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Forest Ecology

and
Management
ELSEVI ER
Forest Ecology and Management 81 (1996) 215-226
The role of exotic conifer plantations in rehabilitating degraded
tropical forest lands: A case study from the Kibale Forest
in Uganda
Robert A. Fimbel *, Cheryl C. Fimbel
Wildlife Conseruation Society, 185th Street and Southern Bouleuard, Bronx, New York, NY 10460, USA
Accepted 21 July 1995
Abstract
Exotic softwood plantations were introduced into the Kibale Forest of western Uganda in the 1960s and early 1970s to
convert grassland areas into wood fiber producing sites. Following establishment, few silvicultural activities were initiated
within these plantings, and today harvesting operations are planned to remove these over-stocked stands to allow for the
development of natural forest communities. During the period May-September, 1994, we inventoried the overstorey and
advanced regeneration in plantations of Pi nus cari beae and Cupressus l usi t ani ca. Within 92 randomly located 0.01 ha plots
per plantation type, we measured the height and diameter of all stems encountered. The pines supported approximately 1.5
times the density and diversity of native woody stems found beneath the cypress (3424 tree and shrub stems ha- l under the
pines), while the pines themselves were characterized by stocking levels, mean heights, and canopy closures which were
higher than those of the cypress. We suggest that the greater natural regeneration development observed beneath the pines,
when compared with the cypress, is a function of pine development (influence upon competition) and stem characteristics
(influence upon seed dispersal mechanisms).
We then compared our regeneration observations with those reported for unlogged and logged areas of the surrounding
natural forest. Within the sapling class (diameter at breast height (dbh) less than 5 cm), which accounted for the majority of
the advanced regeneration under the softwoods, the plantations failed to support the same density of tree-species stems as
found in the natural forest (3077 stems ha-l vs. 2177 and 1111 under the pines and cypress, respectively). The number of
stems ha- ~ in the pines approximated those in 25-year-old logged-over areas of the natural forest. Species richness beneath
the conifers appears to approach the number found in the surrounding forest; however, a closer look at the species
composition suggest several deficiencies in the composition of the advanced regeneration beneath the plantations (especially
within the commercial class).
Following 25 years of development, the advanced regeneration beneath the plantations (especially the cypress stands) is
relatively impoverished compared to the levels of tree diversity and stocking characteristic of both unlogged and logged
natural forest sites. The conditions beneath the pines however, suggest that a diverse community of native species may
eventually exploit this environment through time. The dominance of the fast-growing, light-canopied, N2-fixing Al bi zi a and
Mi l l et i a genera, could help to nurture the further development of a well-structured, well-represented secondary forest
* Corresponding author.
Elsevier Science B.V.
SSDI 0039-6028(95)03637-7
216 R.4. Fimbel, C.C. Fi mbel / Forest Ecology and Management 81 (1996) 215-226
through time. Wi t h the application of silvicultural treatments, there may be management opportunities to further enri ch the
quality of the advanced regenerat i on beneath soft wood plantations. We expl ore future areas of research that coul d contribute
to this end.
Keywords: Cupressus lusitanica; Forest management; Pinus patula; Plantation; Regeneration; Restoration
1. I n t r o d u c t i o n
Re-establishing diverse and productive forests on
lands degraded by previous land conversion practices
such as agriculture, mining and logging, or areas
devastated by natural disasters, have often proved
difficult due to degraded soils and competition from
undesirable species arresting successional processes
(Lieth and Lohmann, 1993). Large energy inputs in
the form of fertility amendments, herbicide applica-
tions, weedings, and enrichment plantings have often
proved necessary to break this ' successional logjam' ,
encouraging the development of economical- a n d / o r
ecological-rich forests. Efforts to establish native
species onto these sites have often been abandoned
in favor of planting exotic conifers or fast-growing
N2-fixing trees, in consideration of their adaptability
to such depauperate conditions (Thompson et al.,
1986; Turnbull, 1987; Evans, 1992). In addition to
their capacity to conserve and enrich disturbed soils
(Yu and Pi, 1985; Jurgensen et al., 1986; Anderson,
1987; Lugo, 1992a), exotic plantations provide the
added benefit of producing timber, pulp, a n d / o r
fuelwood materials in time-frames considered rela-
tively short in comparison with many native tree
species.
