
63121-4499, U.S.A.
and
Francis E. Putz
Department of Botany, University of Florida, 220 Bartram Hall, Gainesville, Florida 32611, U.S.A.

ABSTRACT
The distribution of lianas (woody climbing plants) on trees in a lowland ‘‘liana forest’’ of northeastern Bolivia was
clumped and varied with characteristics of individual trees and tree neighbors. In twenty-four 900-m2 square plots
established to estimate tree ($10 cm DBH [diameter at breast height]) and liana ($2 cm DBH) densities and to
count the number of lianas a tree carried, we estimated a mean of 65 tree species and 51 liana species per hectare.
Mean tree density at the study site (564 trees/ha, SE 5 23.7) was similar to other tropical sites, but mean liana
density was much higher (2471 lianas/ha, SE 5 104.3). Basal area of trees $10 cm DBH was low in Oquiriquia
(19.2 m2/ha) in comparison to other tropical forests. Liana diversity, as expressed by the ratio of liana/tree species,
was higher in this forest than in any other so far reported. Of trees $10 cm DBH, 86 percent carried lianas. Four
tree species (Astrocaryum aculeatum, Euterpe precatoria, Xylopia sericea, and Astronium fraxinifolium) had a lower pro-
portion of liana-infested individuals than expected based on the mean percent of liana infestation in this forest. Forest
plots with similar tree species composition did not have similar liana composition or liana loads per tree, which
suggests that lianas and trees have no specific associations with each other. Lianas showed an aggregated distribution
on trees, suggesting a facilitation process in which new lianas use already established ones to climb trees. Lianas of
four different climbing mechanisms climbed a similar number of trees. Plots in the forest with high palm density also
had high liana density, suggesting that palms and lianas respond positively to common forest conditions in the study
site (perhaps related to successional forest status). Larger-diameter trees carried more lianas than slender trees, but this
relationship was affected by the density of trees 10–30 cm DBH surrounding each tree, which suggests again that the
successional stage of the forest in which a tree grows affects the number of lianas a tree carries. We found little evidence
to support the idea that lianas were more likely to climb some tree species than others. Instead, larger trees and trees
growing in the vicinity of trees 10–30 cm DBH, tended to have more lianas, perhaps as result of longer exposure to
liana infestation.

RESUMEN
La distribución de lianas (bejucos leñosos) en los árboles de un ‘‘bosque de lianas’’ en el noreste de Bolivia fué agregada
y varió de acuerdo a caracterı́sticas individuales y del vecindario en que crecı́a cada árbol. En veinticuatro parcelas
cuadradas de 900 m2 establecidas para estimar las densidades de árboles ($10 cm DAP) y lianas ($2 cm DAP), y
contar el número de lianas por árbol, estimamos un promedio de 65 especies de árboles y 51 especies de lianas por
ha. La densidad promedio de árboles en el sitio de estudio (564 árboles/ha, EE 5 23.7) fué similar a otros sitios
tropicales, pero la densidad promedio de bejucos leñosos fué mucho más alta (2471 bejucos leñosos/ha). El área basal
de árboles $10 cm DAP fué baja en Oquiriquia (19.2 m2/ha) en comparación con otros bosques tropicales. La
diversidad de lianas, expresada como la razón de especies de árboles/especies de lianas leñosas, fué mayor que en
cualquier otro estudio reportado hasta la fecha. De los árboles $10 cm DAP, 86 por ciento cargaban lianas. Cuatro
especies de árboles (Astrocaryum aculeatum, Euterpe precatoria, Xylopia sericea, y Astronium fraxinifolium), tuvieron una
proporción menor de individuos infestados con lianas que lo esperado en base el porcentaje promedio de infección
de lianas en este bosque. Parcelas de bosque con una composición de especies de árboles similar no compartieron una
composición de lianas similar, ni una carga de bejucos similar por árbol, lo que suguiere que las lianas y los árboles
no tienen asociaciones especificas entre sı́. Las lianas presentaron un patrón agregado en los árboles, lo que sugiere un
proceso de facilitación en el que nuevas lianas emplean a otras ya establecidas para trepar en los árboles. Lianas de
cuatro mecanismos distintos para trepar subieron un número similar de árboles. Parcelas del bosque con altas densi-
dades de palmas mostraron altas densidades de bejucos, lo que sugiere que las palmas y los bejucos responden
positivamete a condiciones sucesionales del bosque similares. Arboles de diámetros mayores cargaban más lianas que

1Received 1 February 1999; revision accepted 29 February 2000.

2Current Address: Departamento de Ecologı́a de los Recursos Naturales, Universidad Nacional Autónoma de México,
AP 27-3 (Xangari), C.P. 58089, Morelia, Michoacán, México.

