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ABSTRACT
Although deforestation continues to be a major forest sites derived from abandoned pastures and
threat to tropical biodiversity, abandonment of agri- coffee plantations were similar to old-growth forest
cultural land in Puerto Rico provides an opportunity sites. The species composition of secondary forests
to study long-term patterns of secondary forest derived from abandoned pastures and coffee planta-
regeneration. Using aerial photographs from 1937, tions remained distinct from old-growth forest. In
1967, and 1995, we determined land-use history for addition to historic land use, age and elevation were
2443 ha in the Cayey Mountains. Pastures were the important environmental variables explaining varia-
dominant land cover in 1937 and ⬍20% of the area tion in secondary forest species composition. Non-
was classified as forest. Between 1937 and 1995, indigenous species were common in recently aban-
forest cover increased to 62% due to widespread doned pastures and coffee plantations, but their
abandonment of agriculture. To examine the effect importance declined in the older sites. This study
of historic land use on current forest structure and demonstrates that secondary forests on private land
species composition, we sampled secondary forests can be an important component of the conservation
in 24 abandoned pastures, 9 abandoned coffee of tropical tree biodiversity.
plantations and 4 old-growth forest sites. Sites were
located on two soil types along an elevational Key words: biodiversity; chronosequence; coffee
gradient (125–710 m) and included a chronose- plantation; land use history; non-metric multidimen-
quence from 4 to over 80 years old. After 25–30 sional scaling; pasture; Puerto Rico; secondary suc-
years, basal area and species richness in secondary cession; tropical forests.
Valdosta State University, Valdosta, Georgia 31698, USA. ties of CO2 (Brown and Lugo 1990, Phillips et al.
*Corresponding author; e-mail: jbpascar@valdosta.edu 1998). The recovery of species richness, structural
217
218 J. B. Pascarella and others
ing to the Holdridge classification, the natural veg- assigned to the land-use classes from the 1995
etation is Subtropical Moist Forest in the coastal photographs. Only a few sites assigned to the dense
valleys and Subtropical Wet Forest in the slopes and forest class were actually coffee plantations, but
mountain tops (Ewel and Whitmore 1973). they could not be distinguished from forested sites
We determined land use by analyzing 23 by 23 in the aerial photographs due to a relatively closed
cm, black and white aerial photographs from 1937 canopy created by the shade trees in the plantations.
(scale-1:17,268), 1967 (1:20,000), and 1995 (1: For each photograph, outlines of each land-use
20,000). Interviews with local residents and aerial polygon were traced on acetate overlays. Land-use
photographs from 1951, 1977, and 1988 provided polygons from 1937, 1967, and 1995 were georefer-
additional information on land-use history. The enced, digitized, and entered into the ARC/INFO
photointerpretation was done using a mirror stereo- version 6 geographic information system package
scope and stereopair photographs. Based on fea- (ESRI 1991). The same control points were used in
tures on the photographs, we assigned six land-use each of the three years to maintain accurate registra-
classes (Table 1). Field work in 1996 confirmed that, tion and to allow comparison of land use among the
of the 37 sites we visited, most areas were accurately three study years. The total area of each land-use
220 J. B. Pascarella and others
Table 1. Land-Use Classes and the midpoint of the field to the edge of the closest
Identifying Features forest on the aerial photographs and converted the
distance into meters. Elevation and slope were
Class Criteria recorded for each site.
Sugar cane Even texture, uniform color, no trees, and
distinct plow lines Forest Sampling and Data Analysis
Pasture Dominated by grasses with no or little We characterized the woody vegetation for each site
woody vegetation using two types of samples. We measured individu-
Shrub Dense shrubs with ⬍50% tree cover als ⬎1 cm in diameter at 1.3 m height within four
Open forest ⬎50 to ⬍80% forest tree cover with an
1 ⫻ 50 m belt transects that were located near the
even-structured canopy
Dense forest ⬎80% forest tree cover and uneven
middle of the stand and perpendicular to the slope.
canopy Two additional 10 ⫻ 50 m transects were established
Urban Clusters of buildings and houses, including between the first and second, and the third and
yards fourth belt transects. In these larger transects, we
measured all woody shrubs and trees ⬎10 cm in
Criteria were used to classify land use from aerial photographs of the study area in diameter at 1.3 m height. In total, we sampled all
1937, 1967, and 1995.
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-
class was calculated for each year. The changes in cient to describe the site diversity (unpublished
land use (e.g., sugar cane to pasture) from 1937 to data). Species names follow Liogier (1985, 1988,
1967 and 1967 to 1995 were determined for each 1994, 1995, and 1997). Although woody vines were
class using an overlay method. present in the older stands, they were not sampled.