In recent years, the potential for plantations to
serve as nurse crops for the establishment of native
forest species has been gaining recognition (Jordan
and Famworth, 1982; Hughes and Styles, 1987; Lugo,
1992b; Parrotta, 1992, 1993; Lugo et al., 1993).
While the silvicultural prescriptions required to max-
imize biomass production within many plantation
rotation systems have been extensively researched
(schedules for weeding, thinnings, harvest-regenera-
tion systems, etc.), to date, limited work has been
done to explore treatments designed to maximize the
native forest rehabilitation-restoration capabilities of
these plantings (Lugo, 1992a; Brown and Lugo,
1994).
Within the Kibale National Park in western
Uganda, exotic conifer plantations were established
on small grassland sites that dominate many of the
forest ridgetops, The goal of these plantings was
timber production. Today these plantations are ap-
proaching maturity, and beneath their canopies they
support dense and diverse stands of native saplings
and poles. During the summer of 1994 we invento-
ried this advanced regeneration, with the objectives
of: (a) evaluating the density and diversity of the
natural forest regeneration growing beneath 25-year-
old unmanaged plantations of Pinus caribeae and
Cupressus lusitanica; (b) making general compar-
isons between this advanced regeneration and data
reported by Kasenene (1987) and C. Chapman and
L. Chapman (unpublished data, 1994) for the sur-
rounding natural forest (unlogged and logged); (c)
assessing the role that these plantation species might
play as nurse crops for the rehabilitation-restoration
of a natural forest plant community. This work repre-
sents the first stage in a planned long-term study
designed to examine harvesting options, immediate
impacts of these practices on the advanced regenera-
tion, and post-harvest plant and animal dynamics.
2. Me t h o d s
2.1. Study area
The Kibale National Park (KNP) is 766 km 2 in
size, located just east of the Ruwenzori Mountains in
western Uganda between 013 ' and 041'N and
3019' -3032' E. The newly designated KNP encom-
passes the former Kibale Forest Reserve and a game
corridor to the south, serving as a bridge to the
Queen Elizabeth National Park. The Park ranges in
altitude from 1590 m in the north to 1110 m in the
south, receiving an average of 1470 mm of precipita-
tion annually, distributed over two distinct rainy
R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215-226 217
seasons. Two rivers, the Mpanga and the Dura, drain
the Kibale forest watershed into Lake George.
Kibale National Park is comprised primarily of
mature forest, interspersed with selectively logged
forest, colonizing forest, grasslands, and papyrus
swamps, with exotic softwood plantations distributed
among these natural habitats. The softwood planta-
tions were planted by the Ugandan Forest Depart-
ment in the late 1960s and early 1970s on lands in
the protected forest zone that were previously occu-
pied by ' unproduct i ve' grasslands. The history of
these grasslands is not well documented, but they are
believed to have been the result of long-term agricul-
tural use by local people (Kingston, 1967). Although
programs were originally designed for the mainte-
nance of these plantations, the civil and political
unrest in Uganda during the 1970s and much of the
1980s caused these maintenance programs to be
abandoned. Other than brushing for weed control
during the first 2 - 3 years following their establish-
ment, no other silvicultural practices were conducted
in these plantations. The current policy for the plan-
tations does not include a maintenance program, and
as such, regeneration from the surrounding ' nat ural '
forest is prevalent in the sub-canopy and understorey
levels.
2.2. Vegetation sampling
A total of 184 permanently marked sample plots
for measuring overstorey and understorey vegetation
were established in six softwood plantations (three
were Pinus caribeae and the other three were Cu-
pressus lusitanica). Each stand ranged from 3 to 5 ha
in size, and within species, exhibited similar biotic
and edaphic characteristics.