34

árboles delgados, pero esta relación depende del número de arboles 10–30 cm DAP en la vecindad de cada árbol, lo
que apoya la hipótesis que el estadio sucesional del sitio en que crece cada árbol afecta el número de lianas que éste
carga. Encontramos poca evidencia de asociaciones especı́ficas entre árboles y lianas. Tal parece que árboles de diámetros
mayores, creciendo cerca de árboles 10–30 cm DAP, cargan más bejucos, quizás como resultado de mayor tiempo de
exposición a ser infestados por bejucos.
Key words: Bolivia; liana distribution on trees; liana forests; lianas; liana–tree interactions; tropical forest.
LIANAS ARE WOODY PLANTS that cannot support
themselves in an upright position after they reach
ca 2 m in height (Putz & Holbrook 1991), and
must climb trees or other lianas to reach the top
of a forest canopy (Carlquist 1991, Gentry 1991).
Studies on the distribution of lianas, both on their
supporting trees and across forests, have docu-
mented that lianas have a clumped distribution at
both scales, but no previous study has quantified
the relative contribution of individual (e.g., size and
species) and neighborhood (e.g., canopy structure
and neighboring tree size) attributes in determining
the number of lianas a tree carries. Study of liana
distribution on trees is important because it could
help uncover patterns of association between trees
and lianas, and may clarify which factors enhance
or inhibit liana establishment and distribution
(Clark & Clark 1990, Campbell & Newbery
1993). In addition, understanding these factors is
essential to provide ecologically sound advice on
how to prevent and mitigate the deleterious im-
pacts of lianas on their supporting trees (Putz
1991).
Studies of liana distribution on trees have re-
ported fewer lianas on tree species with architec-
tural characteristics such as high tree flexibility,
long leaves, and high tree bole before first limb
(Boom & Mori 1982, Putz 1984a, Balfour &
Bond 1993, Campbell & Newbery 1993). Palms,
with high scores for these characteristics, seldom
carry lianas (Putz 1984a, Clark & Clark 1990). In
tropical forests, liana density and number of lianas
per tree increase in areas affected by treefall gaps
(Caballé 1986, Hegarty 1989, Hegarty & Caballé
1991). At larger geographical scales, whole forests
can become liana-infested following poorly man-
aged forestry operations (Putz 1991, Uhl et al.
1997, Vidal et al. 1997) or large-scale natural dis-
turbances (Webb 1958). Because liana densities
tend to increase following canopy disturbances, and
palms usually do not carry lianas, it is likely that
liana densities and number of lianas per tree would
be high in forest locations with a low proportion
of palms and a high proportion of canopy distur-
bance.
To understand the factors determining liana
Lianas and Trees in Amazonian Bolivia 35
abundance, two questions need to be addressed.
First, what is the relative importance of host tree
characteristics versus neighboring forest character-
istics in determining the number of lianas a tree
carries? Second, are some liana species more likely
to climb more tree individuals than others (Putz
1984b)? The answer to this latter question is im-
portant because the number of lianas a tree carries
may be determined not by tree characteristics, but
rather by the fact that a tree is located in an area
with high densities of a liana species that climbs
many trees. Because many lianas share the same
climbing mechanisms (Putz 1984b, Putz & Hol-
brook 1991), one can group liana species according
to their climbing mechanism and then ask if lianas
of different climbing mechanisms differ in the
number of trees they climb (Putz 1984b, Pinard &
Putz 1994). Thus, if climbing mechanism is im-
portant, the distribution of liana infestation may
be influenced by composition of lianas.
To describe the distribution patterns of lianas
on trees in a forest, it is essential to include a de-
scription of tree and liana communities at the site
of interest. Tropical forest types vary considerably
in liana density (Gentry 1991, Balfour & Bond
1993), and liana density could affect the distribu-
tion of lianas on trees. At the highest extreme of
liana densities are ‘‘liana forests,’’ a singular and
understudied vegetation type described as occur-
ring on the rim of the Amazon basin and in the
typhoon-affected forests in northeastern Queens-
land, Australia (Webb 1958, Pires 1973, Pires &
Prance 1985, Summerbell 1991). Because of their
high liana densities, liana forests provide ideal sites
to study the interactions between lianas and trees,
and the possible consequences of high liana density
on the tree community (Balée & Campbell 1990).
The objective of this study was to describe the
distribution patterns of lianas on trees in a lowland
liana forest of Bolivia. First, we determined the tree
and liana communities at the study site. Second,
we examined the idea that liana species are associ-
ated with individual tree species by evaluating: (a)
