Ecological Constraints On Rain Forest Management at Bajo Calima, Western Colombia

Forest Ecology and Management, 53 ( 1992 ) 213-244 213
Elsevier Science Publishers B.V., Amsterdam
Ecological constraints on rain forest management

at Bajo Calima, western Colombia
Don Faber-Langendoen
Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166, USA
(Accepted 23 August 1991 )
ABSTRACT
Faber-Langendoen, D., 1992. Ecological constraints on rain forest management at Bajo Calima, west-
ern Colombia. For. Ecol. Manage., 53: 213-244.
A forest harvesting system using skyline cables was evaluated for its effect on secondary forest struc-
ture and tree species richness of lowland rain forests in the Bajo Calima Concession, western Colom-
bia. Forests were sampled using six 0.1 ha plots in mature primary forest and sites 0.4, 4, 8 and 12
years since logging. Clear-cutting reduced overstory (trees greater than or equal to 10 cm dbh) basal
area, biomass and richness to 7%, 4%, and 17%, respectively, of primary forest levels. By the twelfth
year 46% of basal area, 37% of biomass, and 38% of richness had returned. However, 63% ofbiomass
and 50% of richness were composed of'core pioneer' species. The replacement of primary (climax)
species dominants by pioneer species indicates that the early process of regeneration is more appro-
priately described as secondary succession. Age since clear-cutting during the first 12 years of growth
was a significant linear predictor of both pioneer and climax species biomass and basal area, but not
climax richness. Extrapolation of these trends beyond 12 years suggests that overstory basal area and
biomass would equal that of mature rain forests by 30 years (the proposed rotation time). However,
the decline in pioneer growth rates over time, the slow climax growth rates, and the failure to predict
observations in an 18-year-old stand indicate that such a model is unrealistic. Climax species need
longer than 30 years to recover, and biomass may decline between 15 and 30 years if many short-lived
pioneer trees die before climax trees are well established. More consideration needs to be given to
understanding the regeneration of climax species if conservation and harvesting are to be combined.
Putting part of the concession under a longer rotation time may permit climax trees to regenerate
successfully.
INTRODUCTION
Managing species-rich tropical rain forests for forest products presents many
challenges. In particular, several ecologic constraints have complicated ef-
forts to establish a flexible, yet relatively simple, method of forest manage-
ment. These include the low abundance of desirable species, complex popu-
Correspondence to." Don Faber-Langendoen, The Nature Conservancy, Midwest Office, 1313
Fifth St. SE, Minneapolis, MN 55414, USA.
© 1992 Elsevier Science Publishers B.V. All rights reserved 0378-1127/92/$05.00

214 D. FABER-LANGENDOEN
lation dynamics, variable wood quality among species, slow growth rates, long
rotation times, and the need for detailed taxonomic and scientific skills to
harvest and treat the forests (Fox, 1976; Rankin, 1985). Because of these
constraints, the use of natural regeneration has had limited success in many
tropical countries (Mergen and Vincent, 1987). If management goals extend
beyond harvesting and production to include conservation of various biolog-
ical components of the forest, these will become additional constraints on
management (Wyatt-Smith, 1987). Including conservation goals would re-
quire preserving, among other things, some level of alpha diversity, the func-
tion of animal dispersal systems, and overall landscape diversity. These added
goals further complicate the management process, and few management sys-
tems have overcome the associated constraints.
Recently, the system developed by Smurfit Cart6n de Colombia to harvest
rain forests for pulp and paper has attracted attention because of its potential
to include both sustained production and conservation goals (US Dep. Agric.,
1980; Natl. Acad. Sci., 1982; Talbot, 1987; Gradwohl and Greenberg, 1988 ).
This system, developed in the Bajo Calima Concession of western Colombia,
is of interest for two reasons. First, the harvesting method permits rapid re-
generation of forests, and recovery of the original harvest is thought to occur
in as little as 30 years (Mazuera, 1979). Second, through the use of a pro-
posed natural regeneration method, many of the primary rain forest tree spe-
cies appear to regenerate in the secondary forest (Ladrach and Mazuera,
1985), suggesting that timber extraction and conservation are not
incompatible.
Given that conserving species via parks and wilderness areas is difficult in
many countries because of political and economic factors (Mares, 1986 ), the
beneficial features of including both harvesting and conservation in this for-
est development project are plaln enough. If the naturally regenerated forest
cannot be used for pulp and paper production because of ecologic, economic,
or other reasons, then the current operation is no more than a one-time har-
vest of the rain forest, with no subsequent economic or conservational bene-
fits. However, if the management plan works as anticipated, it may provide
one answer to development concerns for tropical countries.
An ecological study was undertaken to assess the use of natural forest man-
agement at Bajo Calima. The objectives of this study were to determine the
structure and diversity of natural regeneration after clear-cutting. Plots were
located in unlogged primary forest and in stands up to 18 years after clear-
cutting. The data provide information on the rates of stand development and
possible sources of the regeneration, and an evaluation is made whether the
patterns fit the management objectives for harvesting and conservation.
BACKGROUND
The Bajo Calima Concession is 10-20 km north of the city of Buenaventura

in the low hills region south of the Rio San Juan and Rio Calima (latitude
ECOLOGICAL CONSTRAINTS ON RAIN FOREST MANAGEMENT 215
3 ° 55'N, longitude 77°0'W). Forests in the concession, which is part of the

Choc6 biogeographic region, are classified by the Holdridge system as transi-
tional between tropical wet and tropical rain forest (Inst. Geogr. 'Augustin
Codazzi', 1977; Forero, 1982). Rainfall data, from 1961 onward, show that
annual rainfall exceeds 7000 mm, and is lowest from December to March
(200-300 m m per month); the lowest recorded monthly rainfall was 139 mm
in January 1963. Annual temperatures average 27.3 °C with average daily
minimum of 21.4 ° C and maximum of 30.3 ° C (Pulpapel, 1972 ).
The forests occur on low hills, with elevations of 50 to 150 m above sea
level, and slopes vary from moderate to steep ( 10 °-50 ° ). The soils, classified
as Entisols (FAO-UNESCO, 1971 ), are a mixture of mottled grey-yellow clays
and coarse Tertiary alluvial deposits, with soil pH of 4.3-5.0. They are nu-
trient poor, with very low concentrations of phosphorus and boron, and high
levels of aluminum (Cannon, 1985 ). Because of the extremely high rainfall
and hilly terrain, land use in the region is restricted primarily to forestry, al-
though agriculture is practiced along the river floodplains (Inst. Geogr. 'Au-
gustin Codazzi', 1983; Anonymous, 1984).
The vegetation is largely undescribed, but recent studies have found the
Bajo Calima forests to be among the most species-rich plant communities in
the world, with over 250 plant species per 0.1 ha for stems of 2.5 cm dbh or
greater (Gentry, 1986) and 252 tree species ha-~ for stems greater than 10
cm dbh (Faber-Langendoen and Gentry, 1991 ). The forests have high den-
sities of small and medium trees and low total above-ground biomass (ap-
prox. 200 tons h a - l ) , basal area (25 m 2 ha-~), and volume (190 m 3 ha -~ )
(Faber-Langendoen and Gentry, 1991). There are few free-climbing lianas
and many hemiepiphytic climbers. The most species-rich woody families are
Fabaceae, Rubiaceae, Sapotaceae, Arecaceae, Myristicaceae, Lauraceae and
Clusiaceae. The same families (except Rubiaceae and Lauraceae) and the
Vochysiaceae, Lecythidaceae and Bombacaceae are dominant in the tree layer
(stems of 10 cm dbh or greater). Tree palms, especially Jessenia bataua (mil-
peso), comprise 20% of tree stems (Faber-Langendoen and Gentry, 1991 ).
The Bajo Calima Concession
The management plan for the Bajo Calima Concession was based on over
15 years of experience in extracting wood for pulp and paper production. With
the support of the Colombian government, the 36 000 ha concession was
placed under a forest management plan in 1974, which required a sustained
production of wood and maximal yield from the forest (Pulpapel, 1972; Frisk,
1978; Ladrach, 1985a). Primary forest volume averaged 113 m 3 h a - ~for bole
volumes of stems greater than 13 cm dbh (but is as high as 146 m 3 h a - ~when
branch wood is included (Mazuera, 1985 ). The plan called for a rotation age
of 30 years, based on the findings of an annual diameter growth of 1 cm year-
2 16 D. FABER-LANGENDOEN
and a wood volume regrowth of approximately 3.8 m 3 ha-~ year-~. Given a