To provide a representative sample of the vegeta- For each site, we determined the total number of
tion in the study area, 37 sites were arbitrarily woody shrub and tree species, calculated basal areas
selected in areas that varied in land-use history, age, and stem densities, plotted the species-site curves,
elevation and soil type (Appendix 1). In addition, and calculated species diversity using H8 (Shannon
site selection depended on accessibility and permis- diversity index) (Ludwig and Reynolds 1988). Spe-
sion of landowners. Thirteen sites were located on cies richness, diversity, number of endemic species,
plutonic soils and 24 on volcanic soils. We used number of non-indigenous species, basal area, and
aerial photographs from 1937, 1951, 1967, 1977, density were compared among the land-use/age
1988, and 1995 to determine the land-use history categories using one-way ANOVA. Basal area and
and time since abandonment at each site. Four sites density were log-transformed (Statistix 1994). These
had continuous forest cover since 1937, twenty- variables were also compared between the different
four had been used as pastures, and nine had been soil types for coffee. Grouped by age and elevation, a
used as coffee plantations. The age of continuously paired t-test was used to compare the effect of soils
forested sites was conservatively estimated at ⬎80 on basal area, stem density, and species richness for
years. The age of sites derived from abandoned forest derived from pasture. A subset regression
pastures was estimated as the midpoint between the model was used to determine the best predictors of
year of the last photograph to show use as pasture the dependent variables density, basal area, species
and the year of the first photograph to show signs of richness, and species diversity (H8) for forests de-
abandonment (either shrub, open or dense forest, rived from pastures. The independent variables
see Table 1). Based on their appearance in the aerial were time since abandonment, the square of the
photographs from 1937 and their location adjacent time since abandonment (to detect non-linear pat-
to existing pastures, we estimated that young forests terns), distance to nearest forest at time of abandon-
in 1937 were derived from abandoned pastures ment, elevation, slope, and time cleared between
(approximately 77 years old). Time cleared was 1937 and 1995.
estimated as the length of time the site was in active To examine the relationship of quantitative and
use as a pasture between 1937–1995. Local resi- qualitative ecological factors on species composi-
dents helped us locate abandoned coffee plantations tion, we used non-metric dimensional scaling
and explained that most plantations were aban- (NMDS) of sites and species’ importance values.
doned in the early 1960s and, thus, we estimated Species’ importance values were calculated as the
the age since abandonment to be 30 yr. We deter- mean of percent relative basal area and percent
mined the distance to forest (open or dense) at time relative density. A PCA analysis was used to gener-
of abandonment by measuring the distance from ate three axes for the initial starting configuration of
Forest Regeneration in Puerto Rico 221
We found significant variation among land-use/age tant in 25-yr old forest from pasture and old-growth
categories in stem density (Fig. 3A, F7,28 ⫽ 4.86, forest. Non-indigenous species occurred in 33 of the
P ⬍ 0.01). In the ANOVA, site 4 was removed from 37 sites (89%). However, only three species, Spatho-
the analysis as a significant outlier. This site was the dea campanulata, Syzygium jambos, and Coffea arabica,
only pasture that had been cleared for a short time, occurred in more than 10 sites. Coffee had not
abandoned, and then recleared. Resprouted, mul- spread to other land-use types except for one adja-
tistemmed shrubs of Miconia prasina produced this cent pasture site. Although coffee plantations had
unusually high stem density. In the regression the greatest number of non-indigenous species, we
analysis, age2 and time cleared explained 44% of found no significant differences in the mean num-
the variance in stem density. In the abandoned ber of non-indigenous species neither among land-
pastures, stem density increased from 4–37 years, use categories nor for forests on the two soil types.
and then declined with age. Based on this pattern, However, using average importance value, non-
we separated abandoned pastures into an establish- indigenous species were most important in young
ment (4–37 years) and a thinning (52–77 years) forest (4–13 yr) derived from pasture and in aban-
phase. In the establishment phase, stem density was doned coffee plantations.
