Ecosystems (2000) 3: 217–228
DOI: 10.1007/s100210000021
Land-Use History and Forest
Regeneration in the Cayey
Mountains, Puerto Rico
John B. Pascarella,*1,3 T. Mitchell Aide,1 Mayra I. Serrano,1
and Jess K. Zimmerman2
1PO Box 23360, Department of Biology, University of Puerto Rico, San Juan, Puerto Rico 00931-3360; 2Institute for Tropical
Ecosystem Studies, University of Puerto Rico, San Juan, Puerto Rico 00936
ABSTRACT
Although deforestation continues to be a major
threat to tropical biodiversity, abandonment of agri-
cultural land in Puerto Rico provides an opportunity
to study long-term patterns of secondary forest
regeneration. Using aerial photographs from 1937,
1967, and 1995, we determined land-use history for
2443 ha in the Cayey Mountains. Pastures were the
dominant land cover in 1937 and ⬍20% of the area
was classified as forest. Between 1937 and 1995,
forest cover increased to 62% due to widespread
abandonment of agriculture. To examine the effect
of historic land use on current forest structure and
species composition, we sampled secondary forests
in 24 abandoned pastures, 9 abandoned coffee
plantations and 4 old-growth forest sites. Sites were
located on two soil types along an elevational
gradient (125–710 m) and included a chronose-
quence from 4 to over 80 years old. After 25–30
years, basal area and species richness in secondary
INTRODUCTION
Deforestation, short-term use in agriculture, and
subsequent abandonment due to declining yields
and poor management have resulted in extensive
areas of secondary tropical forest (Brown and Lugo
1990). Unless land-use practices have severely de-
graded soils (Uhl 1987, Aide and Cavalier 1994),
these forests recover structural characteristics simi-
Received 16 June 1999; Accepted 8 October 1999.
3Author for correspondence and current address: Department of Biology,
Valdosta State University, Valdosta, Georgia 31698, USA.
*Corresponding author; e-mail: jbpascar@valdosta.edu
217
ECOSYSTEMS
r 2000 Springer-Verlag
forest sites derived from abandoned pastures and
coffee plantations were similar to old-growth forest
sites. The species composition of secondary forests
derived from abandoned pastures and coffee planta-
tions remained distinct from old-growth forest. In
addition to historic land use, age and elevation were
important environmental variables explaining varia-
tion in secondary forest species composition. Non-
indigenous species were common in recently aban-
doned pastures and coffee plantations, but their
importance declined in the older sites. This study
demonstrates that secondary forests on private land
can be an important component of the conservation
of tropical tree biodiversity.
Key words: biodiversity; chronosequence; coffee
plantation; land use history; non-metric multidimen-
sional scaling; pasture; Puerto Rico; secondary suc-
cession; tropical forests.
lar to intact forest in a relatively short time (Aide et
al. 1995, 1996, Zimmerman et al. 1995, Finegan
1996, but see Saldarriaga et al. 1988). In addition,
species richness in tropical secondary forests can be
as high or higher than old-growth forest (Brown
and Lugo 1990, Finegan 1996); however, species
composition is often different (Zimmerman et al.
1995, Guariguata et al. 1997). Nevertheless, these
secondary forests function similarly to primary for-
ests by protecting the soil from erosion, retaining
nutrients within the system and fixing large quanti-
ties of CO2 (Brown and Lugo 1990, Phillips et al.
1998). The recovery of species richness, structural
218 J. B. Pascarella and others
characteristics, and ecosystem function indicate a
high degree of resiliency to human perturbations
(Holling 1973), contrary to earlier studies that
emphasized the ‘‘fragility’’ of tropical forests (Rich-
ards 1964, Gómez-Pompa et al. 1972).
Many studies of tropical forest recovery have
focused on slash-and-burn agriculture or recently
abandoned pastures (Uhl 1987, Uhl et al. 1988).
Although these are important land-use practices, a
major limitation in our understanding of long-term
dynamics is the relatively young age (⬍30 years) of
the sites, in part, because many areas of the tropics
have been recently deforested. Only a few studies
have examined long-term forest chronosequences
(Saldarriaga et al. 1988, Corlett and Turner 1997,
Grau et al. 1997) and even when older sites exist, it
is often difficult to accurately determine the age and
