Effects of Forest Fragmentation on Mortality and

Damage of Selected Trees in Central Amazonia

LEANDRO V. FERREIRA AND WILLIAM F. LAURANCE
Biological Dynamics of Forest Fragments Project, National Institute for Research in the Amazon (INPA), C.P. 478,
Manaus, AM 69011-970, Brazil

Introduction
Tropical forests are being cleared at a rate of over
150,000 km2 per year (Whitmore 1997), causing extensive loss and fragmentation of existing wildlife habitats.
Fragmentation has myriad impacts on the dynamics of
tropical ecosystems (e.g., Laurance & Bierregaard 1997)
but its effects on plant communities have received only
limited attention (e.g., Williams-Linera 1990; Laurance
1991, 1997; Malcolm 1994; Turner et al. 1996).
We describe the frequency of mortality and damage in
trees of the family Myrtaceae in fragmented and continuous Amazonian rainforests. By assessing the relative importance of edge and area effects and fragment age, we
can better understand the mechanisms of ecological
change in recently fragmented forests.

Methods
Study Area
This study is part of the Biological Dynamics of Forest
Fragments Project (Fig. 1), a long-term experimental
study of Amazonian forest fragmentation (Lovejoy et al.
1986; Bierregaard et al. 1992). The study area is located
70 km north of Manaus in central Amazonia (28309S,
608W), at 100150 m elevation. Local soils are nutrientpoor. Rainfall ranges from 19002500 mm annually with
a pronounced dry season from June to October. The forest canopy is 3037 m tall, with emergents to 55 m. The
local flora is remarkably diverse in tree species (Rankinde Merona et al. 1992).
The study area is surrounded by large expanses of continuous forest. In the early 1980s, a series of 1-, 10-, and
100-ha fragments (Fig. 1) were isolated by distances of
701000 m from surrounding forest by clearing and often by burning the intervening vegetation to establish
Paper submitted May 17, 1996; revised manuscript accepted October
23, 1996.

cattle pastures. Reserves ranging from 11000 ha in area

were delineated in nearby continuous forest to serve as
experimental controls.
Study Design
From 1980 to 1986 floristic inventories of all trees ($10
cm diameter at breast height) were conducted in 66
square, 1-ha plots in the study area (Rankin-de Merona et
al. 1992). Trees in each plot were marked with numbered aluminum tags and mapped, with leaves, fruits,
and/or flowers collected for every individual. From February to May 1987, one of the authors (LVF) revisited 56
of the plots to assess mortality and damage level to trees
in the family Myrtaceae, for which taxonomic identifications were especially reliable. Every individual was relocated, and any changes in its condition since the initial
inventory were assessed.
A total of 632 Myrtaceae trees of 59 identified species
or morphospecies were examined, averaging 11.365.3
trees ( x 6SD) per plot. Dead or damaged trees were
grouped into three categories: (1) standing dead trees;
(2) physically damaged or fallen trees; and (3) all dead or
damaged trees. We distinguished the first two categories
because standing dead trees may result from altered microclimatic conditions near forest edges (Lovejoy et al.
1986), whereas fallen or damaged trees commonly result
from wind disturbance (Laurance 1991).
Of the 56 plots (Fig. 1), 30 were located in forest fragments and 26 in continuous forest (controls). The fragment plots were in four 1-ha fragments (4 plots); three
10-ha fragments (17 plots); and one 100-ha fragment (9
plots). The control plots were in five 1-ha (5 plots); one
10-ha (3 plots); one 100-ha (9 plots); and one 1000-ha (9
plots) reserves. Plots within fragments were stratified so
that edge and interior areas were both sampled.
We devised four landscape predictors for each plot:
(1) distance to the nearest forest edge; (2) distance to
the nearest east-facing edge (edge aspect 5 701108 );
(3) fragment area; and (4) fragment age (number of years
797
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Fragmentation and Tree Mortality