strongly positively correlated with age (r ⫽ 0.77, We found significant variation among land-use/
P ⫽ 0.001, n ⫽ 15, with site 4 removed as an out- age categories in species richness per site (Fig. 4A,
lier). Abandoned coffee plantations were intermedi- F7,29 ⫽ 7.42, P ⬍ 0.0001) and Shannon diversity
ate in stem density. In coffee plantations, stem indices (H8) (F7,29 ⫽ 6.06, P ⬍ 0.0001) (Appendix
density of coffee (Coffea arabica) ranged from 50– 1). The two measures showed virtually identical
4900 stems/ha and was significantly correlated with patterns. In forests from abandoned pastures, the
overall density (r ⫽ 0.76, P ⬍ 0.05, n ⫽ 9). Old- number of species increased with age (r ⫽ 0.59,
growth forests had significantly lower stem density P ⬍ 0.01, n ⫽ 24), with a peak in intermediate aged
than old forest from pasture (52–77 year) and (37 yr) forests and a slight decline in older forests
abandoned coffee plantations. Stem densities did (Fig. 4A). Age and age2 explained 61% of the
not vary in forest derived from pasture when variation in species number among stands. Species
grouped by soil type, but abandoned coffee planta- richness in old-growth forest and abandoned coffee
tions on plutonic soils had higher stem densities plantations was similar to older forest from pasture.
than abandoned coffee plantations on volcanic soils There was no difference in species richness between
(t ⫽ ⫺3.3, P ⬍ 0.05, df ⫽ 7). coffee or forest from pasture sites that varied in soil
Basal area ranged from 4.08 (site 2) to 67.20 (site type. Shannon diversity indices (H8) ranged from
33) m2/ha, and varied significantly among land-use/ 1.10 (site 8) to 3.01 (site 21) (Appendix 1). Species-
age categories (Fig. 3B, F7,29 ⫽ 19.14, P ⬍ 0.0001, site curves varied among land-use/age categories
Appendix 1). There was no difference in basal area (Fig. 4B). Intermediate and old forest from pasture
between forest from pasture or coffee sites differing sites had the greatest regional accumulation of
in soil type. In abandoned pastures, basal area was species, followed by abandoned coffee, while old-
positively correlated with age (r ⫽ 0.67, P ⬍ 0.001, growth forest and young forest from pasture had the
n ⫽ 24) and time since abandonment and distance lowest.
to remnant forest at time of abandonment ex- Multivariate analysis of quantitative environmen-
plained 61% of the variation in pasture basal area. tal variables separated sites along age and elevation
In the 37 sites, 130 woody species from 42 gradients. Non-metric multidimensional scaling ex-
families were identified, including 118 native spe- plained a total of 92.3% of the variation in species’
cies and 12 non-indigenous species (Appendix 2). A importance values among the 37 sites. Axis I ex-
total of 15 species endemic to Puerto Rico were plained 63.4% of the variance in species’ impor-
found in all sites. We found significant variation tance values and ordered species along an eleva-
among land-use categories in mean number of tional and age gradient (Table 2). Axis II explained
endemic species (F7,29 ⫽ 4.75, P ⬍ 0.01). Old- 14.9% of the variation and separated sites based on
growth forests had the greatest number of endemic years cleared. Axis III explained 14% of the varia-
species, followed by old forest from pasture, interme- tion, separating sites along age, time cleared, eleva-
diate forest from pasture, young forest from pasture, tion, and distance to remnant forest. Sites grouped
and coffee plantations. There was no difference in by major land-use categories were also separated
the mean number of endemic species between sites statistically using Clarke’s resampling method (Fig.
of different soil types in abandoned coffee planta- 5). The observed value of the multivariate statistic R
tions and forest from pasture. Based on average was significantly different from the resampled statis-
importance value, endemic species were most impor- tic (Ro ⫽ 0.35, Rs ⫽ 0.11, P ⬍ 0.01, n ⫽ 37), indicat-
Forest Regeneration in Puerto Rico 223
ing that the different land-uses/age categories dif- Figure 5. Non-metric multidimensional scaling of sites in
fered in species composition. 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.
DISCUSSION
Land-use History and Forest Regeneration study area. The first sites to be abandoned were at
The study region showed a dramatic temporal and the high elevations as has been observed in other
spatial shift in forest cover in the 60-year period, studies in Puerto Rico (Thomlinson et al. 1996). This
from almost complete deforestation to a predomi- pattern of abandonment leads to a positive correla-
nantly forested landscape (⬎60% by 1995). This tion between elevation and time since abandon-
increase in forest cover was greater than the island- ment and eliminates the possibility of an indepen-
wide pattern of approximately 35% forest cover dent analysis of site age and elevation.
noted in the 1980s (Birdsey and Weaver 1987), Although correlated with elevation, time since
most likely due to the mountainous terrain in the abandonment was the best predictor of forest recov-
224 J. B. Pascarella and others
ery. Basal area and species diversity of secondary for the 37 sampled plots. These findings demon-
forests derived from abandoned pastures and coffee strate that small remnant stands of forest (⬍10% of
plantations reached values similar to old-growth the area, excluding the known coffee plantations)
forest in 25–30 yr. Compared to other Neotropical that existed in 1937 were important sources for the
secondary forests in similar lifezones developing on current biodiversity of the area, even though many
abandoned pastures, the rate of recovery of basal late successional species have not yet spread to the
area in Puerto Rico is slower than in Costa Rica secondary forests that now dominate the landscape.