previous land-use history. In addition, few studies
(Weaver and Birdsey 1986) have examined succes-
sional dynamics of abandoned agro-forestry systems
such as shade coffee plantations, although these
land uses are important components of the tropical
landscape. Finally, many studies have taken place in
a landscape context of small areas of secondary
forest that were used for a short term surrounded by
large, intact areas of mature or secondary forest.
Today a more typical tropical landscape would be
small forest remnants in a matrix of urban centers
and agricultural lands, often with abundant non-
indigenous species.
The Caribbean island of Puerto Rico presents an
opportunity to study long-term tropical secondary
forest succession using chronosequences from a
variety of land uses, soil types, and lifezones typical
of the Caribbean and mainland Neotropics. At the
beginning of the 20th century, greater than 90% of
Puerto Rico was in some form of agriculture and
remnant forest was limited to small patches. Since
the 1940’s most of these agricultural areas have
been abandoned as the island’s economy shifted
from agriculture to small industry (Dietz 1986).
Today, much of the island is covered in secondary
forest (Birdsey and Weaver 1987).
To understand better the pattern and process of
secondary forest succession, we developed the fol-
lowing hypotheses: 1) Because the regeneration of
tropical forest depends upon the intensity of previ-
ous land use and proximity to sources of colonizers
(Purata 1986, Uhl et al. 1988), we predicted that the
type of land use and distribution of remnant old-
growth forest patches would determine the rate of
forest recovery and species composition of second-
ary forests. Areas in short-term use and in close
proximity to remnant forest patches should recover
quicker than areas in long-term use or far from
remnant forests. 2) In an agriculture-dominated
landscape, shade coffee plantations contain as much
biodiversity for some animal taxa as forest habitats
(Perfecto et al. 1996). Because abandoned coffee
plantations have initial starting conditions more
similar to forest and might attract seed dispersers for
feeding, resting or nesting activities, we predicted
that basal area, species richness and species compo-
sition would recover faster in abandoned coffee
plantations than in abandoned pastures. 3) Intro-
duced non-indigenous species are important in
agricultural landscapes, either as planted pasture
grasses, tree or shrub crops, ornamentals, shade
trees, or weeds. Several hundred non-indigenous
tree species have been introduced into Puerto Rico
and many are naturalized (Little et al. 1974). Non-
indigenous species can interfere with regeneration
of forests through a variety of mechanisms (Horvitz
et al. 1998) and are important colonizers on other
tropical and subtropical oceanic islands (Smith 1984,
Thebaud and Strasberg 1997, Vitousek et al. 1997)
and mainland (Grau et al. 1997). We predicted that
non-indigenous species would be common in young
secondary forests, particularly those developing from
abandoned pastures where non-indigenous species
are abundant. To test these three hypotheses, we
described the land-use history of an agricultural
region in southeastern Puerto Rico over a 60-year
period (1937–1995) and measured the structure
and species composition of 33 secondary and 4
old-growth forest stands.
METHODS
Study Area, Land Use Analysis
and Study Sites
We conducted this study on private land in the
municipalities of Guayama and Patillas in the Sierra
de Cayey Mountains of southeastern Puerto Rico
(Fig. 1). The study area was limited by the boundary
between the Patillas and Arroyo municipalities to
the south, the Rio Grande de Patillas to the east, the
Carite State Forest Reserve to the north, and state
highway 179 to the west for a total map area of
2,443 ha. Elevations range from 100 m along the
Rio Grande de Patillas to over 700 m near the Carite
Forest Reserve. Rainfall ranges from 1520 mm/yr in
the interior valleys to 2540 mm/yr in the higher
mountains. Soils included volcanic soils, with Los
Guineos series (Ultisols) in the uplands and Caguabo
(Inceptisols) and Naranjito (Ultisols) soils in the
valleys and side slopes, and plutonic soils, primarily
Panduras series (Inceptisols) and limited areas of
Patillas soils (Ultisols) (Boccheciamp 1977). Accord-