whether some tree species are more or less likely to
carry lianas than other tree species; (b) if forest
plots with similar tree species also share similar li-
36 Pérez-Salicrup, Sork, and Putz
ana species; and (c) if tree composition is associated
with mean number of lianas per tree (i.e., liana load
per tree) in forest plots. Third, we explored wheth-
er lianas have a random, clumped or even distri-
bution on trees. Fourth, we evaluated what forest
structural characteristics are associated with higher
liana densities, and assessed whether or not lianas
of different climbing mechanisms differed in the
number of trees they climbed. Fifth, we examined
the relative importance of individual tree diameter
versus neighboring site conditions around each
tree, to explain the number of lianas a tree carried.
STUDY SITE
This study was conducted in a 600,000-ha timber
concession located in the department of Santa
Cruz, Bolivia (148459S, 628009W). The forest,
named Oquiriquia, is situated at 200 m elevation
on the southernmost margins of the Amazon basin.
No meteorological station is located within 100 km
of the study site, but based on regional maps of
temperature and rainfall, mean annual temperature
probably ranges between 20 and 258C, and annual
precipitation between 1200 and 1700 mm (Sán-
chez 1997). Most rainfall in Oquiriquia occurs
from November through April, when portions of
the study area are flooded, and almost no precipi-
tation occurs from May through October (Pérez-
Salicrup 1999). According to the Holdridge vege-
tation classification system (Holdridge 1967), the
study area corresponds to a transition between dry
and moist forest. During the dry months, some tree
and liana species lose their foliage, so this forest is
semi-deciduous. Soils in this region are nutrient-
poor Ultisols derived from the Brazilian shield
(Guamán 1996). Canopy height in the study site
ranges from 15 to 25 m, with canopy emergents
of 35 m (Pérez-Salicrup 1999).
METHODS
DESCRIPTION OF TREE AND LIANA COMMUNITIES.—
From May through June 1995, we randomly es-
tablished twenty-four 900-m2 square plots in a 25-
ha area in Oquiriquia. We, however, excluded plac-
es where a canopy gap (canopy height #5 m) oc-
cupied $50 percent of a plot and sites that had
been affected by logging operations, because these
heavily disturbed places may have had unusually
high liana densities that could have obscured pat-
terns of association between trees and lianas (He-
garty & Caballé 1991). In each of these plots, we
measured diameter at 1.5 m or above buttresses
(DBH, diameter at breast height) of all trees $10
cm DBH and lianas $2 cm DBH. Liana stems can
bend and loop back to the ground and re-root, and
so the number of individual lianas can be easily
overestimated. To avoid this bias, we counted and
measured liana stems only once when it was ob-
vious that it was the same individual. We also
counted and measured all trees (.5 cm DBH in a
10- 3 10-m subplot located at the center of each
of the twenty-four 900-m2 plots.
Tree species composition was obtained by iden-
tifying all trees $10 cm DBH in the 24 plots men-
tioned above, but liana species composition was
based on lianas $2 cm DBH in a subsample of 12
plots. Tree and liana species were identified in the
field to the lowest possible taxonomic level and as-
signed morphospecies numbers. All morphospecies
were collected and taken for further identification
to the Museum of Natural History Noel Kempff
Mercado in Santa Cruz, Bolivia, and the Missouri
Botanical Garden (St. Louis, Missouri).
We constructed 25 species–area accumulation
curves for trees and lianas by randomly changing
the order in which plots were added to the curve
(analogous to a Monte-Carlo simulation). Then we
obtained a mean species–area curve for trees and
lianas 61 SE. We used species area curves to eval-
uate thoroughness of sampling effort and to esti-
mate number of species of trees and lianas per hect-
are. In addition, we calculated the abundance-
based (Chao et al. 1993) and the incidence-based
coverage estimator of species richness (Lee & Chao
1994) as two estimators of the true number of tree
and liana species at Oquiriquia using EstimateS V.5
(Colwell 1997).
To compare the diversity of trees versus lianas,
we contrasted the rate at which additional species
of trees and lianas were added to the sample. We
transformed mean cumulative number of species of
trees and lianas and area sampled to logarithmic
scales and tested if the slopes of cumulative number
of species were different. This last analysis was
based on 12 plots (total area 5 1.08 ha) so that
both life-forms would have the same sampled area.
Diversity of tree and liana species was also con-
trasted by plotting species–individual curves and
comparing the slopes of the lines for both life-
forms (Condit et al. 1996). We also calculated the
Shannon-Wiener diversity index for trees and lia-
nas.
ASSOCIATION BETWEEN TREE AND LIANA SPECIES.—We
counted the number of liana stems $2 cm DBH
that climbed each tree $10 cm DBH in 12 of the