2.3. Overstorey plots
Circular 0.01 ha plots (5.64 m radius) were ran-
domly centered at 20- 50 m intervals within the
softwood plantations, giving a mean plot interval of
one plot per 1225 m z. These plots represent an 8%
sampling intensity of the study area, and no plot was
closer than 20 m to the stand edge or a road. Within
each plot the following plant characteristics were
noted for all stems over 1.3 m in height: species,
height, and dbh in centimeters. In addition, the per-
Table 1
Overstorey and understorey vegetation characteristics beneath exotic softwood plantations of Cupressus lusitanica (CL) and Pinus caribaea
(PC) in the Kibale National Park, Uganda, September 1994. The P-value shows the significance level of a one-way ANOVA test for
differences within and between stands of these two forest types. Values followed by different letters are significantly different at the
P ~< 0.05 level
Plant CL l CL2 CL3 PC 1 PC2 PC3 CL PC P-
parameter a (n = 37) (n = 24) (n = 31) (n = 37) (n = 24) (n = 31) mean mean value
(n = 92) (n = 92)
Conifers
Stems (ha- 1 ) 454 463 417 827 646 620 444a 710b 0.000
BA (m 2 ha- 1) 36.6 42.0 55.7 79,8 66.7 64.7 44.4a 71.3b 0.000
dbh (cm) 31.6 35 41.3 34.1 35.6 37.6 35.8 35.4 0.664
Cover (%) 79 82 78 86 83 86 80a 85b 0.000
Height (m) 24 22 24 35 37 28 23a 33b 0.000
LCR (%) 23 30 27 20 31 31 27 27
Regeneration
Stems (ha- i ) 1535 1545 2780 3089 4709 3019 1958a 3524b 0.000
BA (m 2 ha- i) 4.5 0.9 0.8 3.6 8.4 3.6 2.3a 4.8b 0.004
dbh (cm) 2.1 1.3 2.1 2.9 2,9 2.3 1.6a 2.7b 0.000
Height (m) 3.1 2.3 2.4 3.8 4.2 3.6 2.6 3.9 0.806
Species per plot 5.5 6.1 9.4 9.5 12.7 8.1 6.9a 9.9b 0.000
Trees spp. per plot 2.9 2.7 5.0 7.3 7.7 4.9 3.9a 6.6b 0.000
Shrub spp. per plot 2.3 3.0 3.8 2.8 4.2 3.2 3.0 3.3 0.340
Grd. cover (%) 61 NA NA 70 NA NA 61a 70b 0.018
a BA, basal area; LCR, live crown ratio; grd. cover, ground cover. NA, not available.
218 R~A. Fi mbe l , C. C. Fi mb e l / For e s t Ec ol ogy a n d Ma n a g e me n t 81 ( 1996) 2 1 5 - 2 2 6
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220 R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215 -226
cent live crown and canopy closure (at 1.3 m using a
spherical crown densiometer) were measured for the
plantation species. Physical characteristics of the site,
such as slope (%), aspect (), and site disturbances
were also recorded.
2.4. Understorey plots
Within each overstorey plot the ground vegetation
less than 1.3 m in height was quantified using four
1-m 2 (0.0001 ha) quadrats. The sub-plots were sys-
temically located inside the 0.01 ha plot, and all tree
and shrub seedlings stems falling in these quadrats
were counted. Within 74 of the overst orey plots (37
in the pine and 37 in the cypress), in addition to
recording the species and number of woody stems,
the percent ground cover contributed by all lifeforms
was noted (the quadrat was sub-divided into 25 4%
blocks to facilitate this latter activity). The under-
storey plots represented a sampling intensity of 0.32%
across the study area for all plant paramet ers except
cover (the latter was sampled at an intensity of
0.13 % within the plantations).