ing to the Holdridge classification, the natural veg-
etation is Subtropical Moist Forest in the coastal
valleys and Subtropical Wet Forest in the slopes and
mountain tops (Ewel and Whitmore 1973).
We determined land use by analyzing 23 by 23
cm, black and white aerial photographs from 1937
(scale-1:17,268), 1967 (1:20,000), and 1995 (1:
20,000). Interviews with local residents and aerial
photographs from 1951, 1977, and 1988 provided
additional information on land-use history. The
photointerpretation was done using a mirror stereo-
scope and stereopair photographs. Based on fea-
tures on the photographs, we assigned six land-use
classes (Table 1). Field work in 1996 confirmed that,
of the 37 sites we visited, most areas were accurately
Forest Regeneration in Puerto Rico 219
Figure 1. The distribution of
land-use classes in the study
area during 1937, 1967, and
1995. Dense forest in 1937
included active shade coffee
plantations. Land-use classes
were determined using aerial
photo interpretation. See
Table 1 for classification.
assigned to the land-use classes from the 1995
photographs. Only a few sites assigned to the dense
forest class were actually coffee plantations, but
they could not be distinguished from forested sites
in the aerial photographs due to a relatively closed
canopy created by the shade trees in the plantations.
For each photograph, outlines of each land-use
polygon were traced on acetate overlays. Land-use
polygons from 1937, 1967, and 1995 were georefer-
enced, digitized, and entered into the ARC/INFO
version 6 geographic information system package
(ESRI 1991). The same control points were used in
each of the three years to maintain accurate registra-
tion and to allow comparison of land use among the
three study years. The total area of each land-use
220 J. B. Pascarella and others
Table 1. Land-Use Classes and
Identifying Features
Class Criteria
Sugar cane Even texture, uniform color, no trees, and
distinct plow lines
Pasture Dominated by grasses with no or little
woody vegetation
Shrub Dense shrubs with ⬍50% tree cover
Open forest ⬎50 to ⬍80% forest tree cover with an
even-structured canopy
Dense forest ⬎80% forest tree cover and uneven
canopy
Urban Clusters of buildings and houses, including
yards
Criteria were used to classify land use from aerial photographs of the study area in
1937, 1967, and 1995.
class was calculated for each year. The changes in
land use (e.g., sugar cane to pasture) from 1937 to
1967 and 1967 to 1995 were determined for each
class using an overlay method.
To provide a representative sample of the vegeta-
tion in the study area, 37 sites were arbitrarily
selected in areas that varied in land-use history, age,
elevation and soil type (Appendix 1). In addition,
site selection depended on accessibility and permis-
sion of landowners. Thirteen sites were located on
plutonic soils and 24 on volcanic soils. We used
aerial photographs from 1937, 1951, 1967, 1977,
1988, and 1995 to determine the land-use history
and time since abandonment at each site. Four sites
had continuous forest cover since 1937, twenty-
four had been used as pastures, and nine had been
used as coffee plantations. The age of continuously
forested sites was conservatively estimated at ⬎80
years. The age of sites derived from abandoned
pastures was estimated as the midpoint between the
year of the last photograph to show use as pasture
and the year of the first photograph to show signs of
abandonment (either shrub, open or dense forest,
see Table 1). Based on their appearance in the aerial
photographs from 1937 and their location adjacent
to existing pastures, we estimated that young forests
in 1937 were derived from abandoned pastures
(approximately 77 years old). Time cleared was
estimated as the length of time the site was in active
use as a pasture between 1937–1995. Local resi-
dents helped us locate abandoned coffee plantations
and explained that most plantations were aban-
doned in the early 1960s and, thus, we estimated
the age since abandonment to be 30 yr. We deter-
mined the distance to forest (open or dense) at time
of abandonment by measuring the distance from
the midpoint of the field to the edge of the closest
forest on the aerial photographs and converted the
distance into meters. Elevation and slope were
recorded for each site.
Forest Sampling and Data Analysis
We characterized the woody vegetation for each site
using two types of samples. We measured individu-
als ⬎1 cm in diameter at 1.3 m height within four
1 ⫻ 50 m belt transects that were located near the
middle of the stand and perpendicular to the slope.
Two additional 10 ⫻ 50 m transects were established
between the first and second, and the third and
fourth belt transects. In these larger transects, we
measured all woody shrubs and trees ⬎10 cm in
diameter at 1.3 m height. In total, we sampled all
trees and shrubs down to 1 cm within 200 m2 and all
trees down to 10 cm within 1000 m2. Species-area
curves showed that the sampling scheme was suffi-
cient to describe the site diversity (unpublished
data). Species names follow Liogier (1985, 1988,
1994, 1995, and 1997). Although woody vines were
present in the older stands, they were not sampled.
For each site, we determined the total number of
woody shrub and tree species, calculated basal areas
and stem densities, plotted the species-site curves,
and calculated species diversity using H8 (Shannon
diversity index) (Ludwig and Reynolds 1988). Spe-
cies richness, diversity, number of endemic species,
number of non-indigenous species, basal area, and
density were compared among the land-use/age
categories using one-way ANOVA. Basal area and
density were log-transformed (Statistix 1994). These
variables were also compared between the different
soil types for coffee. Grouped by age and elevation, a
paired t-test was used to compare the effect of soils
on basal area, stem density, and species richness for
forest derived from pasture. A subset regression
model was used to determine the best predictors of
the dependent variables density, basal area, species
richness, and species diversity (H8) for forests de-
rived from pastures. The independent variables
were time since abandonment, the square of the
time since abandonment (to detect non-linear pat-
terns), distance to nearest forest at time of abandon-
ment, elevation, slope, and time cleared between
1937 and 1995.
To examine the relationship of quantitative and
qualitative ecological factors on species composi-
tion, we used non-metric dimensional scaling
(NMDS) of sites and species’ importance values.
Species’ importance values were calculated as the
mean of percent relative basal area and percent
relative density. A PCA analysis was used to gener-
ate three axes for the initial starting configuration of
 Forest Regeneration in Puerto Rico 221