900-m2 plots. Then we estimated the mean pro-
portion of trees with at least one liana and tested
whether individual tree species had a higher or low-
er proportion of liana-infested individuals than the
mean number of infested trees in the forest. To this
end, we estimated the expected proportion of in-
fested individuals per each tree species with ten or
more individuals within the 12 plots in which tree–
liana interactions were studied using a binomial
distribution (Sokal & Rohlf 1995), and then con-
trasted it with the observed proportion of infested
individuals using a goodness of fit chi-square test.
We evaluated whether or not plots in the forest
that had similar tree species composition also had
similar liana species composition by constructing
matrices of Sørensen’s similarity index (Beals 1984)
for tree and liana species and evaluating the rela-
tionship between these two matrices with a Mantel
test (Mantel 1967, Sokal & Rohlf 1995). We also
used a Mantel test to examine the relationship be-
tween the tree species similarity matrix and a ma-
trix of Euclidian distances between plots based on
the number of lianas per tree. Both Mantel tests
were done using PC-ORD V.3.14 (McCune &
Mefford 1997).
DISTRIBUTION PATTERN OF LIANAS ON TREES.—We ex-
amined the distribution pattern of lianas on trees
by comparing the observed distribution of frequen-
cies for trees with different numbers of lianas
climbing on them against the expected Poisson dis-
tribution using a chi-square goodness of fit test
(Sokal & Rohlf 1995). The observed frequency dis-
tribution of trees with different numbers of asso-
ciated lianas was the average number of trees in
each class of liana load within the 12 plots in which
number of lianas per tree was counted.
EFFECT OF TREE COMMUNITY CHARACTERISTICS AND
CANOPY STRUCTURE ON LIANA DENSITY .—We explored
the relative effect of five forest structural features
on density of lianas. For each plot, we estimated
the density of trees $5 cm DBH and then esti-
mated the proportion of trees 10–30 cm DBH (ex-
cluding palms), the proportion of trees $30 cm
DBH (excluding palms), and the proportion of
palms $10 cm DBH, out of the total number of
trees $5 cm DBH per plot. In addition, we esti-
mated the proportion of open canopy (i.e., visible
sky) and that of canopy #5 m. The proportion of
visible sky was measured by taking nine equally
spaced hemispherical photographs 0.4 m from the
ground in each plot. Each hemispherical photo-
graph was analyzed with the HemiView Canopy
Lianas and Trees in Amazonian Bolivia 37
Analysis program (Delta-T devices 1997), and the
mean of the nine photos per plot was used as an
estimator of canopy openness for each plot. Pro-
portion of canopy #5 m was estimated by standing
at 49 equally spaced points in each plot and mea-
suring the highest visible leaf in the canopy above
each point with a range finder. We analyzed the
relative effect of the above five structural variables
on liana density by conducting a stepwise regres-
sion analysis using SYSTATt 6.0 (SPSS 1996).
EFFECT OF CLIMBING MECHANISM ON THE NUMBER OF
TREES CLIMBED PER LIANA .—We evaluated if lianas
of four climbing mechanisms differed in the num-
ber of trees they climbed, by following individual
lianas from their rooting point and counting all
trees on which they climbed. Lianas were selected
outside of the 24 study plots mentioned above in
a stratified-random fashion, selecting individual li-
anas to include a range of 2–12 cm DBH. The
four climbing mechanisms were: (a) stem-twiners
(N 5 95 individuals), (b) branch-twiners (N 5
107), (c) tendril climbers (N 5 87), and (d) spiny
species (N 5 17). Each climbing mechanism was
represented by three liana species, except spiny li-
anas that were represented by two species. By virtue
of sampling design, the liana species sampled for
this part of the study represented the most abun-
dant species for each climbing mechanism among
all liana species at the study site. We conducted an
ANCOVA to compare the number of trees climbed
per liana across the four liana climbing mecha-
nisms, using liana DBH as the covariate. Liana
DBH and number of trees climbed per liana were
log transformed for statistical analyses.
EFFECT OF TREE DIAMETER VERSUS NEIGHBORHOOD
CHARACTERISTICS ON LIANA LOAD PER TREE.—We
conducted a contextual analysis (Boyd & Iversen
1979, Heisler & Damuth 1987) to evaluate the
relative contribution of tree DBH and site char-
acteristics on the mean number of lianas that a tree
carried. The first step in the contextual analysis was
to conduct an ANCOVA to compare the number
of lianas per tree between plots using individual
tree DBH as covariate. After plot and tree DBH
(but not their interaction) were found significant,
we conducted a multiple regression using tree
DBH and the same five structural variables per plot
used before as independent variables (see above).
The effect of each variable was then evaluated in
terms of its statistical significance and its relative
contribution to the model R 2. For this analysis, tree
DBH was transformed to logarithmic scale and the
38 Pérez-Salicrup, Sork, and Putz
FIGURE 1. (a) Species–area accumulation curves; (b)
log10 scale species–area accumulation curves; and (c) log10
scale species/individual accumulation curves, for trees and
lianas in Oquiriquia timber concession, Bolivia.
number of lianas per tree was square root trans-
formed.
RESULTS
DESCRIPTION OF TREE AND LIANA COMMUNITIES.—
Mean tree density was estimated as 564 (SE 5
23.7) trees $10 cm DBH/ha, whereas mean liana
density at the site was 2471 (SE 5 104.3) lianas
$2 cm DBH/ha. Mean basal area for trees $10
cm DBH was estimated as 19.2 (SE 5 0.82) m2/
ha, and for lianas $2 cm DBH 1.9 (SE 5 0.15)
m2/ha, yielding a liana/tree basal area ratio of 0.10.
We found 84 and 52 species of trees and lianas,
respectively (Fig. 1a; Appendices 1 and 2). The
abundance-based coverage estimator of tree species
richness was 105, while that for liana species rich-
ness was 54. Similarly, the incidence-based coverage
estimator of tree species richness was 107, and for
liana species richness was 57. Based on the species–
area curves, 1 ha of this forest had ca 65 tree and
51 liana species (Fig. 1a), yielding a liana/tree spe-
cies ratio of 0.78. The rate at which new species
were added to the sample as area increased was
higher for trees than for lianas (no. of tree species
5 (area)0.505, R2 5 0.982, df 5 1, 10; F 5 557.7,
P , 0.001; no. of liana species 5 (area)0.371, R2 5
0.962, df 5 1, 10; F 5 252.8, P , 0.001; differ-
ence in slopes, df 5 1, 20; F 5 33.8, P , 0.001;
Fig. 1b). Similarly, the slope of the log transformed
species-individual curve was higher for trees than
for lianas (no. of tree species 5 (area)0.507, R2 5
0.984, df 5 1, 10; F 5 607.8, P , 0.001; no. of
liana species 5 (area)0.332, R2 5 0.964, df 5 1,
10; F 5 268.2, P , 0.001; difference in slopes: df
5 1, 20; F 5 38.7, P , 0.001, Fig. 1c). The
Shannon-Wiener diversity index for trees was esti-
mated as 3.53 with an evenness of 0.80, and for
lianas was 3.40 with an evenness of 0.86.
ASSOCIATION BETWEEN TREE AND LIANA SPECIES.—Of
trees $10 cm DBH, 86.3 percent (SE 5 2.45)
supported at least one liana. Of the 21 tree species
with at least 10 individuals in the 12 plots where
we studied tree-liana interactions, 4 species of trees
had a higher proportion of liana-free individuals
than the mean proportion of liana-infested trees for
the 12 plots based on a binomial distribution, but
none had a higher proportion of liana-infested in-
dividuals (Table 1). Two of these tree species with
fewer individuals carrying lianas than expected
were the palms Astrocaryum aculeatum and Euterpe
precatoria. The other two species were Xylopia ser-
icea (Annonaceae) and Astronium fraxinifolium
(Anacardiaceae).
The matrices of similarities for tree and liana
species showed no correlation (Mantel r 5 0.22, P
5 0.20). Similarly, the matrices for tree species and
mean number of lianas per tree per plot did not
have a significant relationship (Mantel r 5 20.07,
P 5 0.64).
DISTRIBUTION PATTERN OF LIANAS ON TREES.—The
modal number of lianas per tree was 2, but the
range was large (0–34). The observed distribution
of number of lianas per tree was different from the
expected Poisson distribution (P , 0.001), with a
variance-to-mean ratio of 4.8, which indicated that
lianas had a clumped distribution on trees (Fig. 2).
This clumped distribution pattern persisted even
when the four species having a lower proportion of
individuals carrying lianas than expected were ex-
cluded from the analysis (A. acuelatum, E. preca-
toria, X. sericea, and A. fraxinifolium).
EFFECT OF TREE COMMUNITY CHARACTERISTICS AND
CANOPY STRUCTURE ON LIANA DENSITY .—Liana den-
 Lianas and Trees in Amazonian Bolivia 39