30 year rotation, cutting 1200 ha year- 1 would theoretically permit a sustain-
able harvest. At the end of the 30 year rotation (actually 26 years as 5280 ha
within the concession had already been harvested or colonized by settlers),
the natural forest will be transformed into a managed forest with units rang-
ing from 1 to 30 years of age. In addition, two permanent forest reserves of
1772 and 2372 ha have been set aside at the edges of the concession.
The harvest method
Harvesting is accomplished using a clear-cut method followed by natural

regeneration from seedlings and saplings, with no silvicultural treatments. The
concession was divided into two sections (A and B). Each section has 30
fronts, grouped into six units of five fronts. Each front is 600 ha, plus adjacent
forest edge. One front from each section is cut each year and the front extends
1000 m on either side of a 3 km stretch of road. Primary forest is left on the
outer edge of the front to provide a source of regeneration. The actual extent
of forest edge surrounding a front varies, depending on the location of other
fronts; for example, the 4-year-old front in the B-5 unit has less primary forest
edge than the 0.4-year-old front (Fig. 1 ).
To harvest the forests, teams of cutters manually fell almost all trees over
13 cm dbh with axes or chainsaws, clean off the bark and branches, and sec-
tion the boles into 1.5-m lengths. Larger branches are also cut, adding 20-
30% to the harvestable volume (Mazuera, 1985 ). Bundles of wood are placed
along lines in the clear-cut zone and lifted to the road using skyline cables
(see Frisk, 1978 and Ladrach, 1985a for details). Truckers haul the hand-
loaded timber 140 km to pulp processing mills in the city of Cali.
Not all trees are cut. For example, palms (which do not produce good pulp
fiber), Brosimum guianensis (which has an irritating latex in the trunk), badly
deformed trees or trees with boles or stems less than 13 cm dbh are usually
left behind.
Four major features of the plan warrant special attention: ( 1 ) natural re-
generation after clear-cutting leads to rapid regrowth of the forest; (2) al-
though clear-cutting damages much of the understory, the skyline cable sys-
tem prevents damage to the remaining seedlings and saplings, as well as the
soil base. All bark and branches left behind are a nutrient source for regener-
ation; (3) the use of natural regeneration permits at least some native rain
forest species to regrow; (4) the forest management plan is based on a sus-
tained yield system that is designed to ensure production in the future. Thus,
in sum, this plan overcomes some of the ecological constraints on combining
harvesting and conservation objectives in one plan.
Social and political tensions may disrupt the plan. Despite the government
having granted the concession to Smurfit Cart6n de Colombia, local people
,5" ~"
:"- \ ' - '<.. q

~: . . . . . . \~8 y,.:L \ /
i ) ~ .... 4:,', ) -'~., ~ ~.
i i "~ III! pF" " ~ d e road ~ wl~', l ~ ' a ~ |

~". . . . . J I 1 ,,,~, ~.,~ , ~'ulpape
r'Ulpapel~,j
I ~, 0.4 yr II ~.- station
~ - J ( 3 3
" - KILOMETRES
Fig. 1. Map of a portion of the Bajo Calima Concession showing the location of sample plots
within cutting fronts. Five fronts are found within a larger unit (cuartel), such as B V (B 5).
There are 12 such units in the entire concession, divided in half by the main SE-NW road. The
northern units are labelled A, the southern B. Two fronts (labelled 1 or 2 on the map) are
selected for cutting each year usually from a unit in either the A or B half of the concession.
Fronts are labelled 1 if they were located in or near the BV unit (except for Primary Forest
Front 2 ). Fronts are labelled 2 if they are outside this area. Thus each age is labelled as 1 or 2.
Each front covers 600 ha plus some forest edge. Inset map shows the location (arrow) of the
concession in Colombia.
still exert severe pressures o n these forests. At the age o f 8 - 1 0 years, regener-
ating forests p r o v i d e excellent pole-sized trees for c o n s t r u c t i o n a n d m i n e - p r o p
materials. In fact, m o s t sites w i t h i n the c o n c e s s i o n o l d e r t h a n 1 0 - 1 2 years
h a v e a l r e a d y b e e n cut o v e r again, u n d e r m i n i n g a t t e m p t s to m a k e the opera-
t i o n sustainable. T h e C o l o m b i a n g o v e r n m e n t has not clarified h o w it will
m a i n t a i n the c o n c e s s i o n for forestry uses after the c o m p a n y has c o m p l e t e d its
initial cutting. T h e s e larger social a n d political aspects o f the r e g e n e r a t i o n are
not dealt with f u r t h e r here, b u t will c e r t a i n l y o v e r w h e l m any ecological con-
siderations. A n y claims for the usefulness o f these m a n a g e m e n t plans de-
p e n d s u l t i m a t e l y o n the interests o f the C o l o m b i a n g o v e r n m e n t a n d S m u r f i t
C a r t 6 n de C o l o m b i a in using this land for sustainable forestry.
Research at Bajo Calirna on natural regeneration
Early studies highlighted the r a p i d r e g e n e r a t i o n after cutting. T h e s e include

an i n v e n t o r y in 1964 o f 4-year-old forest ( P u l p a p e l , 1 9 7 2 ) , a s t u d y in 1973
of natural regeneration from 6 to 8 years of age (Benitez, 1974), and studies

in 1975 and 1979 of natural regeneration up to 15 years of age (Ladrach,
1976; Mazuera, 1979). The latter studies were especially important in dem-
onstrating that, by 15 years, total volume was half of the primary forest before
cutting. This information was used to support a 30 year rotation plan, assum-
ing that, if in 15 years total volume was half, by 30 years it would be full. The
regeneration was considered of adequate quality for pulp (Benitez, 1974). In
addition, the early pioneer dominants, such as Cecropia, Vismia, Miconia,
and Goupia were being replaced by the original primary forest (climax) spe-
cies. Such rapid regrowth in the early stages of secondary tropical forest re-
generation has been reported elsewhere in the tropics (Richards, 1952; Swaine
and Hall, 1983; Whitmore, 1984; de Graaf, 1986; Uhl, 1987), though it has
been called secondary succession, rather than regeneration, because the early
stages of regrowth are dominated by a set of species that do not dominate in
the primary forest.
Sources of the natural regeneration were investigated in an important fol-
low-up study by Ladrach and Mazuera ( 1985 ). They suggested that regener-
ation of all species appeared to depend heavily on seed dispersal mechanisms
from mature trees along the edge of the clear-cut. Thus, both climax and pi-
oneer seedlings that appeared after clear-cutting colonized the site.
Two major problems were encountered in carrying out previous research at
Bajo Calima. First, identification of tree species was difficult because their
taxonomy is poorly understood (Monsalve, 1985; Gentry, 1988). Common
names used by local foresters were used in all the studies, hampering descrip-
tions of the forest, and making assessment of regeneration patterns difficult.
Second, the source of regeneration, whether from advance regeneration on
the site or from colonization by dispersal from off the site, remained un-
known because survival of seedlings and saplings present before cutting was
not followed during the cutting and subsequent years of regeneration. Hence,
the importance of dispersal for regeneration of primary rain forest tree spe-
cies was impossible to assess. Dispersal was not addressed directly in this study,
but the role of advance regeneration in the regrowth of primary forest trees is
described.
METHODS
Site selectionfor chronosequence
Sites were chosen by age since clear-cutting. As two cutting fronts of 600 ha
have been cleared annually beginning in 1974, two series of 0.4 (immediately
after clear-cutting), 4, 8, and 12-year-old sites were selected as a chronose-
quence. Chronosequence data must be interpreted carefully, as both the spa-
tial and temporal differences among the sites can affect the sequence of events
(Austin, 1981 ).
Where site and disturbance differences can be minimized, such data can
provide a useful description of regeneration and succession. Three 0.1 ha plots
were located within each front by picking three points at 0.5- or l-km inter-
vals along the road, starting from the edge of the adjacent cutting front (as
determined from company records) (Fig. 1 ). The forest was entered along
the ridge nearest to the point and areas were chosen that had no signs of fur-
ther disturbance after clear-cutting. Plots of 10 × 100 m (0.1 ha) were located
parallel to or angled along a representative location of the upper slope of the
ridge. Plots were located on upper slopes to minimize differences in altitude,
soils and moisture (Lieberman et al., 1985).
In each 0.1 ha plot, dbh was measured for all woody individuals of 10 cm
dbh or greater (overstory). Diameters of individuals between 2.5 cm and 9.9
cm dbh (understory or treelets) were measured in 5 quadrats of 10× 10 m 2
systematically located every other 10 m within the 0.1 ha plot (total under-
story sample equaled 0.05 ha). Height was visually estimated with the aid of
a 12 m clipper pole used to collect voucher specimens.
Comparative data for the primary forest areas were available for a 1.0 ha
and 0.5 ha macroplot (Faber-Langendoen and Gentry, 1991 ). Three 0. l-ha
plots were chosen in the upland portions of each macroplot to provide com-
parable sample sizes with the secondary forest plots. Data from these 0.1-ha
plots are adequate for our analyses, as emphasis is placed on the means of the
three plots for a given front (see below). In all there were 30 0.1-ha plots of
ages 0.4-12 years and primary forest.
One 18-year-old site was available for study. This 1 ha site behind the for-
estry station had been fenced and protected from pole cutting by the local
people (Fig. 1 ). However, it differed in two respects from other sites. First,
logs were removed after clear-cutting using a skidding system. Second, de-
spite protection, it has suffered some intervention by local people. As pioneer
tree species are especially desirable as poles, the second factor has had an
impact on their abundance, but less on overall diversity (as judged by 0.1 ha
plot data gathered in completely unprotected 18-year-old forests). As this site
was the only 18-year-old site available, it is included in this study for compar-
ative purposes. Two 0.l-ha plots were located in the site, and their values
were averaged. Plot locations were selected to avoid severely damaged areas.
Life history analyses
Voucher specimens were sent to the Missouri Botanical Garden for han-
dling and identification. Because many specimens were sterile and some were
of undescribed species, all undetermined specimens were sorted into mor-
phospecies based on leaf characteristics. Ongoing floristic work at the conces-