Figure 2. Total area (ha) and percentage of study area

covered by each land-use class from 1937 to 1995. The
different patterns in each bar and in the legend show the
relative contributions of other land uses from the previous
year to each land use listed on the abscissa. Data were not
available for 1937.

the NMDS. We used three axes, an Euclidean

distance measure, and 50 iterations to find the final
configuration of the ordination (McCune and Med-
ford 1997). A second matrix of environmental
variables versus sites was also created. Environmen-
tal variables considered were elevation, age (time
since abandonment), time cleared since 1937, dis-
tance to nearest forest at time of abandonment, and
slope. For statistical comparison of species composi-
tion, we grouped sites into five land-use/age catego-
ries: abandoned coffee, old-growth forest, young
forest from pasture (4–13 years), intermediate forest
from pasture (25–37), and old forest from pasture
(52–77 years). To examine the importance of quali-
tative group variables on variation in species compo-
sition within and among multiple samples, we used
a multivariate resampling technique (Clarke 1993)
to compare the variation among land-use/age cat-
egories.
RESULTS
Land use changed greatly between 1937, 1967, and
1995 (Figs. 1, 2). Forest, including both open and
dense forest, increased in area from 20% in 1937 to
62% in 1995. In 1995, most of the forest area was
classified as dense forest. Area in pastures remained
relatively unchanged from 1937 to 1967, then
declined in 1995. Shrub cover declined from 28% of
the area in 1937 to 13% in 1995. Sugar cane
cultivation declined in area from 5% in 1937 to
0.2% in 1967 and 0% in 1995. Only a small urban
area had developed by 1995. Most land-use catego-
ries, with the exception of shrubs, exhibited unidi-
rectional change over the entire 60-yr period. Nearly
Figure 3. Land-use/age categories and (A) mean stem
density (⬎1 cm)/ha (⫾1 SE) and (B) mean basal area
(m2/ha) (⫾1 SE). Lowercase letters indicate significant
differences among the groups (n ⫽ 4 except for coffee
(n ⫽ 9)).
all of the sugarcane was converted into pastures,
pastures changed to various forest categories, forests
remained forests and succeeded to dense forests.
Shrubs either were cleared for pasture or under-
went succession to open and dense forest.
Areas used for agriculture were not abandoned at
the same time nor were they abandoned equally at
all elevations. Because higher elevation pastures
were abandoned first, age of secondary forests from
pastures was positively correlated with elevation
(r ⫽ 0.73, P ⬍ 0.0001, n ⫽ 24) and years cleared
was strongly negatively correlated with elevation
(r ⫽ ⫺0.92, P ⬍ 0.0001, n ⫽ 24) (Appendix 1). All
old-growth forest sites were located above 650 m.
Abandoned coffee sites were found across the eleva-
tion gradient, ranging from 125 m to 650 m.
Stem density varied among land uses and showed
a strong relationship to age within the abandoned
pastures (Fig. 3A). Stem density ranged from 1,930
(site 3) to 16,030 (site 4) stems ha⫺1 (Appendix 1).
222 J. B. Pascarella and others
We found significant variation among land-use/age
categories in stem density (Fig. 3A, F7,28 ⫽ 4.86,
P ⬍ 0.01). In the ANOVA, site 4 was removed from
the analysis as a significant outlier. This site was the
only pasture that had been cleared for a short time,
abandoned, and then recleared. Resprouted, mul-
tistemmed shrubs of Miconia prasina produced this
unusually high stem density. In the regression
analysis, age2 and time cleared explained 44% of
the variance in stem density. In the abandoned
pastures, stem density increased from 4–37 years,
and then declined with age. Based on this pattern,
we separated abandoned pastures into an establish-
ment (4–37 years) and a thinning (52–77 years)
phase. In the establishment phase, stem density was
strongly positively correlated with age (r ⫽ 0.77,
P ⫽ 0.001, n ⫽ 15, with site 4 removed as an out-
lier). Abandoned coffee plantations were intermedi-
ate in stem density. In coffee plantations, stem
density of coffee (Coffea arabica) ranged from 50–
4900 stems/ha and was significantly correlated with
overall density (r ⫽ 0.76, P ⬍ 0.05, n ⫽ 9). Old-
growth forests had significantly lower stem density
than old forest from pasture (52–77 year) and