TABLE 1. Percent of liana-infested individuals $10 cm

DBH for tree species with ten or more indi-
viduals in 12 plots of 900 m2 in Oquiriquia
timber concession, Bolivia. The first four species
in the list had significantly lower proportions
of liana-infested individuals than the forest
mean of 86.3 percent based on the binomial
distribution.

% of trees
Species N with lianas
Astrocaryum aculeatum 13 7.69***
Euterpe percatoria 11 18.18***
Xylopia sericea 16 56.25***
Astronium fraxinifolium 22 72.73**
Senna sp. 1 14 71.43
Miconia sp. 1 13 84.62
Tabebuia heptaphylla 17 88.24
Physocalymma scaberrimum 47 89.36 FIGURE 2. Mean percentage of trees (columns; error
Casearia sylvestris 40 90.00 bars 5 1 SE) per liana-load category in twelve 900-m2
Spondias sp. 10 90.00 plots in Oquiriquia timber concession, Bolivia, compared
Ampelocera ruizii 21 90.48 to the expected Poisson distribution (continuous line).
Licania canescens 11 90.91
Vochysia haenkeana 11 90.91
Aspidosperma rigidum 16 93.75
Ocotea guianensis 16 93.75 lyzed in the context of five plot characteristics, the
Senna multijuga 50 94.00 number of lianas per tree increased significantly
Inga sp. 18 94.45 with the proportion of trees 10–30 cm DBH in a
Sapium marmieri 47 95.75 plot. None of the other four neighborhood plot
Aspidosperma cylindrocarpon 15 100
Luehea candicans 11 100 variables affected the number of lianas a tree carried
Roupala sp. 10 100 (Table 3).
** P , 0.01.
*** P , 0.001. DISCUSSION
DESCRIPTION OF TREE AND LIANA COMMUNITIES.—
sity increased in plots with a higher proportion of Tree density in Oquiriquia was similar to other
palms, and was not significantly correlated with Neotropical sites, but liana density was much high-
any other of the remaining four community and er (Fig. 3). High liana density in Oquiriquia was
structure variables analyzed (Table 2). also evident in the liana/tree basal area ratio of
0.10, higher than the ratio estimated for other for-
EFFECT OF CLIMBING MECHANISM ON THE NUMBER OF ests, which ranged from 0.02 to 0.04 (Gentry
TREES CLIMBED PER LIANA .—The modal number of 1983, Putz 1983). Because sites with large canopy
trees climbed per liana in this forest was two, with gaps were excluded, these estimates may be low for
a range of one to six. Analysis of covariance re- liana density and high for tree density in this forest.
vealed that the number of trees climbed per liana Nevertheless, the combination of high liana density
did not differ among the four mechanisms studied, and typical tree density suggests that liana density
but large diameter lianas climbed more trees in Oquiriquia neither depends on nor reduces tree
(climbing mechanism: df 5 3, 301; F 5 0.3, P 5 density. It is difficult to evaluate if in other liana
0.80; liana DBH: df 5 1, 301; F 5 35.3, P , forests, elevated liana density reduces tree density,
0.001). because quantitative accounts of other liana forests
are not available in the literature.
EFFECT OF TREE DIAMETER VERSUS NEIGHBORHOOD Basal area of trees $10 cm DBH in Oquiriquia
CHARACTERISTICS ON LIANA LOAD PER TREE .—Analysis (19.2 m2/ha) was low compared to other seasonally
of covariance revealed that both tree diameter and dry or wet tropical forests (range 5 17–75 m2/ha;
plot were significant factors in determining the Murphy & Lugo 1995, Kellman & Tackaberry
number of lianas per tree (plot: df 5 11, 538; F 1997). It is possible that high liana density reduces
5 3.5, P , 0.001; log tree DBH: df 5 1, 538; F tree biomass (Pérez-Salicrup 1999); however, data
5 126.7, P , 0.001). When tree DBH was ana- from other liana forests would be necessary to cor-
40 Pérez-Salicrup, Sork, and Putz

TABLE 2. Summary statistics from stepwise regression for effect of plot characteristics on liana density in Oquiriquia timber
concession, Bolivia (N 5 24, R2 5 0.17). Proportion of palms, trees 10–30 cm DBH, and trees $ 30 cm
DBH were estimated from the total number of trees $ 5 cm DBH in each 900-m2 plot.

Plot characteristic Coefficient F P

Proportion of palms 247.17 4.4 0.047
Proportion of trees 10–30 cm DBH 20.13 0.3 0.568
Proportion of canopy #5 m 0.24 1.3 0.260
Proportion of open canopy 0.08 0.1 0.714
Proportion of trees $30 cm DBH 20.14 0.4 0.514

roborate this hypothesis. Forest fragments near Ma-
naus, Brazil, have experienced reduced tree biomass
and higher liana densities as result of fragmentation
(Laurence et al. 1997, 2001), but there is no direct
evidence that tree biomass was reduced by the in-
crease in liana density.
Liana forests apparently originate following
large-scale human or natural disturbances. Inten-
sive logging, fragmentation, and introduction of
nonnative invasive species are human disturbances
that produce liana-infested forests (Boring et al.
1981; Appanah & Putz 1984; Baars & Kelly 1996;
Laurence et al. 1997, 2001; Uhl et al. 1997; Vidal
et al. 1997). Based on charcoal found in the soil
profile, Balée and Campbell (1990) have suggested
that liana forest in the Amazon basin may have
resulted from past human activity. Large-scale nat-
ural disturbances associated with liana-infested for-
ests include hurricanes and fires (Webb 1958, Sum-
merbell 1991, Pinard et al. 1999). For Oquiriquia,
there are no records of human land use in the past
500 years. When forestry operations began in
1994, the site was already a liana forest. We found
charcoal in the soil profile of Oquiriquia, but char-
coal also has been found in the profile of non-liana
forests in the Amazon basin and other regions of
the Neotropics (Sanford et al. 1985, Kennedy &
Horn 1997). Similarly, there is no documentation
to indicate that this forest is prone to hurricanes
or fires. Thus we do not have enough information
to suggest the origin of this liana forest.
In comparison with other tropical sites, the
Oquiriquia site is not floristically diverse. Only 84
species of trees were recorded, with a Shannon-
Wiener diversity index of 3.5 that is much lower
than that reported for most tropical and subtropi-
cal sites, which tend to be #4.1 (Gentry 1988,
Faber-Langendoen & Gentry 1991, Wright et al.
1997). Low tree floristic diversity in Oquiriquia
could be related to its relatively low annual rainfall.
This forest apparently receives only 1200 to 1700
mm of rain per year, and its tree species richness
fell within the values expected for this precipitation
(Gentry 1988, Phillips et al. 1994, Clinebell et al.
1995).
The liana community in Oquiriquia was less
species-rich than the tree community. This inter-
pretation is supported by the steeper slope of the
tree species versus the slope of liana species accu-
mulation curves plotted against area or individuals
in logarithmic scale (Fig. 2b, c), and by the higher
number of tree species versus liana species obtained
with both coverage-based estimators of species rich-
ness. However, the ratio of liana/tree species (0.78)
in 1 ha of this forest was much higher than any
other reported in the literature (0.10–0.4; Gentry
1983, Putz 1984b, Putz & Chai 1987, Gentry
1988, Appanah et al. 1993, Lieberman et al. 1996,

TABLE 3. Summary statistics from contextual analysis for effect of individual tree diameter and surrounding site char-
acteristics on the number of lianas carried per tree in Oquiriquia forest concession, Bolivia (N 5 551, R2 5
0.3). Proportion of palms, trees 10–30 cm DBH, and trees $ 30 cm DBH were estimated from the total
number of trees $5 cm DBH in each 900-m2 plot.