sion should continue to improve our identifications.
Tree species were separated into one of three life-history categories - - palm,
pioneer, or climax species - - based on field observations, previous work by
Ladrach and Mazuera ( 1985 ), and the literature (Whittaker, 1975; Denslow,
1987; Uhl, 1987; Swaine and Whitmore, 1988). Thus, except for the pi-
oneers, all broad-leaved evergreen tree species were lumped together as cli-
max rain forest species. Pioneer was defined narrowly as a set of 18 'core
pioneer' species of the genera Cecropia (four species ), Isertia (one species ),
Miconia (six species), Ochroma (one species), Pourourna (two species),
Psychotria (one species ) and Vismia (three species).
Basal area, density (no. of stems), species richness, and the Shannon di-
versity index (Whittaker, 1975) were calculated for each plot. Allometric
equations for estimating total above-ground biomass were taken from the lit-
erature for the following species groups; stemless palms, light-wood pioneers
and medium-wood pioneers (see table 1 in Uhl and Jordan, 1984 ); stem palms
(see table V in Folster et al., 1976 ); climax (Uhl et al., 1988a) (see also table
1 in Faber-Langendoen and Gentry, 1991 ). For climax species, an average
wood density of 0.604 g c m - 3 was used, based on the average wood densities
from Ladrach ( 1985b ) of the 30 most abundant species in two primary forest
plots (Faber-Langendoen and Gentry, 1991 ). These equations, although not
species specific, take into account major differences in growth form and wood
density among species. Volume was calculated as follows: V equals
Height(m) ×basal area(cm 2) ×0.5 (Dawkins ( 1961 ) in Whitmore (1984) ).
This represents volume for above-ground wood and bark (primarily the big-
ger woody parts greater than 4 cm diameter), and is comparable with biomass
estimates in that much of the tree is included. Another measure of volume,
used at Bajo Calima by Mazuera (1979), gave comparable results for the nat-
ural regeneration (see Faber-Langendoen, 1990 ).
Statistical analyses
Data for the 30 plots were averaged by front, the front being the unit of
replication (Hurlbert, 1984). As two fronts are cut per year, only two repli-
cates per year are possible. Fronts of 0.4-12 years and primary forest in one
area of the concession, 'Juanchaco' (BV unit) and 'Juanchaco Norte' (BI - -
west side) are grouped as a set (front 1 ), as they are closer together than the
other fronts (AIII, BI - - east side) (Fig. 1 ).
Trends in basal area, biomass, volume, density and richness were analyzed
by year for both overstory and understory. Basal area is emphasized as it was
based on direct field measurements. Biomass measures, however, provide a
more consistent measure of tree growth because species with the same dimen-
sions can differ widely in wood density (Uhl and Jordan, 1984; Ladrach,
1985b).
Linear and quadratic regressions were used to test whether age after clear-
cutting was a good predictor of regeneration patterns. They were applied sep-
arately by life history groups and overstory-understory. A nested one-way
ANOVA was used with plots nested within cutting fronts to determine if dif-
ferences between fronts were significant. A one-way ANOVA was also used to
examine the effects of age on richness and diversity, this time using the pri-
mary forest data as a control. All analyses were done using Statistical Analysis
Systems ( 1985 ).
Ordination was used to summarize the major patterns of species changes
among the plots. This analysis places plots with similar species composition
close together and dissimilar plots far apart in ordination space. Ordination
was performed using DECORANA, a Fortran program for detrended corre-
spondence analysis (DCA) (Hill and Gauch, 1980). Despite the somewhat
arbitrary method of detrending, DCA provides an ecologically interpretable
set of results (Gauch, 1982; Peet et al., 1988). Separate ordination analyses
were performed on overstory ( 133 species) and understory ( 146 species) data
for all 32 plots (0-18 years). Species average basal area per plot was used,
and rare species were downweighted.
For the two ordinations, biplots of the first two axes were made. Points
representing the average score of the three plots within a front were plotted.
These points were connected sequentially by age to suggest a possible trajec-
tory over time. If through natural regeneration the average species composi-
tion of secondary forests begins to resemble composition of the primary for-
est, then the trajectory will point back to the initial primary forest scores.
Following Halpern ( 1988 ), values for resistance were obtained by measuring
the ordination distance from the initial primary forest scores to the secondary
forest scores farthest away in ordination space. A second measure, from pri-
mary forest scores to the oldest secondary forest scores, gives a value for the
resilience of the forests - that is, how well they return to the original primary
forest composition.
Nutrient analyses
Soil samples were taken from the top 10 cm in three to five of the quadrats
in the 0.1 ha plot, pooled, and mixed. They were partially air dried and sub-
sequently placed in a plant drier at 60°C for 48 h. Soil nutrient and textural
analyses were completed by the soils division of Centro International Agri-
cultura Tropical (C.I.A.T.) in Cali, Colombia. Organic C was measured using
the Walkey-Black method. Soil pH was determined in a 1 : 1 mixture of soil
and water. Soils to be analyzed for phosphorus and potassium were extracted
using the Bray II method. Phosphorus was determined colorimetrically, and
potassium was determined by atomic absorption. Soils analyzed for Mg, Ca,
and AI were extracted using KC1 ( 1 N ) . Mg and Ca were determined using
atomic absorption, and aluminum was determined by titration. Boron was
extracted using hot water and determined colorimetrically. Manganese was
extracted by the double acid method and determined using atomic absorption.
Soil changes in relation to age were analyzed from 0 (primary forest) to 12
years (n = 10). Primary forest stands were included as they provide the start-
ing point for subsequent soil changes after clear-cutting.
RESULTS
Changes in basal area after clear-cutting

Clear-cutting reduced basal area in the overstory (trees of 10 cm dbh or
greater) from 27 m 2 h a - i to less than 2.0 m 2 h a - 1 and in the understory from
5.3 m 2 ha -1 to 1.5 m 2 ha -1 (averages of two fronts from Table 1, Fig. 2a).
During regeneration overstory basal area reached a maximum at 12 years of
12.5 m 2 ha-1; the understory reached a maximum at 8 years of 9.2 m 2 ha-1.
Age was a significant predictor of overstory and total basal area using linear
TABLE1
Changes in above-ground basal area, biomass, volume and density during natural regeneration after
clear-cutting. Values are given for overstory (over) (stems of 10 cm dbh or greater) and understory
(under) (stems 2.5-9.9 cm dbh) by year since clear-cutting. Values represent the mean of three plots
within a front
Front age Front Basal area Biomass Volume Density

(years) (m2ha -I ) (tons ha -t ) (m3ha -1 ) (No. ha -I )
Over Under Over Under Over Under Over Under
0.4 1 1.5 1.3 7.8 3.1 10.8 4.2 90 530

2 2.0 1.7 6.6 3.4 9.2 4.8 130 1090
4 1 2.0 8.1 7.5 24.6 11.6 23.6 120 6530

2 1.2 6.4 7.9 19.3 9.6 21.7 80 5540
8 1 6.9 7.9 29.0 21.8 43.1 25.3 480 5050

2 10.4 10.2 61.3 33.3 92.8 42.4 690 5790
12 1 10.9 7.2 66.7 22.5 97.9 29.9 690 3660

2 14.2 9.8 93.1 32.0 146.0 40.0 800 5220
18 1 12.0 8.6 63.7 23.1 97.5 30.3 470 5230
PF 1 29.7 5.3 235.9 16.2 329.5 22.0 790 3960

2 24.4 5.3 187.7 12.4 259.9 17.6 770 3460
PF, primary forest.