abandoned coffee plantations. Stem densities did
not vary in forest derived from pasture when
grouped by soil type, but abandoned coffee planta-
tions on plutonic soils had higher stem densities
than abandoned coffee plantations on volcanic soils
(t ⫽ ⫺3.3, P ⬍ 0.05, df ⫽ 7).
Basal area ranged from 4.08 (site 2) to 67.20 (site
33) m2/ha, and varied significantly among land-use/
age categories (Fig. 3B, F7,29 ⫽ 19.14, P ⬍ 0.0001,
Appendix 1). There was no difference in basal area
between forest from pasture or coffee sites differing
in soil type. In abandoned pastures, basal area was
positively correlated with age (r ⫽ 0.67, P ⬍ 0.001,
n ⫽ 24) and time since abandonment and distance
to remnant forest at time of abandonment ex-
plained 61% of the variation in pasture basal area.
In the 37 sites, 130 woody species from 42
families were identified, including 118 native spe-
cies and 12 non-indigenous species (Appendix 2). A
total of 15 species endemic to Puerto Rico were
found in all sites. We found significant variation
among land-use categories in mean number of
endemic species (F7,29 ⫽ 4.75, P ⬍ 0.01). Old-
growth forests had the greatest number of endemic
species, followed by old forest from pasture, interme-
diate forest from pasture, young forest from pasture,
and coffee plantations. There was no difference in
the mean number of endemic species between sites
of different soil types in abandoned coffee planta-
tions and forest from pasture. Based on average
importance value, endemic species were most impor-
tant in 25-yr old forest from pasture and old-growth
forest. Non-indigenous species occurred in 33 of the
37 sites (89%). However, only three species, Spatho-
dea campanulata, Syzygium jambos, and Coffea arabica,
occurred in more than 10 sites. Coffee had not
spread to other land-use types except for one adja-
cent pasture site. Although coffee plantations had
the greatest number of non-indigenous species, we
found no significant differences in the mean num-
ber of non-indigenous species neither among land-
use categories nor for forests on the two soil types.
However, using average importance value, non-
indigenous species were most important in young
forest (4–13 yr) derived from pasture and in aban-
doned coffee plantations.
We found significant variation among land-use/
age categories in species richness per site (Fig. 4A,
F7,29 ⫽ 7.42, P ⬍ 0.0001) and Shannon diversity
indices (H8) (F7,29 ⫽ 6.06, P ⬍ 0.0001) (Appendix
1). The two measures showed virtually identical
patterns. In forests from abandoned pastures, the
number of species increased with age (r ⫽ 0.59,
P ⬍ 0.01, n ⫽ 24), with a peak in intermediate aged
(37 yr) forests and a slight decline in older forests
(Fig. 4A). Age and age2 explained 61% of the
variation in species number among stands. Species
richness in old-growth forest and abandoned coffee
plantations was similar to older forest from pasture.
There was no difference in species richness between
coffee or forest from pasture sites that varied in soil
type. Shannon diversity indices (H8) ranged from
1.10 (site 8) to 3.01 (site 21) (Appendix 1). Species-
site curves varied among land-use/age categories
(Fig. 4B). Intermediate and old forest from pasture
sites had the greatest regional accumulation of
species, followed by abandoned coffee, while old-
growth forest and young forest from pasture had the
lowest.
Multivariate analysis of quantitative environmen-
tal variables separated sites along age and elevation
gradients. Non-metric multidimensional scaling ex-
plained a total of 92.3% of the variation in species’
importance values among the 37 sites. Axis I ex-
plained 63.4% of the variance in species’ impor-
tance values and ordered species along an eleva-
tional and age gradient (Table 2). Axis II explained
14.9% of the variation and separated sites based on
years cleared. Axis III explained 14% of the varia-
tion, separating sites along age, time cleared, eleva-
tion, and distance to remnant forest. Sites grouped
by major land-use categories were also separated
statistically using Clarke’s resampling method (Fig.
5). The observed value of the multivariate statistic R
was significantly different from the resampled statis-
tic (Ro ⫽ 0.35, Rs ⫽ 0.11, P ⬍ 0.01, n ⫽ 37), indicat-
 Forest Regeneration in Puerto Rico 223