Effect Partial R2 Tolerance P

Tree DBH 0.474 0.98 0.001
Proportion of palms 0.036 0.53 0.485
Proportion of trees 10–30 cm DBH 0.133 0.58 0.007
Proportion of trees $30 cm DBH 20.071 0.57 0.153
Proportion of canopy #5 m 20.047 0.42 0.418
Proportion of open canopy 20.071 0.57 0.153

FIGURE 3. Tree and liana densities in four Neotrop-
ical sites. Tree densities were obtained from Lieberman et
al. (1986), Lang and Knight (1983), and Rankin et al.
(1990) for La Selva, Costa Rica, Barro Colorado Island,
Panama, and Manaus, Brazil, respectively. Liana density
for Barro Colorado Island was obtained from Putz
(1984b), and data for La Selva and Manaus are courtesy
of the Missouri Botanical Garden Alwyn Gentry transect
data set. Densities for Oquiriquia are from this study.
Wright et al. 1997). Thus the Oquiriquia forest
was not only very dense in lianas, but also relatively
rich in liana species.
ASSOCIATION BETWEEN TREE AND LIANA SPECIES.—As
expected by the high density of lianas, liana infes-
tation in Oquiriquia was also high; 86.3 percent
(SE 5 2.45) of trees $10 cm DBH carried at least
one liana. In comparison, 42, 49, and 57 percent
of trees carried lianas in Rı́o Negro (Venezuela),
Barro Colorado Island (Panama), and Sarawak
(Malaysia), respectively (Putz 1983, 1984b, Camp-
bell & Newbery 1993).
Of the 21 tree species with ten or more indi-
viduals, only 4 had a higher proportion of liana-
free individuals, suggesting that these 4 species may
have characteristics that allow them to shed or
avoid lianas. Two of these species were the palms
A. acueleatum and E. precatoria. Palms free them-
selves of lianas as new leaves grow and old leaves
Lianas and Trees in Amazonian Bolivia 41
are shed (Putz 1984a). The two other species, X.
sericea and A. fraxinifolium, were soft-wooded, fast-
growing species that may grow rapidly and shed
liana-infested branches; however, other species with
high liana loads were also soft-wooded and fast
growing. Thus, the reason why X. sericea and A.
fraxinifolium trees had fewer lianas than expected
is unknown. None of the tree species analyzed had
a significantly higher proportion of liana-infested
individuals than the forest mean (Table 1). This
result could be explained, at least in part, by the
very high frequency of liana infestation in the for-
est.
There was no significant relationship between
the plots with similar tree species and plots with
similar liana species composition. Similarly, the fact
that no relationship was found between plots with
similar tree species composition and the mean
number of lianas per tree supports the idea that no
tree species is particularly prone to carry many li-
anas. Thus, in this study site, the distribution and
abundance of liana species, and the number of li-
anas on a given tree, did not seem to be associated
with individual tree species.
DISTRIBUTION PATTERN OF LIANAS ON TREES.—Lianas
in the Oquiriquia forest had a clumped distribu-
tion on trees. The aggregated pattern of lianas on
trees suggests that once a tree is infested by one
liana, it has a higher probability than expected to
be climbed by a second liana. This pattern has been
noted in other tropical forests (Putz 1983, 1984b,
Clark & Clark 1990) and is potentially the result
of a facilitation process by which new invading li-
anas use the stem of the first established liana to
reach the crown of the tree (Pinard & Putz 1994).
If lianas use other lianas to reach the top of the
canopy, the possibility of finding specific associa-
tions between tree and liana species would be fur-
ther reduced; because at any one time, a tree will
be carrying lianas that have climbed on that tree,
plus lianas that climbed on the first established li-
anas.
EFFECT OF TREE COMMUNITY CHARACTERISTICS AND
CANOPY STRUCTURE ON LIANA DENSITY .—Contrary to
our expectations, plots with a higher proportion of
palms also had more lianas in this forest. Palms
tend to have no lianas (Putz 1984a); so apparently,
both palms and lianas tended to co-occur in this
forest in response to some other environmental fac-
tor, and not because palms facilitated the presence
of lianas. It is possible that both lianas and palms
become abundant at a certain stage of forest suc-
42 Pérez-Salicrup, Sork, and Putz
cession in Oquiriquia, but data to support this idea
are not available.
EFFECT OF CLIMBING MECHANISM ON THE NUMBER OF
TREES CLIMBED PER LIANA –Lianas utilizing the four
climbing mechanisms compared in this study did
not differ in the number of trees they climbed.
Thus, trees were not likely to carry more lianas as
an effect of growing in a neighborhood with a high
density of lianas having a particular climbing
mechanism.
Larger-diameter lianas climbed more trees, al-
though liana diameter only accounted for 10 per-
cent of the variation in this relationship. Thus oth-
er factors could affect the number of trees a liana
climbs. We suggest that two important possibilities
are the distance between tree crowns and the num-
ber of lianas already present in neighboring trees.
To colonize an additional tree, a liana must either
grow into a new crown from above, or fall from
the first tree, survive the fall, send a new shoot to
a second tree, and finally climb it (Peñaloza 1983,
1984, Putz 1984b). The number of lianas on
neighboring trees is likely to be important because
of the biomechanical facilitation process mentioned
above.
EFFECT OF TREE DIAMETER VERSUS NEIGHBORHOOD
CHARACTERISTICS ON LIANA LOAD PER TREE .—The
contextual model, which included individual tree
DBH and site structural features, revealed that larg-
er-diameter trees carried more lianas than slender
trees but that this relationship was affected by the
neighborhood conditions in which a particular tree
was growing. When tree diameter was analyzed in
the context of plot structural characteristics, trees
growing in plots with a higher proportion of trees
10–30 cm DBH also had more lianas. Larger-di-
ameter trees, which presumably are older, tended
to have more lianas in this forest, perhaps because
of longer exposure to liana infestation. This rela-
tionship was affected by the number of surround-
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ACKNOWLEDGMENTS
This research was possible through a grant from BOL-
FOR, a joint project of the Bolivian government and US-
AID, aimed at improving forest management operations.
Sigma-Xi and the Compton Foundation provided addi-
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 Lianas and Trees in Amazonian Bolivia 45

APPENDIX 1. Species of trees sampled in twenty-four 900-m2 plots in Oquiriquia forest concession, department of Santa
Cruz, Bolivia, and their quantitative parameters. N: total number of individuals; P: number of plots in
which that species was found; D: density; BA: basal area; RIV: relative importance value [(relative N +
relative BA)/2 3 100]. Species are ranked by descending RIVs.