35
0
Over
30 •
Under
J~ 25
< 20
ua
o I I L I /'/
0 4 8 12 16 20 24 28 PF
3O
"£: 25
C'a
£
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',~15
0 •
O3 10 0
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4 8 12 16 20 24 28 PF
AGE (years)
Fig. 2. Changes in basal area after clear-cutting by (a) structural category and (b) life-history
group. Data from the first 12 years are analyzed by regression (Tables 3, 5). Regression lines
are shown only if they were significant (P< 0.05 ). Data for the 18-year-old site and the primary
forests (placed at what would be equivalent to the 30 year rotation) are shown for comparison.
regression (Table 2 ). For the understory, a quadratic regression was a signif-

icant improvement over linear regression (Table 2). Total basal area did not
vary significantly between fronts within age (one-way nested ANOVA;
F=2.65, P>0.05).
Extrapolation of overstory trends beyond 12 years suggests that secondary
forest basal area would equal that of the primary forest in less than 30 years.
Mean annual basal area growth varied from 0.9 m 2 ha-t year-1 to 1.3 m 2
h a - I year- 1 in 8 and 12-year-old forests. However, the value for the 18-year-
old site was lower than expected (Fig. 2a).
Examination by life-history groups showed that climax species were re-
duced by clear-cutting from a total basal area of 28.3 m 2 ha-1 in the primary
forest to 2.3 m 2 ha -I. By 12 years they had reached 6.7 m 2 ha -1 (Fig. 2b,
Table 3); the single 18-year-old site showed further, and large, increases to
12.9 m 2 ha- '. Palm species, which were damaged during the removal of other
TABLE 2
Summary of regression analyses for structural characteristics of regeneration. Data are analyzed for
total, overstory, and understory from fronts of 0.4-12 years ( n = 8 ) . Yis measure of abundance, Xis
• age. The quadratic equation (RQ) is provided only if it was a significant ( P < 0 . 0 5 ) improvement
over the linear
Dependent variable Equation r2
Total Basal area Y= 2.95 + 1.60X 0.87***

Biomass Y= 2.51 + 8.61X 0.84***
Volume Y= - 1.65+ 12.83X 0.81"*
Density RQ: Y= 59.29 + 159.38X- 10.27 X 2 0.82**
Overstory Basal area Y= - 0 . 0 8 + 1.02X 0.82**

Biomass Y= - 5.49 + 6.64X 0.78**
Volume Y= - 9.68 + 10.21X 0.77**
Density Y= 0.68 + 6.20X 0.82**
Understory Basal area Y=0.86+ 1.95X-0.11 X 2 0.86**

Biomass Y= 1.40+6.11X-0.33 X 2 0.79**
Density NS
**P < 0.01; ***P< 0.001; NS, not significant.
trees, attained values by the twelfth year comparable with those in the pri-
mary forest (Table 3 ). Pioneer species had low values in the primary forest
( 1.1 m 2 ha- 1), and, after clear-cutting, increased rapidly to values of 11.3 m 2
ha- ' in 12-year-old forests. Their proportion of total basal area increased from
3% in the primary forest to more than 50% in the 8- and 12-year-old forests.
For all three life-history groups, age was a significant predictor of basal area
using linear regression (Table 4, Fig. 2b). Climax species had a lower rate of
increase in basal area than did pioneer species. If recovery of basal area to
levels in the primary forest depends ultimately on climax species, or if their
recovery to these levels is of conservational interest, then extrapolation of
linear regression trends suggest that at least 56 years would be required for
recovery. If just overstory climax species' basal area is considered, the length
of time required increases to 97 years (Table 4, Fig. 3a). However, the climax
species value in the 18-year-old site was higher than predicted from 0.4- to
12-year trends.
Changes in biomass, volume and density with age
For the overstory and understory, changes in biomass and volume were
similar to those observed for basal area above (Table 1 ), and regressions
showed similar trends (Table 2 ). However, rates of change are predicted to
be slower for these measures than for basal area. Overstory biomass growth
rates ranged from 3.6 to 7.8 tons ha -1 year -1 in 8- and 12-year-old forests.
TABLE 3
t-
O
Comparison of above-ground basal area, biomass, volume, and density characteristics for life-history groups by year since clear-cutting. Values represent e
combined overstory and understory measures (see Table 1 for further explanations of overstory and understory)
t-
Front age Front Basal area Biomass Volume Density 70.

(year) (m 2 h a - ' ) (tons h a - ' ) (m 3 ha -~ ) (No. h a - l )
>
Climax Palm Pioneer Climax Palm Pioneer Climax Palm Pioneer Climax Palm Pioneer
o
0.4 1 2.3 0.4 0.2 8.7 1.5 0.7 11.8 2.0 1.3 530 70 20 z
2 2.3 1.3 0.1 6.6 3.3 0.3 8.9 4.8 0.3 700 490 30
7~
4 1 3.0 0.9 6.3 8.3 1.3 22.5 11.1 2.9 21.7 1490 910 4440 O'~
2 2.9 1.3 3.5 9.0 5.4 12.8 12.0 7.1 12.2 1620 920 3080
8 1 4.9 2.2 7.7 16.2 7.1 27.5 21.7 9.4 37.2 1820 1420 2290
2 7.9 2.4 10.3 32.4 8.4 53.7 43.7 13.9 77.5 2850 3190 2230
O
12 1 6.6 2.3 9.1 27.4 9.8 52.1 37.0 16.5 75.5 2000 940 1380 K
m
2 6.8 3.5 13.6 28.9 15.1 81.0 39.1 22.1 124.7 2080 1650 2290 Z
18 1 12.9 1.3 6.4 56.1 2.3 28.3 76.5 4.0 47.4 4090 720 890
PF 1 30.6 3.2 1.2 230.9 15.9 5.3 319.9 21.4 10.3 3530 1030 190
2 25.9 2.9 0.9 186.6 9.2 4.4 257.5 13.9 4.0 2940 1210 80
PF, primary forest.

TABLE 4
Summary of regression analyses by life history for structural characteristics of regeneration. Data are
analyzed for fronts of 0.4-12 years (n = 8 ). Y is measure of abundance, X is age
Life history Dependent variable Equation r2
Palm Basal area Y= 0.71 + 0.18X 0.78"*

Biomass Y= 1.88 + 0.84X 0.83**
Volume Y= 2.12 + 1.35X 0.86**
Density NS
Pioneer Basal area

Total Y=0.44+0.97X 0.88*
Over Y = - 1.02 + 0.70X 0.85***
Under RQ: Y = - 0.23 + 1 . 3 1 X - 0.08X 2 0.72**
Biomass Y= - 3.47 + 5.71X 0.83***
Volume Y= - 9.70+ 8.72X 0.81"*
Density NS
Climax Basal area

Total Y= 1.96+0.43X 0.74**
Over Y= 0.67 + 0.22X 0.66**
Under Y= 1.26+0.21X 0.68**
Biomass Y= 5.02 + 2.00X 0.71"*
Volume Y= 5.98 + 2.78X 0.72"*
Density Y= 85.05 + 12.88X 0.57**
RQ: Y= 41.56 + 4 0 . 1 7 X - 2.19X 2 0.78"*
*P< 0.05; **P< 0.01; ***P< 0.001; NS, not significant.
Extrapolation of the biomass regression beyond 12 years for overstory values