Table 2. Kendall Correlation of Environmental

Variables with NMDS Ordination Axes 1–3

Environmental Axis 1 Axis 2 Axis 3

Variable (63.4%) (14.9%) (14.0%)

Age ⫺0.364 a 0.094 ⫺0.517 b

Years cleared 0.072 0.227 c 0.567 b
Distance to forest 0.073 0.118 0.465 b
Elevation ⫺0.423 a 0.13 ⫺0.472 b
Slope 0.109 ⫺0.151 0.003

The percent variation in the ordination explained by each axis is shown.

Probabilities indicate the significance of the variable in each axis. The total sample
size was 37.
aP ⬍ 0.001.
bP ⬍ 0.0001.
cP ⬍ 0.05.

Figure 4. Land-use/age categories and (A) mean species

richness (direct counts of species) (⫾1 SE) and (B)
species-site curves. For species richness, lowercase letters
indicate significant differences among the groups (n ⫽ 4
except for coffee (n ⫽ 9)). For species-site curves, second-
ary forests derived from pastures were combined into
young (4–13 yr), intermediate (25–37 yr), and old stands
(52–77 yr). All sites had 0.12 ha sampled.

ing that the different land-uses/age categories dif-
fered in species composition.
DISCUSSION
Land-use History and Forest Regeneration
The study region showed a dramatic temporal and
spatial shift in forest cover in the 60-year period,
from almost complete deforestation to a predomi-
nantly forested landscape (⬎60% by 1995). This
increase in forest cover was greater than the island-
wide pattern of approximately 35% forest cover
noted in the 1980s (Birdsey and Weaver 1987),
most likely due to the mountainous terrain in the
Figure 5. Non-metric multidimensional scaling of sites in
species’ space. The three axes (I–III) represent the location
of the 37 sites in multivariate space, based on the species’
importance values found in each site.
study area. The first sites to be abandoned were at
the high elevations as has been observed in other
studies in Puerto Rico (Thomlinson et al. 1996). This
pattern of abandonment leads to a positive correla-
tion between elevation and time since abandon-
ment and eliminates the possibility of an indepen-
dent analysis of site age and elevation.
Although correlated with elevation, time since
abandonment was the best predictor of forest recov-
224 J. B. Pascarella and others
ery. Basal area and species diversity of secondary
forests derived from abandoned pastures and coffee
plantations reached values similar to old-growth
forest in 25–30 yr. Compared to other Neotropical
secondary forests in similar lifezones developing on
abandoned pastures, the rate of recovery of basal
area in Puerto Rico is slower than in Costa Rica
(Guariguata et al. 1997). The slower rate of recovery
is probably due to a longer period of use (decades in
Puerto Rico versus years in Costa Rica). Given that
the dominant land use in the region was cattle
pastures, often for more than 50 years, we believe
there was little regeneration by resprouting of old-
growth forest tree species, as has been described
following short-term slash and burn agriculture
(Uhl 1987).
Our hypothesis that the distance to remnant
forest at time of abandonment would have a signifi-
cant effect on forest recovery was not supported. All
sites were either adjacent to or within 200 m of
remnant forest at the time of abandonment and,
thus, there was little variation in this variable. The
wide variation in species composition in very young
abandoned pastures suggests that chance events
determine which species arrive and initially capture
a site (Purata 1986). Two main groups of colonizing
species were found in abandoned pastures. One
group included small-seeded animal dispersed shrub
or small tree species (e.g., Casearia spp., Miconia spp.,
and Psidium guajava) that are all able to resprout
after being cut. This characteristic may allow these
species to remain in active pastures as sprouts until
the site is abandoned. A second group included two
large trees, Tabebuia heterophylla and Spathodea cam-
panulata, that are often used as living fences and
whose seeds are widely dispersed by wind. Many
large-seeded species that persisted in the remnant
forests (e.g., Dacryodes excelsa, Prestoea montana, Ma-
nilkara bidentata, and Sloanea berteriana) were rare or
absent in the secondary forest. Their absence may
be due to changes in frugivore species composition,
abundance, and foraging behavior (Wunderle 1997),
as well as an inability of some species typical of
mature forest to survive in early successional com-
munities (Uhl 1987, Brown and Lugo 1990, but see
Nepstad et al. 1996).
The study region represents 0.2% of the total land
area of Puerto Rico yet contains 118 native trees and
shrubs, or 22% of the woody species diversity of the
island (Little et al. 1974). Fifteen endemic species,
including a federally endangered endemic tree,
Eugenia haemetocarpa, occurred in the study sites.
The shape of the species-site curves for the different
land-use categories suggests that the total regional
diversity of the area may even be higher than that
for the 37 sampled plots. These findings demon-
strate that small remnant stands of forest (⬍10% of
the area, excluding the known coffee plantations)
that existed in 1937 were important sources for the
current biodiversity of the area, even though many
late successional species have not yet spread to the
secondary forests that now dominate the landscape.
Similar results have been found in regenerating
forests in Hong Kong and Singapore (Turner and
Corlett 1996, Corlett and Turner 1997).
Forest Recovery in Shade Coffee
Versus Pastures
Abandoned coffee plantations had the highest mean
basal area for all land-use/age categories and lower
mean species richness in comparison to similarly
aged forest derived from pasture. The major differ-
ence between abandoned coffee plantations and
abandoned pastures is the starting conditions when
they were abandoned. The coffee plantations al-
ready have a high density of woody plants and
shade trees that account for the rapid increase in
basal area. Although frugivorous birds may increase
seed dispersal into shade coffee plantations, this
apparently does not result in higher species diver-
sity, suggesting that seed germination or seedling
survival of dispersed seeds is low. Although active
shade coffee plantations are better habitats for some
animal species in comparison with cattle pastures
(Perfecto et al. 1996), once these land uses are
abandoned, forest from pasture accumulates plant
biodiversity more rapidly.
Instead of facilitating colonization, the existing
forest structure of abandoned coffee plantations
appears to inhibit the recruitment of native woody
species. Similar patterns have been observed in
abandoned coffee plantations and pastures in the
Cordillera Central and karst regions of Puerto Rico
(Birdsey and Weaver 1982, Rivera and Aide 1998)
and in abandoned cacao plantations and pastures in
the Dominican Republic (Rivera et al., in press).
Despite considerable variation in elevations (120–
675 m), coffee stands in the region were remarkably
similar in composition due to the dominance of the
Meliaceous tree Guarea guidonia. Guarea guidonia has
a rapid growth response to increased soil nitrogen
availability (Fernandez 1997), which is typical of
abandoned coffee plantations. This may explain
why Guarea guidonia dominates abandoned coffee
plantations throughout Puerto Rico (Weaver and
Birdsey 1986, Zimmerman et al. 1995). The high
abundance of Guarea guidonia seedlings in aban-
doned coffee plantations suggests that this species
will continue to dominate these sites for many years
(Rivera and Aide 1998).