Density (61 SE)

Species N P trees $10 cm DBH/ha BA m2/ha RIV
Physocalymma scaberrimum 130 22 60.2 6 11.70 2.8405 12.72
Sapium marmieri 130 16 60.2 6 19.99 2.6962 12.34
Senna multijuga 91 19 42.1 6 6.69 0.9161 6.11
Brosimum guianense 68 4 31.5 6 20.82 0.9713 5.31
Tabebuia heptaphylla 61 18 28.2 6 5.27 0.9799 5.05
Casearia sylvestris 69 18 31.9 6 6.83 0.6370 4.49
Inga sp. 1 47 22 21.8 6 3.98 0.4780 3.17
Ficus sp. 1 13 12 6.0 6 1.33 0.8991 2.87
Miconia sp. 1 32 15 14.8 6 3.76 0.5721 2.80
Aspidosperma cylindrocarpon 26 13 12.0 6 2.91 0.6461 2.75
Astronium fraxinifolium 44 15 20.4 6 4.99 0.3171 2.63
Senna sp. 1 33 15 15.3 6 3.71 0.3000 2.13
Ampelocera ruizii 28 10 13.0 6 5.08 0.3548 2.07
Aspidosperma rigidum 24 16 11.1 6 2.11 0.3576 1.91
Licania canescens 34 11 15.7 6 4.96 0.1911 1.89
Ocotea guianensis 23 6 10.6 6 5.11 0.3475 1.85
Astrocaryum aculeatum 23 12 10.6 6 2.63 0.1901 1.44
Dendropanax cuneatus 13 5 6.0 6 2.84 0.3475 1.44
Vochysia haenkeana 19 11 8.8 6 2.22 0.2332 1.39
Luehea candicans 19 6 8.8 6 3.78 0.2161 1.34
Xylopia sericea 21 9 9.7 6 4.09 0.1446 1.24
Dipteryx alata 9 5 4.2 6 1.87 0.3164 1.19
Cordia alliodora 13 5 6.0 6 2.91 0.2196 1.10
Pisonia aculeata 21 8 9.7 6 3.74 0.0573 1.01
Clarisia ilicifolia 18 10 8.3 6 2.70 0.1024 1.00
Didymopanax morototoni 5 5 2.3 6 0.94 0.2658 0.90
Trichilia anaequilatera 10 7 4.6 6 1.76 0.1718 0.86
Chorisia speciosa 8 6 3.7 6 1.44 0.1921 0.83
Hymenaea courbaril 5 5 2.3 6 0.94 0.2287 0.80
Ficus sp. 2 3 3 1.4 6 0.77 0.2549 0.79
Acacia loretensis 10 5 4.6 6 2.50 0.1404 0.78
Ocotea sp. 1 10 8 4.6 6 1.48 0.1397 0.77
Spondias sp. 1 13 5 6.0 6 2.67 0.0818 0.75
Acacia sp. 1 8 5 3.7 6 1.97 0.1460 0.71
Cariniana estrellensis 5 5 2.3 6 0.94 0.1815 0.68
Euterpe precatoria 11 6 5.1 6 2.50 0.0826 0.67
Alseis floribunda 11 8 5.1 6 1.77 0.0697 0.63
Vitex cymosa 2 2 0.9 6 0.64 0.2047 0.61
Senna sp. 2 5 4 2.3 6 1.15 0.1461 0.58
Ceiba samauma 7 7 3.2 6 1.05 0.1092 0.57
Spondias mombin 6 5 2.8 6 1.21 0.1002 0.51
Roupala sp. 1 1 1 0.5 6 0.46 0.1726 0.49
Amaioua sp. 1 6 4 2.8 6 1.53 0.0767 0.45
Pouteria hispida 5 4 2.3 6 1.15 0.0904 0.44
Swietenia macrophylla 6 4 2.8 6 1.38 0.0694 0.43
Pouteria sp. 1 7 4 3.2 6 1.70 0.0372 0.38
Maclura tinctoria 3 2 1.4 6 1.02 0.0912 0.36
Cecropia sp. 1 4 4 1.9 6 0.86 0.0645 0.33
Schizolobium amazonicum 2 2 0.9 6 0.64 0.0732 0.27
Lunania sp. 1 4 1 1.9 6 1.85 0.0241 0.23
Salacia sp. 1 4 4 1.9 6 0.86 0.0254 0.23
Sp. 11 1 1 0.5 6 0.46 0.0632 0.21
Phyllostylon rhamnoides 1 1 0.5 6 0.46 0.0599 0.20
Qualea sp. 1 4 3 1.9 6 1.09 0.0143 0.20
Cordia glabrata 1 1 0.5 6 0.46 0.0539 0.18
Amaioua intermedia 3 2 1.4 6 1.02 0.0316 0.16
Amburana cearensis 2 1 0.9 6 0.93 0.0471 0.16
Jacaratia spinosa 1 1 0.5 6 0.46 0.0136 0.16
46 Pérez-Salicrup, Sork, and Putz

APPENDIX 1. Continued.

Density (61 SE)