suggests that it would take 30-35 years to reach primary forest values; once
again, the 18-year-old site had values lower than predicted from 0.4- to 12-
year trends.
For life-history groups, pioneer species represented more than 60% of bio-
mass values at 12 years, compared with 2% in the primary forest (Table 3 ).
Again, if recovery of primary forest biomass depends ultimately on climax
species, at least 90 years would be required for their recovery, based on trends
extrapolated from regression models during the first 12 years (Table 4). These
data, like those for basal area, indicate that climax species regenerate slower
than do pioneers in the early stages of regrowth.
Clear-cutting reduced the overstory density from 780 stems ha-1 to 110
stems ha- 1and reduced understory density from 3710 stems h a - l to 810 stems
ha-~ (Table 1 ). The overstory values based on 0.1-ha plots are comparable
with values found in recently clear-cut 1.0-ha macroplots (64-94 stems ha-~;
Faber-Langendoen, 1989). Over 60% of the overstory stems remaining were
13 cm dbh or less, the lower limit of clear-cutting. By the twelfth year of re-
ECOLOGICALCONSTRAINTSON RAIN FORESTMANAGEMENT 227
301
(3
Climax •
/ i
25
20
vE
<
<
O3
<
n~
/ ~ ~ ~//
4 8 12 16 20 24 28 PF
b
i Ove
8O
_i Under
O'3 7O A
(13
A
~: 60
212 "
0 5O
rt-
09 40
W
~ 3o
n
(/3 2o
|
I I [ [ I I I //-..~
4 8 12 16 20 24 28 PF
A G E (years)
Fig. 3. Changes in climax (a) basal area and (b) species richness after clear-cutting. (a) Basal
area values are shown along with their regression lines (see Table 5), as are species richness
values (see Table 7 ). See Fig. 2 for further details.
growth, overstory density had returned to primary forest levels, but now pi-
oneers and climax species were codominants (each group had about 35% of
the total number of stems, the rest being palms) (Table 3 ).
Changes in species richness
Overstory tree species richness was reduced to less than 10 species per 0.1
ha after clear-cutting. By the eighth and twelfth year of regrowth, richness had
significantly increased to more than 20 species (Table 5, Fig. 4a). Understory
species richness was reduced to 25 species per 0.1 ha, but had almost tripled
by the eighth to twelfth years of regrowth and was comparable with values in
the primary forest (Fig. 4a) Values for total richness (which is less than sim-
ple addition of understory and overstory values, as the same species could
occur in both stories) peaked at 8 years, then declined. Quadratic regression
was a significant improvement over linear regression (r 2 increased from 0.68
to 0.85, Table 6, Fig. 4a), indicating that a maximum level had been reached.
TABLE 5
Comparison of diversity (tree species richness) for overstory (greater than 10 cm dbh) and under-
story (2.5-9.9 cm dbh) tree plots. Richness for the overstory was measured in 0.1 ha plots. Richness
for the understory was measured in 0.05 ha subplots. Values represent the mean of three plots within
a front
Year Front Diversity Diversity
Over Under Total Climax Palm Pioneer
0.4 1 8 23 30 25 4 1
2 10 28 35 29 5 1
4 1 9 50 53 37 6 10
2 6 51 53 38 4 11
8 1 18 59 64 45 7 12
2 22 75 82 66 5 11
12 1 20 59 70 53 6 11
2 20 63 66 49 7 10
18 1 24 95 104 86 7 11
PF 1 54 83 119 104 9 6
2 50 51 78 67 9 3
PF, primary forest.
Non-linear regressions may provide a more suitable model for these data as
the 18-year-old site had richness values exceeding even those of the primary
forest (Table 5, Fig. 4a). Overstory richness was best predicted by linear
regression. Diversity measurements using the Shannon index provided a very
similar trend with respect to age since clear-cutting (Faber-Langendoen, 1989,
1990).
Palm species richness was not significantly affected by clear-cutting; by the
twelfth year of regeneration seven species per 0.1 ha were present, compared
with primary forest levels of nine species (Table 5, Fig. 4b). Pioneer species
richness, initially low, very quickly reached maximum values by the fourth
year, then remained constant. As pioneers were defined as a specific set of 18
species (see Methods), this decline in numbers of new pioneer species indi-
cates that most have established early in regeneration. Climax values ranged
from 25 species per 0.1 ha immediately after clear-cutting to between 45 and
66 species ha- ~ 8-12 years later. Richness in the 18-year-old forest was even
higher, with 86 climax species, identical to the average primary forest values
(Table 5, Fig. 4b). Most of this total climax richness comes from the under-
story stems (Fig. 3b). In fact, overstory changes in climax species richness
were not significant using regression analysis (Table 6). Similarly, when us-
ing one-way ANOVA (with age since clear-cutting considered a treatment ef-
© O
110
©
100 u~e_r
oi
UJ7 80 L3
I •
0 70 .--2"-7--'©.
-
rr" 60 .".
. -" ; " , 5• .... ".~"'-.
k'~ "
w 5O
W 40 ," /
O- )" /
O9 30 ~/
20~ J - m - - f -
10
i I I i 1 I I //x~.-
4 8 12 16 20 24 28 PF
110
~z ao
"t-
O 70
rr" 60
W 50
ILl 40
0..
CO 3O
20
10
0 l I I I /'
4 8 12 16 20 24 28 PF
AGE (years)
Fig. 4. Changes in species richness by structural category and life-history groups. (a) Overstory
richness includes stems 10 cm dbh or greater in 0.1 ha plots. Understory richness includes stems
2.5-9.9 cm dbh in 0.05 ha subplots. Total richness represents their combined values. (b) Life-
history group richness includes all stems in 0.05 ha subplots and in 0.1 ha plots. See Table 6 for
regression equations.
fect and the primary forest serving as a control), the effects of clear-cutting
on richness remained significant even at 12 years ( n = 10, F = 82.9, P<0.0001,
range equals 5.674). In addition, fronts of 0.4, 8, and 12 years did not differ
significantly in species richness, indicating that reductions in overstory spe-
cies richness after clear-cutting remained unchanged over the first 12 years.
Extrapolation of regression data for total overstory richness alone suggests
that about 40 years are required to regain original primary forest richness
(Table 6, Fig. 4a); however, overstory climax richness showed no significant
change over time (Fig. 3b).
Species abundance patterns
Changes in species abundance differed partially, depending on their life
histories. In the overstory, palm species were least affected by clear-cutting
TABLE 6
Summary of regression analyses for species richness for fronts 0.4-12 years (n = 8). Data are pre-
sented by size category and life-history. Yis measure of abundance, X i s age (see Table 2 for details)
Richness Equation r2
Total RL: Y= 36.92 + 3.23X 0.68**

RQ: Y = 27.83 + 8 . 9 3 X - 0.46X 2 0.85**
Overstory RL: Y= 6.92 + 1.18X 0.67**
Understory RQ: Y = 2 0 . 5 7 + 10.02X-0.55X 2 0.93**
Climax
Total RL: Y = 2 8 . 6 8 + 2 . 3 1 X 0.58*
Overstory RL: NS
Understory RL: Y= 24.49 + 2.44X 0.62*
Pioneer
Total RL: Y= 3.83 + 0.74X 0.49*
RQ: Y=O.26+ 2 . 9 9 X - O . 1 8 X 2 0.93**
Overstory RL: Y=0.50+0.55X 0.69**
Understory RQ: Y= 0.67 + 2 . 6 2 X - 0.16X 2 0.76**
*P< 0.05; **P< 0,01.
(Table 7 ). The dominant palm species, J. bataua, varied in abundance with

age, but was consistently present in all fronts and ages. Other palms were more
sporadic.
Pioneer species in the overstory responded similarly as a group; that is,
while nearly absent in the overstory of primary forest, they were very abun-
dant 8-12 years after clear-cutting (Table 7 ). The abundance of Cecropia sp.
no. 13 peaked at 8 years; others, including Vismia panamensis, Isertia alba,
Miconia centronoides, Miconia s.p. no. 39, and Pourouma sp. no. 19, were still
increasing at 12 years. Understory regeneration of pioneers was also consis-
tently high (see below). Densities for all pioneers decreased in the single 18-
year-old forest, although basal area continued to increase for Pourouma and
M. centronoides. Several species classified as climax showed pioneer-like re-
sponses, reaching their maximum values at 8-12 years, e.g. Apeiba membran-
acea, Inga sp. no. 86, and Simarouba amara.
A number of climax species that were dominant in the primary forest over-
story were not seen in the regenerating forests, notably Pouteria buenaventu-
rensis, Eschweilera panamensis, and Tovomita weddelliana (Table 7). Their
regeneration in the understory was either non-existent or very low. Other spe-
cies, such as Otoba lehmanii, Goupia glabra, and Qualea lineata did reappear
in the overstory, either immediately or after 4-8 years of regeneration. Gou-
pia and Qualea were especially abundant in the understory by the twelfth year.
Dominant understory species that only attain small stature when mature
were generally less affected by clear-cutting. Climax species, such as Mabea
ECOLOGICALCONSTRAINTSON RAIN FOREST MANAGEMENT 231
TABLE7
Regeneration patterns of dominant overstory (10 cm dbh or greater) species in the Bajo Calima
Concession. All species are shown that occurred in more than three of 32 plots and had an average
density of at least three stems per 0.1 ha in one front. Each mean value represents the average of two
fronts. A blank means the species was not found in that age group
Life Species Age (years)