(Guariguata et al. 1997). The slower rate of recovery Similar results have been found in regenerating
is probably due to a longer period of use (decades in forests in Hong Kong and Singapore (Turner and
Puerto Rico versus years in Costa Rica). Given that Corlett 1996, Corlett and Turner 1997).
the dominant land use in the region was cattle
pastures, often for more than 50 years, we believe Forest Recovery in Shade Coffee
there was little regeneration by resprouting of old- Versus Pastures
growth forest tree species, as has been described Abandoned coffee plantations had the highest mean
following short-term slash and burn agriculture basal area for all land-use/age categories and lower
(Uhl 1987). mean species richness in comparison to similarly
Our hypothesis that the distance to remnant aged forest derived from pasture. The major differ-
forest at time of abandonment would have a signifi- ence between abandoned coffee plantations and
cant effect on forest recovery was not supported. All abandoned pastures is the starting conditions when
sites were either adjacent to or within 200 m of they were abandoned. The coffee plantations al-
remnant forest at the time of abandonment and, ready have a high density of woody plants and
thus, there was little variation in this variable. The shade trees that account for the rapid increase in
wide variation in species composition in very young basal area. Although frugivorous birds may increase
abandoned pastures suggests that chance events seed dispersal into shade coffee plantations, this
determine which species arrive and initially capture apparently does not result in higher species diver-
a site (Purata 1986). Two main groups of colonizing sity, suggesting that seed germination or seedling
species were found in abandoned pastures. One survival of dispersed seeds is low. Although active
group included small-seeded animal dispersed shrub shade coffee plantations are better habitats for some
or small tree species (e.g., Casearia spp., Miconia spp., animal species in comparison with cattle pastures
and Psidium guajava) that are all able to resprout (Perfecto et al. 1996), once these land uses are
after being cut. This characteristic may allow these abandoned, forest from pasture accumulates plant
species to remain in active pastures as sprouts until biodiversity more rapidly.
the site is abandoned. A second group included two Instead of facilitating colonization, the existing
large trees, Tabebuia heterophylla and Spathodea cam- forest structure of abandoned coffee plantations
panulata, that are often used as living fences and appears to inhibit the recruitment of native woody
whose seeds are widely dispersed by wind. Many species. Similar patterns have been observed in
large-seeded species that persisted in the remnant abandoned coffee plantations and pastures in the
forests (e.g., Dacryodes excelsa, Prestoea montana, Ma- Cordillera Central and karst regions of Puerto Rico
nilkara bidentata, and Sloanea berteriana) were rare or (Birdsey and Weaver 1982, Rivera and Aide 1998)
absent in the secondary forest. Their absence may and in abandoned cacao plantations and pastures in
be due to changes in frugivore species composition, the Dominican Republic (Rivera et al., in press).
abundance, and foraging behavior (Wunderle 1997), Despite considerable variation in elevations (120–
as well as an inability of some species typical of 675 m), coffee stands in the region were remarkably
mature forest to survive in early successional com- similar in composition due to the dominance of the
munities (Uhl 1987, Brown and Lugo 1990, but see Meliaceous tree Guarea guidonia. Guarea guidonia has
Nepstad et al. 1996). a rapid growth response to increased soil nitrogen
The study region represents 0.2% of the total land availability (Fernandez 1997), which is typical of
area of Puerto Rico yet contains 118 native trees and abandoned coffee plantations. This may explain
shrubs, or 22% of the woody species diversity of the why Guarea guidonia dominates abandoned coffee
island (Little et al. 1974). Fifteen endemic species, plantations throughout Puerto Rico (Weaver and
including a federally endangered endemic tree, Birdsey 1986, Zimmerman et al. 1995). The high
Eugenia haemetocarpa, occurred in the study sites. abundance of Guarea guidonia seedlings in aban-
The shape of the species-site curves for the different doned coffee plantations suggests that this species
land-use categories suggests that the total regional will continue to dominate these sites for many years
diversity of the area may even be higher than that (Rivera and Aide 1998).
Forest Regeneration in Puerto Rico 225
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Forest Regeneration in Puerto Rico 227
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
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.