Ferreira & Laurance

Figure 1. Map of study

area in central Amazonia
showing locations of 1-, 10-,
and 100-ha forest fragments and continuous forest reserves (controls).
Fragments and controls
used in the study are
shaded, whereas stippled
areas indicate cattle pastures or young regrowth,
and unstippled areas are
primary rainforest.

since isolation). We contrasted the two edge-distance predictors because our study area receives prevailing easterly
winds (V. Kapos, personal communication) that could
cause increased windthrow along east-facing edges. Edgedistances were measured from the center of each plot.
Plots were pooled into discrete categories to facilitate
statistical analysis. There were four categories for the
edge-distance measures (1 5 #60 m; 2 5 61100 m; 3 5
101500 m; 4 5 .500 m) and for fragment area (1 5 1 ha;
2 5 10 ha; 3 5 100 ha; 4 5 control) and three categories
for fragment age (1 5 3 years; 2 5 4 years; 3 5 7 years).
The rationale for the edge-distance categories is that microclimatic changes can occur within 60 m of forest
edges in the study area (Kapos 1989), whereas wind-disturbance may be pronounced within 100200 m of
edges and detectable up to 500 m from edges, at least in
some Australasian forests (Laurance 1991).
Statistical Analysis
Tree mortality and damage were patchy in nature, yielding a strongly positively skewed (approximately negative binomial) data distribution. We ranked the dependent variables, which reduced both skewness and
heteroscedasticity among samples. Ranked data were
compared between treatments using one-way ANOVAs,
followed where appropriate by Tukeys tests to contrast
sample means.
The factors describing fragment area and edge-distance were intercorrelated (rs 5 0.660.87; Spearman
rank correlations). To assess effects of area independently of edge effects, all plots ,150 m from any edge
were excluded, and the remaining plots were used to
contrast damage levels in fragmented and continuous
forest. The two edge-distance measures also were inter-

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Volume 11, No. 3, June 1997

correlated (rs 5 0.68) but their efficacy as predictors is

compared here as alternative explanations for the observed variation in tree damage.
The interval between the initial tree inventory and our
resurvey of each plot varied from 17 years. Plots with
longer intervals should accrue more dead trees via natural mortality processes, independent of other factors. To
assess the importance of this effect, we used data from
27 non-edge plots to estimate the natural mean mortality rate (M ) of Myrtaceae trees in the study area, using a
logarithmic model (M 5 ln(S )/t, where S 5 proportion
of individuals surviving the interval and t 5 time in
years; Lieberman et al. 1985). The expected natural mortality was calculated for each plot (expected mortality 5
[12M]t), and expected and observed values were compared using least-squares regression analysis.
A mathematical core-area model (Laurance 1991;
Laurance & Yensen 1991) was used to predict the impacts of edge effects on fragments of varying sizes and
shapes. The model generates accurate (.99%) predictions of the size of unaffected core-area for any fragment,
using three parameters: a fragment shape-index (SI ), fragment area (TA), and empirical knowledge of the distance
(d ) to which edge effects penetrate into fragments.

Results
Nearly half (25/56) of the plots had no damage, whereas
the remainder had 657% dead or damaged trees. Annual
mortality rates were low (0.56 6 0.26%, x 6 SE) in forestinterior plots (.100 m from edge) but were nearly seven
times higher (3.85 6 0.72%) in edge plots. Annual rates
of tree damage were over eight times higher in edge
(2.96 6 1.06%) than interior (0.32 6 0.32%) plots.

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Fragmentation and Tree Mortality

799

Table 1. Summaries of one-way ANOVAs used to assess effects of landscape features on tree mortality and damage in fragmented and
continuous forests in central Amazonia.
All dead or
damaged trees
Factor tested *
Dist. nearest edge
Dist. eastern edge
Fragment area
Fragment age

Standing dead
trees only

Fallen or
damaged trees

14.39
6.38
0.00
2.71

,0.0001
0.0009
0.9920
0.0840

6.33
8.56
0.77
1.09

0.0010
0.0001
0.3900
0.3490

12.87
2.46
1.33
8.24

0.0003
0.0730
0.2600
0.0016

* Sample sizes varied between treatments and comparisons, but were reasonably large (n 5 730 plots) for all treatments except non-edge plots
in fragments (n 5 4).