history
0.4 4 8 12 18 PF
Palm Jessenia bataua 1 1 1 3 1 5

Wettinia cf. quinaria 1 1 2
Socratea exorrhiza 1 2
Pioneer Cecropia sp. no. 13 4 3 1
Visrnia panamensis 1 12 15 3
Isertia alba 9 10 1
Pourouma sp. no. 19 4 6 3
Miconia centronoides 4 10 7
Micona sp. no. 39 1 1 6 1 1
Climax Apeiba membranacea 2 2 2
Inga sp. no. 86 1 3
Simarouba amara 3
Goupia glabra 1 3 1
Qualea lineata 3 2
Otoba lehmannii 1 1 2
Eschweilera panamensis 1 2
Tovomita weddeUiana 2
Pouteria buenaventurensis 1
occidentalis, Mabea sp. no. 6, Guatteria sp. no. 19, Macrolobium archerii, and
the palms Orbignya cuatracasana and Amandra decasperma, dominated in
the understory prior to cutting. They were reduced somewhat by clear-cut-
ting, but had values at 8-12 years that were 50-100% of their primary forest
values. Other climax species, such as Guatteria sp. no. 44, Inga sp. no. 43, or
G. glabra, were not c o m m o n in the primary forest understory, but regenerated
well in secondary forests.
Many of the same pioneer species that dominated in the overstory regen-
eration also dominated in the understory. In addition, Psyehotria, Cecropia
no. 14, Miconia reducens, and Vismia sp. no. 4, all small-statured pioneer tree
species, were common.
Ordination analyses
The two ordinations each contained two maj or axes of variation (overstory
eigenvalues were: Axis 1 - - 0.677, Axis 2 m 0.415, Axis 3 - - 0.195; under-
story eigenvalues were: Axis 1 0.449, Axis 2 - - 0.243, Axis 3 - - 0.180). - -
The overstory ordination had a greater degree of species turnover after dis-
turbance than did the understory, as measured by the size of their eigenvalues
(Gauch, 1982 ). Both fronts of the overstory show similar trends in their tra-
jectories, with the greatest displacement from primary forest characteristics
occurring at 8 and 12 years (Fig. 5a). The single 18-year-old forest was more
similar to primary forest. Dominant species in 12-year-old overstory plots
were predominantly the pioneer species, such as Cecropia sp. no. 13, I. alba,
M. reducens, 111.panamensis, Ochroma lagopus, Cecropia sp. no. 14, M. cen-
tronoides, and Pourouma sp. no. 19, as well as a number of climax species,
Pouteria sp., S. amara, A. membranacea, Inga sp. no. 88, and Ficus trigonata,
as well as one palm species, Bactris sp. (see also Table 7). Thus, the first
200
,-, .o;T2
150 /,. '0.4 "\
/" N
./" N,
¢u /.. /' 'N",\
~I00
<
/" 'N.
F 4 ""~''" "~1 8
5O 4
I i I
0
0 50 100 150 200 250 300 350
AXIS 1
200 D
4 8
12
150 : ..' . !.
(xl
co 1oo
X "',.., '..,.
• ,~, ..
50 "" %. \,.
FRONT 1 FRONT2 "%"~
-- 4
,--@--. •
i J , i , J i
00 50 100 150 200 250 300
AXIS 1
Fig. 5. Ordination results from detrended correspondence analysis. Each point represents the
average of three plots within a front, except for the single 18 year site, which was sampled with
two plots (see Methods; site selection). All Front l or Front 2 ages are connected as if they
represented changes over time (chronosequence). Arrows from the 12-year-old fronts indicate
the expected direction of the trajectory based on the composition of the 18-year-old site. (a)
Overstory (stems l0 cm dbh or greater) fronts from primary forest to 18 years; (b) Understory
(stems 2.5-9.9 cm dbh) fronts of the same ages.
ECOLOGICALCONSTRAINTSON RAIN FORESTMANAGEMENT 233
TABLE8
Measures of resistance and resilience for rain forests at Bajo Calima. Resistance is measured using the
maximum Euclidean Distance (ED) between primary forest and the most distant age group on the
ordination in Fig. 5. Resilience is measured using the ED between primary forest and the oldest age
group ( 18 years) on the ordination in Fig. 5. Distance values given in the table are in standard devia-
tion (SD) units defined by Detrended Correspondence Analysis (see Methods: Statistical analyses).
Understory distances are weighted by the percentage difference in the eigenvalue of the first two axes
between overstory and understory (see Results: Ordination analyses)
Ordination Maximum ED to
ED oldest plot
Overstory Front 1 2.58 2.10

Front 2 2.56 2.02
Understory Front 1 1.62 0.61

Front 2 1.67 0.71
ordination axis represents a gradient from climax-dominated plots to pi-

oneer-dominated plots. The second axis separated plots within the primary
and recently cut years, indicating that plots are not very homogeneous in spe-
cies composition.
The understory was most dissimilar to primary forest at 4 years, and there
was little change for the next 8 years (Fig. 5b). Species responsible for the
separation of 4- to 12-year-old plots from the primary forests were similar to
those listed for the overstory above. The 18-year-old site, however, was placed
much closer to the primary forest (Fig. 5b). In effect, the trajectory of sec-
ondary forests begins to return to a composition resembling the primary for-
est. The initial divergence in composition between 4-year-old sites in differ-
ent fronts (second axis variation on the ordination) is owing to the dominance
of M. reducens and Carapa guianensis in Front 1 and Cecropia alvarezii and
Miconia sp. no. 39 in Front 2. Both sets of dominants are absent in the other
front. By the 12th year these differences disappeared. Thus, pioneer recruit-
ment into the clear-cuts may be initially variable.
The distance measures of resistance and resilience were lower for the un-
derstory than for the overstory, indicating that the understory had a higher
relative resilience and relative resistance (Table 8). Thus, the species com-
position of the secondary forest understory was less displaced and began to
resemble the primary forest more quickly than for the overstory. This latter
trend, however, was almost entirely dependent on inclusion of the single 18-
year-old forest.
Soil texture and nutrients
Soils were virtually all clay loam, with an occasional area of sandy clay loam.
In general, soil properties did not vary significantly with age. Variation be-
TABLE9
Comparison of soil nutrients in the primary forest and natural regeneration after clear-cutting. Values
are given for the top soil layer ( 0 - 1 0 cm) (x_+ 1 SD). Each mean represents the average for the three
plots of a front
Soil variable Front Age (years)
0.4 4 8 12 18~ PF r 2'b
Percent organic matter 1 7.2 7.4 9.7 9.0 7.7 13.7 0.08
2 12.4 12.0 12.9 10.0 13.2
pH 1 4.4 4.4 4.4 4.4 4.3 4.0
2 4.3 4.4 4.5 4.4 4.6
Total P ( p p m ) 1 4.9 5.5 4.4 4.7 2.5 4.1 0.08
2 2.6 2.9 2.1 4.5 2.1
Ca (mmol per 100g) 1 0.91 0.96 1.40 1.30 0.60 1.15 0.49 c
2 0.82 2.39 2.50 1.24 0.54
Mg (mmol per 100g) 1 0.49 0.46 0.46 0.44 0.29 0.60 0.10
2 0.60 0.58 0.91 0.33
K (mmol per 100g) 1 0.14 0.16 0.15 0.15 0.12 0.26 0.28
2 0.43 0.17 0.19 0.13 0.21
A1 (mmol per 100g) 1 3.4 3.2 4.0 3.4 4.1 4.6 0.50 c
2 4.6 2.8 2.9 4.2 5.7
B (ppm) 1 0.66 0.60 0.67 0.44 0.40 0.83 0.02
2 0.41 0.37 0.38 0.50 0.43
Mn ( p p m ) 1 12.9 12.0 23.0 14.3 6.9 26.6 0.20
2 8.0 32.5 41.1 30.7 16.3
aMean value from two plots on the same site.