Non-indigenous Species and
Secondary Forests
The hypothesis that non-indigenous species would
be important components of secondary forests devel-
oping from pastures was only partially supported.
Although non-indigenous species were diverse and
abundant in young forests derived from abandoned
pastures (ⱕ25 yr old) and in abandoned coffee
plantations, their diversity and importance declined
in older secondary forests derived from pasture and
in the old-growth forests. For example, the wind-
dispersed tree Spathodea campanulata dominated the
forest canopy layer, while Psidium guajava was com-
mon in the shrub layer in young secondary forests
derived from pasture. Nevertheless, in the 25-yr old
sites, both species appeared to be unable to recruit
seedlings and saplings below a dense canopy (Pas-
carella, personal observation). Both species declined
in relative importance value in abandoned pastures
37 yrs or older. These species appear to facilitate
invasion of secondary forests by shade-tolerant
native species (Wadsworth and Birdsey 1985).
In contrast to Spathodea campanulata, the non-
indigenous tree Syzygium jambos (Myrtaceae) was
primarily found in old pastures, abandoned coffee
plantations, and old forest, indicating it is able to
invade closed-canopy forests and tolerate deep
shade. Syzygium jambos was also an important com-
ponent of secondary forests in the Luquillo region,
often forming dense groves (Little and Wadsworth
1964, Aide et al. 1996). The coppicing ability of
Syzygium jambos may allow it to dominate areas
following hurricane disturbance (Aide et al. 1996)
and the shade-tolerant seedling can dominate the
seedling-layer regeneration niche (Horvitz et al.
1998). Long-term monitoring and experimental
removals are needed to verify the role of introduced
non-indigenous species in secondary succession.
Conclusions
In spite of almost complete deforestation, this tropi-
cal agricultural landscape recovered to a forest
dominated landscape in 60 yr. Our study suggests
that conservation of small forest remnants, even in
highly deforested landscapes, is important in conser-
vation of regional biodiversity. In areas continuing
to undergo deforestation, financial incentives to
farmers to maintain small areas of native forest
cover within larger agricultural landscapes should
be encouraged.
ACKNOWLEDGMENTS
We thank the landowners in the Sierra de Cayey
who allowed us to work on their properties and
Forest Regeneration in Puerto Rico 225
provided information on agricultural practices.
Anaeli Cintron, David Baez, Humfredo Marcano,
and Caleb Gonzalez assisted with field work and
data entry. Frank Axelrod of the University of
Puerto Rico-Rio Piedras helped identify plant speci-
mens. Manuel Guariguata and Katherine Ewel pro-
vided comments on an earlier draft of this manu-
script. This research was funded by a grant under
the NASA Institutional Research Award program to
the University of Puerto Rico and the NSF-CREST
program.
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 Forest Regeneration in Puerto Rico 227

Appendix 1. Environmental and Stand Characteristics of Sampled Forests

Time Basal Species

Age Soil Elevation Cleared Area Density Number of Diversity
Site (y) Type (m) (y) (m2 /ha) (ha) Species (H8)

1p 4 p 300 54 6 2980 7 1.12

2p 4 v 350 54 4.1 3130 8 1.59
3p 4 p 400 54 8.4 1930 15 2
4p 4 v 620 19 7.7 16030 11 1.1
5p 13 p 300 46 21 5330 12 1.58
6p 13 v 325 46 30 5170 11 1.29
7p 13 p 500 46 24 7890 22 1.66
8p 13 v 550 46 18 6670 25 2.79
9p 25 p 300 35 29 8720 16 1.83
10p 25 v 350 35 54 9670 23 1.88
11p 25 p 500 35 17 7643 27 2.55
12p 25 v 625 35 22 8330 29 2.63
13p 37 p 500 22 23 8060 28 2.8
14p 37 v 550 22 29 5060 32 3.01
15p 37 p 600 22 25 12150 25 2.37
16p 37 v 625 22 38 12,520 31 2.35
17p 52 v 650 7 39 6310 31 2.76
18p 52 v 650 7 41 3380 16 2.19
19p 52 v 670 7 32 9220 33 2.91
20p 52 v 700 7 28 7430 28 2.74
21p 77 v 625 0 27 9260 33 3.01
22p 77 v 650 0 49 4090 22 2.28
23p 77 v 700 0 36 3570 27 2.67
24p 77 v 710 0 36 4500 21 2.44
25c 30 p 125 0 51 5920 23 2.19
26c 30 v 325 0 27 4630 23 2.47
27c 30 v 325 0 32 5630 20 2.24
28c 30 p 450 0 32 5850 20 2.36
29c 30 p 500 0 51 8240 23 2.33
30c 30 p 550 0 40 8660 22 2.2
31c 30 p 650 0 34 8990 19 2.31
32c 30 v 675 0 41 2610 21 2.44
33c 30 v 650 0 67 4670 21 2.26
34f ⬎80 v 650 0 35 3620 27 2.7
35f ⬎80 v 670 0 45 4360 27 2.57
36f ⬎80 v 680 0 30 4590 29 2.74
37f ⬎80 v 700 0 49 3550 25 2.73

Secondary forests included sites 1–24p derived from abandoned pastures and sites 25–33c derived from abandoned coffee plantations. Sites 34–37f are old-growth forests. Soil
types are plutonic (p) and volcanic (v). Within a given age and land use, sites are arranged in order of increasing elevation. All sites had 0.12 ha sampled.
228 J. B. Pascarella and others

Appendix 2. Mean Importance Value of Woody Species by Land-Use/Age Category

Land Use and Age

Old-
Forest Derived from Pasture (y) Growth
Forest Abandoned
Species (Family) 4 13 25 37 52 77 (⬎80 y) Coffee