Variation in census interval appeared to have little effect on mortality and damage estimates. When both
dead and damaged trees were considered, the length of
the sampling interval accounted for only 5.3% of the total variation in the data set (F 5 3.03, p 5 0.087; leastsquares regression analysis).
One-way ANOVAs revealed highly significant effects
of edge-distance on most mortality and damage parameters (Table 1). Edge effects appeared to have a major influence on tree mortality and damage (Fig. 2). Distance
to the nearest east-facing edge, however, was not a better predictor than simple edge-distance. For overall mortality and damage, edge-distance accounted for more of
the variation in the data set (45.3%) than did distance to
nearest eastern edge (26.9%; based on sums of squares
from the ANOVAs).
Tukeys tests suggested tree mortality and damage
were significantly elevated within 100 m of fragment
margins. When total mortality and damage were considered, samples from both 060 m and 61100 m from
edges had significantly ( p , 0.01) higher values than
those farther from edges. Results were identical when
standing dead and fallen or damaged trees were analyzed separately (Fig. 2).

Fragment area effects per se appeared weak. When

only sites .150 m from edges were compared between
fragmented and continuous forest, the analysis was nonsignificant (Table 1), accounting for only 3.9% of the
variation in overall damage and mortality (ANOVA sums
of squares). Although this test was conservative because
there were only four non-edge sites in fragments, total
damage and mortality was virtually identical in fragmented (2.5 6 4.3%) and continuous (2.8 6 5.8%) forest.
Fragment age also had limited effects on tree damage
and mortality (Table 1). The effects of age were significant only for fallen and damaged trees, and the trend
was actually opposite to that expected, with more fallen
and damaged trees in younger (3- or 4-year-olds) than
older (7-year-olds) fragments ( p , 0.05; Tukeys test).
Fragment age accounted for 16.7% of the variation in total
mortality and damage (ANOVA sums of squares).
To generate the core-area model, we used a value of
d 5 100 m because tree mortality and damage were significantly increased within 100 m of edges (Fig. 2). The
model (Fig. 3) suggests edge effects may have a substantial impact on Amazonian forest remnants, even upon
larger (5001000 ha) remnants. This is especially apparent when realistic fragment shapes (i.e., SI 5 24) are
used in the model, rather than a square fragment (SI 5
1.13), which is unusual in real landscapes. For example,
using realistic shapes (Fig. 3), a fragment of 1000 ha is
predicted to have 2242% of its total area influenced by
edge effects, whereas a fragment of 500 ha would have
3058% of its area influenced.

Discussion
Edge and Area Effects

Figure 2. Effects of edge-distance on tree damage and

mortality ( x 6 SE) in central Amazonia.

Our findings suggest edge effects can cause sharply elevated tree mortality and damage in recently fragmented
forests, at least among species of Myrtaceae in Amazonia. Rates of tree death and damage were seven to eight
times higher in edge than non-edge sites. When all dead
or damaged trees were considered, nearly half (45%) of

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Causes of Tree Mortality

Figure 3. A core-area model for rainforest fragments

in central Amazonia. The empirical estimate for the
penetration-distance of edge effects (d 5 100 m) was
based on observed patterns of tree mortality and damage. According to the model, fragments with realistic
shapes (SI 5 24) are expected to become increasingly
vulnerable to edge effects once fragment area falls below about 500 ha.