Species N P trees $10 cm DBH/ha BA m2/ha RIV
Dendropanax sp. 1 2 2 0.9 6 0.64 0.0269 0.15
Brosimum gaudichaudii 2 2 0.9 6 0.64 0.0232 0.14
Himatanthus attenuatus 2 2 0.9 6 0.64 0.0204 0.14
Sp. 13 1 1 0.5 6 0.46 0.0345 0.13
Sp. 2 1 1 0.5 6 0.46 0.0359 0.13
Dendropanax arboreus 2 1 0.9 6 0.93 0.0130 0.12
Sp. 12 2 2 0.9 6 0.64 0.0143 0.12
Sp. 3 2 2 0.9 6 0.64 0.0140 0.12
Sp. 6 2 2 0.9 6 0.64 0.0130 0.12
Trichilia elegans 2 1 0.9 6 0.93 0.0116 0.11
Calycophyllum spruceanum 1 1 0.5 6 0.46 0.0231 0.10
Rollinia sp. 1 2 2 0.9 6 0.64 0.0077 0.10
Sapium sp. 1 1 1 0.5 6 0.46 0.0209 0.10
Sp. 14 1 1 0.5 6 0.46 0.0222 0.10
Sp. 4 1 1 0.5 6 0.46 0.0052 0.10
Sp. 9 2 1 0.9 6 0.93 0.0233 0.10
Rhamnidium elaeocarpum 1 1 0.5 6 0.46 0.0207 0.09
Combretum sp. 1 1 1 0.5 6 0.46 0.0114 0.07
Randia sp. 1 1 1 0.5 6 0.46 0.0060 0.06
Sloanea sp. 1 1 1 0.5 6 0.46 0.0067 0.06
Sp. 7 1 1 0.5 6 0.46 0.0057 0.06
Sp. 1 1 1 0.5 6 0.46 0.0062 0.06
Garcinia madruno 1 1 0.5 6 0.46 0.0046 0.05
Guarea sp. 1 1 1 0.5 6 0.46 0.0039 0.05
Sp. 10 1 1 0.5 6 0.46 0.0038 0.05
Sp. 5 1 1 0.5 6 0.46 0.0040 0.05
Total 1219 19.2370 100.00
 Lianas and Trees in Amazonian Bolivia 47

APPENDIX 2. Species of lianas sampled in twelve 900-m2 plots in Oquiriquia forest concession, department of Santa
Cruz, Bolivia, and their quantitative parameters. N: total number of individuals; P: number of plots
where species was found; D: density; BA: basal area; RIV: relative importance value [(relative N 1
relative BA)/2 3 100)]. Species are ranked by descending RIVs.

D (61 SE)
Species N P lianas $2 cm DBH/ha BA m2/ha RIV
Mascagnia sp. 1 153 12 141.7 6 39.71 0.1964 8.55
Tynanthus schumannianus 189 12 175.0 6 22.77 0.1565 8.31
Tetrapteris sp. 1 134 10 124.1 6 42.08 0.1218 6.18
Machaerium sp. 1 105 11 97.2 6 20.29 0.1153 5.36
Tanaecium sp. 1 148 12 137.0 6 28.34 0.0721 5.20
Connarus sp. 1 99 9 91.7 6 26.70 0.1001 4.83
Bignoniaceae sp. 1 120 12 111.1 6 17.88 0.0775 4.71
Forsteronia sp. 1 103 12 95.4 6 15.68 0.0825 4.46
Serjania sp. 1 86 9 79.6 6 29.80 0.0845 4.13
Hippocratea sp. 2 109 12 100.9 6 20.67 0.0623 4.06
Doliocarpus dentatus 98 8 90.7 6 31.30 0.0536 3.59
Combretum sp. 1 58 8 53.7 6 17.34 0.0749 3.25
Mussatia hyacinthina 63 9 58.3 6 18.51 0.0618 3.02
Hiraea sp. 1 74 10 68.5 6 24.80 0.0508 2.98
Mascagnia sp. 2 53 9 49.1 6 22.70 0.0480 2.44
Malpighiaceae sp. 4 57 5 52.8 6 22.73 0.0410 2.35
Hiraea sp. 2 50 3 46.3 6 40.50 0.0439 2.27
Arrabidaea sp. 1 45 6 41.7 6 18.20 0.0401 2.05
Machaerium sp. 2 41 8 38.0 6 12.42 0.0387 1.93
Mascagnia platyrachis 33 9 30.6 6 9.19 0.0377 1.72
Paullinia sp. 1 36 3 33.3 6 18.40 0.0327 1.66
Bignoniaceae sp. 2 39 5 36.1 6 24.47 0.0292 1.64
Doliocarpus sp. 1 40 7 37.0 6 13.77 0.0229 1.49
Dalbergia sp. 1 27 2 25.0 6 17.94 0.0297 1.38
Uncaria guianensis 22 8 20.4 6 7.22 0.0250 1.15
Malpighiaceae sp. 3 43 2 39.8 6 38.82 0.0030 1.04
Strychnos sp. 1 17 4 15.7 6 8.59 0.0247 1.02
Forsteronia sp. 2 21 8 19.4 6 7.51 0.0135 0.82
Malpighiaceae sp. 2 3 2 2.8 6 1.99 0.0259 0.74
Combretum fruticosum 15 5 13.9 6 6.43 0.0151 0.73
Solanaceae sp. 1 15 5 13.9 6 8.33 0.0143 0.71
Tanaecium jaroba 13 5 12.0 6 5.55 0.0148 0.68
Celtis sp. 1 15 6 13.9 6 4.95 0.0123 0.66
Acacia sp. 1 16 2 14.8 6 12.10 0.0108 0.64
Leguminosae sp. 4 14 1 13.0 6 12.96 0.0115 0.61
Curarea sp. 1 10 1 9.3 6 9.26 0.0134 0.57
Dichapetalum sp. 1 11 6 10.2 6 4.21 0.0089 0.48
Leguminosae sp. 3 5 1 4.6 6 4.63 0.0119 0.42
Doliocarpus sp. 2 10 5 9.3 6 4.06 0.0069 0.40
Leguminosae sp. 1 9 5 8.3 6 3.38 0.0033 0.29
Bauhinia sp. 1 6 4 5.6 6 2.56 0.0036 0.23
Maripa sp. 1 3 1 2.8 6 2.78 0.0060 0.22
Combretum sp. 2 5 2 4.6 6 3.74 0.0036 0.20
Mascagnia sp. 3 3 1 2.8 6 2.78 0.0024 0.13
Malpighiaceae sp. 1 1 1 0.9 6 0.93 0.0036 0.12
Leguminosae sp. 2 3 3 2.8 6 1.45 0.0021 0.12
Serjania sp. 2 3 2 2.8 6 1.99 0.0021 0.12
Sp. 1 3 3 2.8 6 1.45 0.0021 0.12
Machaerium jacarandifolium 2 2 1.9 6 1.25 0.0009 0.07
Trichostigma sp. 1 2 2 1.9 6 1.25 0.0009 0.07
Hippocratea sp. 1 1 1 0.9 6 0.93 0.0007 0.04
Martinella obovata 1 1 0.9 6 0.93 0.0003 0.03
Total 2232 1.9177 100.00