br2 values for linear regression unless otherwise indicated (all were NS at P > 0.05 ).
Or2 for quadratic regression ( P < 0.1 ).
tween fronts was often greater than variation between ages. For example, soil
organic matter, which was 13.4% in the primary forest, ranged from 7.2 to
12.4% in the 12-year-old forest (Table 9 ). Total phosphorus was lowest in the
primary forest, and highest in the 12-year-old forest. Two nutrients, calcium
and aluminum, showed quadratic responses with ages (P< 0.1 ): calcium val-
ues increased from 0.78 in the primary forest to 1.93 mmol per 100 g in the
8-year-old forest, then declined, whereas aluminum values decreased from
5.2 to 3.0 mmol per 100 g in the 4-year-old forest, then increased to 3.8 mmol
per 100 g in the 12-year-old forest. Soil values from the single 18-year-old
forest supported these trends.
DISCUSSION
Changes in structure and life histories after clear-cutting
Secondary forests in the Bajo Calima Concession grew rapidly during the
first 12 years after clear-cutting. Changes in basal area, biomass, and volume
indicated that the regeneration attained values by the twelfth year that were
at least half of the primary forest. These results lend support to previous stud-
ies at Bajo Calima by Mazuera (1979) and Ladrach (1976) that suggested
that harvestable basal area for pulp and paper was half that of the primary
forest by 15 years. However, much of the regrowth was due to a small set of
pioneer species, and the early process of regeneration at Bajo Calima is more
frequently described as secondary succession, rather than natural regenera-
tion (Grubb, 1977; Swaine and Hall, 1983; Whitmore, 1984; Uhl, 1987). This
distinction does not imply that natural regeneration may not ultimately be
possible, but only that the observed changes in the shorter time period are
better described as species replacement than regeneration.
Of fundamental importance to the interpretation of stand development is
whether linear trends in basal area from 0 to 12 years can simply be extrapo-
lated over the next 15 years to support a 30 year rotation. The single 18-year-
old forest did not support such an extrapolation in all cases, but it had been
harvested differently and subject to selective cutting during regeneration, so
it may not be typical. One-year measurements of growth rates in tree diame-
ters in this stand, showing values less than the required 1 cm year-1 to pro-
duce values equal to primary forest by 30 years (Faber-Langendoen, 1989),
could also be questioned because of the differences in the stand's history. In
other studies, patterns of regeneration over periods of time greater than 20
years have shown non-linear trends. Uhl (1987) found linear trends during
the first 10 years of succession for many tropical sites, including his own at
San Carlos in Venezuela. By contrast, in an 80 year study of succession after
slash-and-burn agriculture at San Carlos, Saldarriaga et al. (1988) found that,
although biomass values at San Carlos were a third of the primary forest by
20 years, after 60 years of succession they were still below the primary forest
values. Biomass values for the first 12 years of succession at Bajo Calima are
similar to those at San Carlos (Fig. 6). Soils at San Carlos, like those at Bajo
Calima, are nutrient poor.
Beyond these comparisons, other factors suggest that linear trends cannot
simply be extrapolated. The data from Bajo Calima clearly indicate that pi-
oneer species are responsible for the rapid rates of growth (Figs. 2a,b). Pi-
oneer species establish quickly and begin flowering and reproducing at a very
young age, as early as 3-5 years (Longman, 1985; Martinez-Ramos and A1-
varez-Buylla, 1986). Pioneers occupy more than 50-60% of basal area and
biomass in the 12-year-old forest and may remain dominant in 30 year sec-
ondary forests, even if overall basal area is comparable with primary forest.
However, many pioneer species are short-lived (Richards, 1952; Whitmore,
1984), and declines in pioneer density and growth rates after 5-10 years of
succession have been documented elsewhere (Swaine and Hall, 1983; Lad-
rach, 1985c; Uhl, 1987). While the short life of pioneer species may eventu-
ally lead to decreased pioneer dominance, it is not clear that climax growth
236 D. F A B E R - L A N G E N D O E N
• BAJO CALIMA
D SAN CARLOS (ST)
• SAN CARLOS(LT)
" ~ 150 ii /
/
/
t-
O
~ IO0
.r ~
/
0
~ 5o
0 10 20 30 40 50 60
YEAR
Fig. 6. Changes in biomass during natural regeneration compared between San Carlos and Bajo
Calima. The San Carlos short term (ST) data are taken from Uhl ( 1987 ); the San Carlos long-
term (LT) data are taken from Saldarriaga et al. (1988).
will compensate for this loss, as estimates based on regression models for the
first 12 years of regrowth also suggest that climax species will be at only half
their original basal area by 30 years. Thus, life-history trends predict that basal
area should decline sometime after 15 years before presumably increasing
again.
The decline in pioneer species growth may also reflect declining soil nu-
trient availability. Uhl et al. ( 1982 ) suggested that declines in the growth of
Cecropia could be caused by declining levels of nutrients after the initial flush
from clear-cutting or cutting and burning (their study included both). At Bajo
Calima, the decomposing litter left after clear-cutting may prolong the rapid
growth of pioneer species (see Buschbacher et al., 1988). This input of nu-
trients may also explain why no significant differences in nutrient levels were
observed between plots of various ages (Table 9).
Further consideration needs to be given to climax species regeneration as
conservational goals require that they be able to mature in the long run.
Abundances of climax species increased slowly, if at all, during the first 12
years after clear-cutting. Whether their regeneration is slowed by competition
from the pioneer canopy is not known. Species such as E. panamensis will
regenerate to some extent, but at 12 years, few individuals were very large
(Faber-Langendoen, 1989 ). As it is unlikely that they will flower and repro-
duce within 30 years (Longman, 1985), the 30 year rotation is too short to
permit successful regeneration. Hartshorn (1989a) also predicted that slow
maturing climax species would be lost from the strip-cutting system being
used in the Palcazu Valley of Peru.
Changes in species richness
Levels of species richness were very high in secondary forests, primarily

because of the high number of understory saplings and treelets that survived
and/or regenerated after clear-cutting (Table 1 ). By contrast, overstory rich-
ness increased slowly (Fig. 4a) and, for overstory climax species, did not in-
crease significantly at all over a 12 year period (Fig. 3b). Climax richness
may increase later as seedlings and saplings recruit into the overstory; there-
fore the long-term prospects for recovery of overstory climax richness may be
better.
The rapid increase in total species richness after clear-cutting is not entirely
unusual in forest regeneration (Hartshorn, 1989a,b). Swindel et al. (1984)
argue that cutting itself increases diversity in the regenerating forest when-
ever clear-cutting reduces, but does not eliminate, previously abundant spe-
cies. Almost every species becomes scarcer. In addition, species richness, al-
though relatively easy to measure, simply indicates presence in a stand and
says nothing about the critical aspects of population persistence and repro-
duction. Long-term monitoring of tree plots are essential for these
determinations.
Resilience to clear-cutting
Resilience and resistance of the understory to clear-cutting was greater than

that of the overstory, as indicated by the ordination trajectories (Fig. 5 ). The
degree to which understory resilience will aid overall resilience will depend
on how rapidly understory seedlings and saplings can pass through the smaller
size classes into the overstory. A more dynamic approach than the one used
here would be necessary to permit projections of future stand composition.
The 18-year-old site provided one indication that the secondary forests may
eventually become more similar to the primary forest. As stated above, the
effects of disturbance by the local people on this site are unknown; if they
removed the more common pioneer species, they may have improved the
similarity in composition to primary forest, while simultaneously decreasing
the basal area. At any rate, more 18-year-old sites are needed to verify the
recovery patterns observed here.
In general, clear-cutting with minimal soil disturbance followed by natural
regeneration encourages rapid regeneration, especially in comparison with
clear-cut and burn, clear-cut and bulldoze, or clear-cut, burn and farm (Jor-
dan, 1987 ). The soil nutrient data from this study suggest that disturbance to
the soils by clear-cutting was minimal, and improved resilience. Good recov-
ery in the understory after clear-cutting has also been reported in other forest
systems (Halpern, 1988; Hutchinson (1982) in Longman and Jenik, 1987).
In fact, for Halpern's study of clear-cut forests that had no soil disturbance or
238 D.FABER-LANGENDOEN
burning, measures ofunderstory resistance ( 1.63 ) and resilience (0.87) were