the total variation in the data set (using ranked data) was
explained by the distance of plots to forest edge (Fig. 2).
The distance of plots to the nearest easterly-facing
edge (which receives persistent tradewinds) was a less
effective predictor, explaining 27% of the total variation.
This may occur because windstorms that damage trees
can come from virtually any direction (V. Kapos, pers.
comm.) and may cause complex patterns of forest disturbance (Boose et al. 1994) that obviate any simple relationship between edge aspect and tree mortality. In addition, microclimatic changes in fragments, such as
reduced humidity and increased temperature variability
near edges, may not vary greatly between easterly edges
and those with other aspects.
Edge and area effects are rarely discriminated in studies of fragmented ecosystems (Temple 1986; Laurance &
Yensen 1991; Didham 1997), but in this study we demonstrated that area effects per se had little apparent effect on tree mortality and damage, accounting for only
4% of variation in the data set. Area effects that might influence tree persistence in fragments include population-level processes, such as losses of small populations
via random genetic or demographic events (Shafer 1981),
and community-level phenomena, such as declines in reproduction following losses of specialized pollinators or
seed-dispersers (Powell & Powell 1987; Aizen & Feinsinger
1994). In general, however, we suspect such changes require longer time-scales to be manifested than in the recent (3- to 7-year-olds) fragments examined in this study.
At least for this data-set, edge effects apparently swamped
area effects as the proximate cause of tree damage and
mortality.

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There are two likely causes of elevated tree mortality

and damage near fragment margins. The first is microclimatic changes. Working in newly created forest fragments in the same study area, Kapos (1989) demonstrated that hotter, drier conditions more typical of
outside areas penetrated at least 4060 m into fragment
interiors. Microclimatic changes are often lessened in
older edges which become sealed by pioneer and secondary vegetation (Williams-Linera 1990; Kapos et al.
1993), but they are probably an important cause of tree
mortality in new fragments (Lovejoy et al. 1986). The
large numbers of standing dead trees near edges (Fig. 2)
may have been killed by sudden shifts in temperature,
relative humidity, or soil moisture that exceeded their
physiological tolerances. The observation that leaf-fall increases dramatically near recent edges (Lovejoy et al.
1986; Sizer 1992) suggests affected trees experienced
severe water-stress.
A second likely cause of tree damage is wind turbulence. When forests are cleared and fragmented, the
edges of remnants are exposed to increased windspeed,
turbulence, and vorticity, which often lead to elevated
windthrow (Chen et al. 1992) and forest-structural damage (Laurance 1991, 1997). Winds striking an abrupt forest edge cause an increase in downwind turbulence, resulting in pronounced wind-eddies for at least 210
times the height of the forest edge (Savill 1983). In tropical Queensland, Australia, a region subjected to occasional cyclones, Laurance (1991, 1997) suggested that
wind forces caused detectable increases in forest damage up to 500 m from fragment edges and marked damage within 200 m of edges. Unlike microclimatic changes,
wind damage is unlikely to lessen over time as fragment
edges become less permeable because downwind turbulence usually increases as edge permeability is reduced
(Bull & Reynolds 1968; Savill 1983).
Conservation Implications
Our results suggest that in central Amazonia, edge effects in tree mortality and damage penetrate about
100 m into fragment interiors (Fig. 2). Our study was not
explicitly designed to measure the penetration-distance
(d ) of edge effects, however, which can be assessed
most precisely using replicated edge-interior transects
(Laurance & Yensen 1991). Thus our estimate of d 5
100 m should be regarded as approximate.
The core-area model (Fig. 3) suggests edge effects
should increase rapidly in intensity as fragment area falls
below about 500 ha. Fragments below this size-range
may still have important conservation values (Turner &
Corlett 1996), but their ecological characteristics are
likely to differ markedly from those in intact forest. Such
differences may include the decline of wind- and drought-

Ferreira & Laurance

sensitive tree species near forest edges and a general

shift toward disturbance-adapted vegetation in fragments (Laurance 1991, 1997). A sudden increase in tree
mortality could potentially cause ecological distortions
that drive initial species losses in fragments (Terborgh et
al. 1997), especially among animals that have mutualistic
relationships with vulnerable tree species.

Acknowledgments
C. Gascon, S. Lewis, W. Magnusson, J. Chambers, and
three anonymous referees commented on earlier drafts
of the manuscript. This study was supported by World
Wildlife Fund-U.S., National Institute for Research in the
Amazon, Ministerio de Ciencia e Tecnologia (MCT-Brazil), Smithsonian Institution, and the Mellon Foundation.
This is publication number 175 in the BDFFP technical
series.

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