Alchornea latifolia (Euphorbiaceae) 0 0.69 0.44 3.15 4.08 3.6 2.4 5.46
Andira inermis (Fabaceae) 8.42 4.37 2.35 0 0 0.7 0 2.1
Artocarpus altilis a (Moraceae) 0 0 0 0 0 0 0 3.16
Casearia arborea (Flacourtiaceae) 1.06 0 3.4 1.65 7.58 3.8 5.2 0.88
Casearia guianensis (Flacourtiaceae) 10.64 5.64 10.68 4.1 0.11 0 0 0.19
Casearia sylvestris (Flacourtiaceae) 0.38 11.09 14.55 5.36 1.63 0.7 0.2 0.68
Cecropia schreberiana (Moraceae) 0 0.67 0.66 3.02 6.54 3.1 2.1 3.79
Ceiba pentandra (Bombacaceae) 0 0 0 0 0 0 0 1.33
Cinnamomum elongatum (Lauraceae) 6.32 0.23 0.59 1.12 0.09 0 0 2.53
Citharexylum caudatum (Verbenaceae) 0 0 0.24 0.52 4.71 2.2 0 0
Citharexylum fruticosum (Verbenaceae) 1.54 1.17 1.28 0.95 0 0 0 0.05
Coccoloba swartzii (Polygonaceae) 0 0 0 2.33 0.35 0.4 0 0
Coffea arabica a (Rubiaceae) 0 0 0 0.64 0.09 0 0 13.2
Cordia alliodora (Boraginaceae) 0 0 0 0.91 2.65 0 0 3.12
Cordia borinquensis b (Boraginaceae) 0 0.15 0.16 0.17 2.55 1.4 4.2 0.03
Dacryodes excelsa (Burseraceae) 0 0 0 0 0.17 0 5.2 0
Daphnopsis americana (Thymeleaceae) 1.85 1.49 1.04 1.18 0 0 0 0
Dendropanax arboreus (Araliaceae) 0.87 0.08 1.68 0.87 0.31 0 0 5.95
Drypetes glauca (Euphorbiaceae) 0 0 0 0 0 0.2 4.2 0
Faramea occidentalis (Rubiaceae) 0 0 3.25 2.25 0 0.7 0 0
Genipa americana (Rubiaceae) 0 0 0 0 0 0 0 0.93
Guarea guidonia (Meliaceae) 0 2.68 7.58 3.2 0.62 0.3 0 19.6
Hibiscus pernambucensis (Malvaceae) 0 0 0 2.1 1.87 0 0 0.35
Homalium racemosum (Flacourtiaceae) 3.07 0 1.87 2.59 0.17 1.9 3.3 0.03
Inga laurina (Fabaceae) 0 0.15 0.27 0.17 3.04 8.9 0.7 0.06
Inga vera (Fabaceae) 0 0 0 1.25 1.67 0.5 0 3.41
Ixora ferrea (Rubiaceae) 0 0 0.53 0.18 0 0.3 1.3 0
Micropholis guyanensis (Sapotaceae) 0 0 0 0 0 2.2 6.2 0
Miconia prasina (Melastomataceae) 17.68 3.21 0.5 0.67 1.28 0.3 0.2 0
Myrcia deflexa (Myrtaceae) 0 0.56 2.07 3.37 8.37 6.4 1.5 0
Myrcia splendens (Myrtaceae) 0.98 4.56 1.13 9.78 2.55 3.2 0 2.02
Ocotea leucoxylon (Lauraceae) 0.22 2 2.09 12.85 14.9 11 7.2 8.8
Pouteria multiflora (Sapotaceae) 0 0 0 0 0.31 0 0 2.28
Prestoea montana (Arecaceae) 0.99 0.18 0 0.79 14.06 17 25 4.87
Psidium guajava a (Myrtaceae) 13.98 2.71 0.48 0.07 0 0 0 0
Quararibea turbinata (Bombacaceae) 0 0 0 0 0 0 0 2.01
Schefflera morototoni (Araliaceae) 0 1.19 1.47 1.56 2.25 0.5 2.7 0.95
Sloanea berteriana (Eleocarpaceae) 0 0 0 0 0 0.3 5.3 0
Spathodea campanulata a (Bignoniaceae) 20.62 29.1 20.67 0 0.1 0 0 2.36
Syzygium jambos a (Myrtaceae) 0 1.36 0.14 0.53 1.22 7.3 1.7 0.52
Tabebuia heterophylla (Bignoniaceae) 5.7 16.56 2.98 16.73 3.41 8 0 0
Tetrazygium urbanii b (Melastomataceae) 0.2 0 6.08 0.26 0 1.4 1 0
Trichilia pallida (Rosaceae) 0 0.9 0.76 2.78 0 0.2 0 0.9
Urera baccifera (Urticaceae) 0 0 0 0 0.3 0 0 1.5
Zanthoxylum martinicense (Rutaceae) 0 0 0.74 0.43 0.19 0 1 1.02

Species’ importance values were calculated as the mean of percent relative basal area and percent relative density. Only species that had greater than 5% importance value in at
least one site are listed. Peak importance is highlighted in boldface.
aNonindigenous species.
bPuerto Rican endemic species.