very similar to those reported here ( 1.65 and 0.66, respectively, Table 8 ).
The patterns of rain forest resilience to clear-cutting, as measured by tree
species composition, suggest a slower rate of recovery than when measured
by structural characteristics of basal area or biomass. Ewel's (1984) com-
ments are instructive: "How does the resilience of wet, lowland tropical for-
ests compare with that of other forests? If... 'maturity' is expressed as physi-
cal structure, such as biomass, then tropical ecosystems are probably as
resilient as most others. If, however, the 'maturity' axis incorporates not just
any kind of high diversity, but diversity consisting of the same array of species
that occupied the site prior to disturbance, then tropical lowland forests may
be among the world's most fragile ecosystems."
Dispersal and the source of diversity
The rapid recovery of understory richness and diversity raises the question
as to its source. From shortly after clear-cutting to 4 years of age, climax rich-
ness in the understory (stems 2.5-9.9 cm dbh) increased from 26 to 36 spe-
cies (Fig. 3b). Given the size of the stems, these species undoubtedly came
from advance regeneration. Over the next 8 years, only 10-15 climax species
were added to the understory. As seedlings less than 2.5 cm dbh were not
sampled during this study, and stump sprouts from stems of 10 cm dbh or
greater can add approximately nine species per 0.1 ha (Faber-Langendoen,
1989 ), it would appear that climax tree richness can be accounted for almost
entirely from advance regeneration and not dispersal (see also Uhl et al.,
1988b; Hartshorn, 1989b). Similarly, eight climax species remained in the
overstory immediately after clear-cutting; by the twelfth year overstory levels
had only increased to 20 species. These additional species seem to have been
added by recruitment from the understory advance regeneration. Thus, ad-
vance regeneration is the predominant source of regeneration for many of the
climax species observed in the secondary forest.
The contribution of seed dispersal to climax species regeneration during
early phases of regeneration may be small, Large-seeded trees, such as E. pan-
amensis and Pouteria spp. require larger bird and mammal dispersers that are
less likely to venture into large openings. By contrast, many of the pioneer
species have small seeds that are dispersed primarily by birds and bats, which
would more likely traverse large gaps or clearings (Schupp et al., 1989). In
addition, pioneer seeds may persist in the soil seed bank for longer periods of
time, and contribute to rapid establishment immediately after clear-cutting.
Pioneer seedling density in soils from the primary forest numbered 200 m -2,
while climax species numbered less than 50 m -2 (Faber-Langendoen, 1989).
It may be that the role of animal dispersers is important to the long-term
regeneration of many climax species (Gomez-Pompa and Vasquez-Yanes,
1981; Whitmore, 1984; Foster et al., 1986), but the rotation period is rela-
tively short and will not permit such a role. It will be necessary for manage-
ment policy to address what level of diversity is desirable or feasible in these
forests, and to determine whether adequate remnants exist along the edges of
clear-cuts to permit dispersal.
The role of the two forest reserves in protecting plant diversity in the
concession is completely unknown. No floristic, structural or diversity studies
have been done to determine whether these areas are representative of the
concession. Studies of the remnant forests surrounding each clear-cut front
are necessary to evaluate their role in promoting recovery of site diversity.
Combining conservation and forestry
The prospects for extracting forest products for pulp and paper within a 30
year rotation period and conserving the complete diversity of primary rain
forest species seem slim. If a 30 year rotation plan is retained, conservation
of tree diversity will be minimal. That is not to say that even limited mainte-
nance of such diversity has no value. But clearly many tree species (not to
mention other plant or animal species) would not be retained within these
secondary forests under the current plan.
Of more immediate concern to forestry management is the sustainability
of the 30 year rotation plan for pulp and paper production. Questions persist
about the likelihood that harvestable basal area or volume will have returned
in 30 years to that of either the original rain forest or sufficient for reharvest-
ing. The death rate of pioneer trees between 20 and 30 years may well outpace
the growth rates of the climax trees that are in the understory (Uhl, 1987). If
this is so, a 30 year rotation period is too long, and it may be more economical
to harvest at 20 years. Evidence from trends in forest growth after 15 years
are woefully inadequate to decide on future growth potential.
Regardless of which basal area projections are correct, the 30 year rotation
period is too short to meet significant conservational goals, as the overstory
climax richness would have just begun to mature. There is, however, another
option. Once the entire concession has been logged it might be desirable in
the next cutting cycle to change harvesting procedures. The secondary forests
could be divided into two groups, one to be cut on a long-term rotation, the
other on a short-term rotation. The short-term rotation would utilize the fast
growth rates of the pioneer trees, and forests could be harvested when growth
rates began to decline (possibly in 20-30 years). The long-term, e.g. 60 year
rotation, would be allowed to continue growing, encouraging the greater re-
generation of climax species. The use of long rotations in forest management
has been developed for temperate forests by Harris (1984). This two-level
rotation strategy may permit some economic return on part of the concession
and enhance conservational objectives on the remaining areas.
CONCLUSIONS
(1) The species-rich rain forests of Bajo Calima show vigorous succes-
sional growth after clear-cutting under the present system of harvesting, de-
spite the fact that clear-cutting reduces above-ground basal area and biomass
by over 90%.
(2) Patterns of secondary forest growth after clear-cutting indicate that tree
life-history groups differed in their response to clear-cutting. Early regrowth
during a 12 year period is dominated by pioneers, with several climax species
as subdominants. Several species are little affected by the clear-cut, either be-
cause they are palms (which are not harvested for pulp) or are understory
species that do not grow very large.
(3) Total (combined overstory and understory) regrowth during the first
12 years is sufficiently rapid that extrapolations of total basal area, biomass,
or volume values to 30 years suggests that complete recovery of primary for-
est levels would occur. If, instead of total regrowth, the growth of life-history
groups are examined, then a different pattern emerges. Much of the rapid
regrowth is accounted for by a small set of pioneer species. For climax species,
regeneration is slow, based on any of the regression models used. Extrapola-
tion of data beyond the 12 year period suggests that 90 years would be re-
quired to recover biomass levels of the primary forest. The single 18-year-old
site had values lower than expected. If the regeneration is cut in 30 years,
climax species will not have had time to mature and reproduce, and will be
lost from the system.
(4) Understory and total species richness is very high after 12 years of re-
growth, comparable with that of the primary forest. Resilience of the under-
story is high because clear-cutting does not damage the soil and many seed-
lings and saplings remain. Overstory species richness increased more slowly,
and climax overstory species richness did not change at all over the 12 year
period. Recovery of overstory climax richness, like that of climax basal area
and biomass, is slow.
( 5 ) Evaluation of the relative contributions of advance regeneration versus
seed dispersal suggest that advance regeneration of climax species from seed-
lings and saplings plays a very significant role in maintaining climax species
richness in these plots and probably accounts for almost all of the climax rich-
ness observed in 0.4- to 12-year-old secondary forests.
ACKNOWLEDGEMENTS
I thank AI Gentry for establishing the initial contacts that led to this coop-
erative study between the Missouri Botanical Garden and Smurfit Cart6n de
Colombia. His guidance in the field was indispensable in getting me started.
I thank Smurfit Cart6n de Colombia for providing all logistic support and for
the efforts made by Clem Lambeth, Edgar Mejia, and Carlos Barrera to see
that my needs were met during my stay in Colombia. Julio Nifio and the crew
at the forestry station are to be thanked for their cooperation and companion-
ship throughout my field research. Miryam Monsalve, Eulises Renteria, and
Jose Amobia provided much needed field assistance. The staff at the Missouri
Botanical Garden kindly examined many specimens, and Alan Brant gra-
ciously processed many of the specimens. I thank Robert Peck, William Lad-
rach, J.B. Hall and two anonymous reviewers for their comments and insights
into some of the ideas presented here. The study was completed as partial
fulfillment of the requirements for the Ph.D. degree in Biology at St. Louis
University.
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Forest Ecology and Management, 53 ( 1992 ) 213-244 213

Elsevier Science Publishers B.V., Amsterdam

Ecological constraints on rain forest management

© 1992 Elsevier Science Publishers B.V. All rights reserved 0378-1127/92/$05.00

The Bajo Calima Concession is 10-20 km north of the city of Buenaventura

3 ° 55'N, longitude 77°0'W). Forests in the concession, which is part of the

The Bajo Calima Concession

and a wood volume regrowth of approximately 3.8 m 3 ha-~ year-~. Given a

The harvest method

Harvesting is accomplished using a clear-cut method followed by natural

:"- \ ' - '<.. q

i ) ~ .... 4:,', ) -'~., ~ ~.

i i "~ III! pF" " ~ d e road ~ wl~', l ~ ' a ~ |

Research at Bajo Calirna on natural regeneration

Early studies highlighted the r a p i d r e g e n e r a t i o n after cutting. T h e s e include

of natural regeneration from 6 to 8 years of age (Benitez, 1974), and studies

Site selectionfor chronosequence

Life history analyses

phospecies based on leaf characteristics. Ongoing floristic work at the conces-

Changes in basal area after clear-cutting

Front age Front Basal area Biomass Volume Density

Over Under Over Under Over Under Over Under

0.4 1 1.5 1.3 7.8 3.1 10.8 4.2 90 530

4 1 2.0 8.1 7.5 24.6 11.6 23.6 120 6530

8 1 6.9 7.9 29.0 21.8 43.1 25.3 480 5050

12 1 10.9 7.2 66.7 22.5 97.9 29.9 690 3660

18 1 12.0 8.6 63.7 23.1 97.5 30.3 470 5230

PF 1 29.7 5.3 235.9 16.2 329.5 22.0 790 3960

PF, primary forest.

regression (Table 2 ). For the understory, a quadratic regression was a signif-

Dependent variable Equation r2

Total Basal area Y= 2.95 + 1.60X 0.87***

Overstory Basal area Y= - 0 . 0 8 + 1.02X 0.82**

Understory Basal area Y=0.86+ 1.95X-0.11 X 2 0.86**

**P < 0.01; ***P< 0.001; NS, not significant.

Changes in biomass, volume and density with age

Front age Front Basal area Biomass Volume Density 70.

PF, primary forest.

Life history Dependent variable Equation r2

Palm Basal area Y= 0.71 + 0.18X 0.78"*

Pioneer Basal area

Climax Basal area

*P< 0.05; **P< 0.01; ***P< 0.001; NS, not significant.

Extrapolation of the biomass regression beyond 12 years for overstory values

Changes in species richness

Year Front Diversity Diversity

Over Under Total Climax Palm Pioneer

PF, primary forest.

Total RL: Y= 36.92 + 3.23X 0.68**

*P< 0.05; **P< 0,01.

(Table 7 ). The dominant palm species, J. bataua, varied in abundance with

Life Species Age (years)

Palm Jessenia bataua 1 1 1 3 1 5

story eigenvalues were: Axis 1 0.449, Axis 2 - - 0.243, Axis 3 - - 0.180). - -

Overstory Front 1 2.58 2.10

Understory Front 1 1.62 0.61

ordination axis represents a gradient from climax-dominated plots to pi-

Soil texture and nutrients

P < 0.01; *P< 0.001; NS, not significant.

P< 0.05; P< 0.01; P< 0.001; NS, not significant.