Plants were identified using a combi nat i on of the
following resources: botany keys, local knowledge,
and Ugandan botanists on site and at the Makerere
University herbarium. Voucher specimens of uncom-
mon or questionable plant species were deposited at
the Makerere University Biological Field Station in
KNP.
2.5. Soil sampling
Within four overstorey plots in each of the six
plantations (n = 12 for each conifer species), four
samples were systematically excavat ed to a depth of
20 cm using an Oakfield soil auger, sieved through a
2 mm screen, and combined. In the natural forest
adjacent to each softwood plantation, four randoml y
located sites were selected and the same soil extrac-
tion procedures repeated. Samples were oven dried at
600C until a constant weight, and then transported
to the soil testing facilities at Rutgers University
(New Brunswick, NJ) where the macro- and micro-
nutrients were analyzed using a Spectra Span 7 DCP
spectrometer (Applied Research Laboratories, Valen-
cia, CA).
2.6. Statistical analysis
All treatments were compared using a one-way
ANOVA to identify significant differences within
and between the conifer species. Tukey' s Studen-
tized Range Test (HSD) was used to identify signifi-
cant differences ( P < 0.05) between treatment means
when F values in the ANOVAs were significant
( a = 0.05). Assumpt i ons of the statistical models
were satisfied.
3. Results
3.1. Cypress vs. pi ne--overst orey
Pines averaged 710 stems ha -~, and a basal area
of 71.29 m 2 h a - t ; approxi mat el y 1.6 times the mean
values exhibited by the cypress (Table 1, P < 0.001).
Within each species there was variability among the
three replications (stands); however, the pines consis-
tently demonstrated higher values for the above vari-
ables across all stands examined. The pines were
also taller than the cypress ~ = 33 m vs. 23 m,
P < 0.001) and supported a slightly higher canopy
cover (-~= 85% vs. 80%, P < 0.001). The two species
had similar mean tree diameters ( 35- 36 cm dbh) and
live crown ratios (27%, Tabl e 1).
3.2. Cypress vs. pi ne--nat ural regeneration in un-
derstorey
There was greater natural regeneration develop-
ment beneath the pines compared with the cypress
plantations. We found an average of 3524 stems
ha - 1 under the pine, with a mean basal area of 4.78
m 2 ha -1. These means were roughly twice as high
as those for cypress (Table 1, P < 0.001). Per 0.01
ha plot (n = 92 per plantation type), the pines sup-
ported more native species ~ = 9.9 vs. 6.9 under
cypress, P < 0.001). Roughly 60% of these species
were trees ( ~= 6.6 under pine, 3.9 under cypress),
and both of the softwoods supported similar numbers
of shrub species per plot ~ = 3.3 and 3.0 respec-
tively, Table 1). Ground cover, consisting of woody
and herbaceous stems under 1.3 m tall, was higher
under the pines than the cypress ~ = 70.4% vs.
R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 8l (1996) 215-226 221
61.5%, P ~< 0.018). In no plots did we observe the
regeneration of the exotic softwood species.
The number of tree species developing under the
pines appeared higher (46 known, one unknown)
than under the cypress (40 known, two unknowns),
with a species overlap of approximately 70% be-
tween the two softwoods (Table 2). Those species
that did not overlap were relatively uncommon (the
maximum observed number was 11 stems per species
ha-1 within the non-overlapping group of species).
A total of 15 commercial species established beneath
the softwoods (11 in the pine and 13 in the cypress),
with Blighia wel wi t schi i / uni j ugat a (we were not
able to clearly differentiate between these two com-
mercial species, perhaps owing to hybridization be-
tween them), Celtis africana, Fagaropsis angolensis,
Funtumia africana, and Olea welwitschii accounting
for over 90% of the stems in this category. Commer-
cial species comprised roughly 15- 20% of the ad-
vanced regeneration stems. A total of 44 noncom-
mercial species was observed in the plantations: 36
in the pines and 29 under the cypress (Table 2). Of
this latter group of species, Albizzia grandibracteata,
Celtis durandi, Diospyros abyssinica, Milletia dura,
and Teclea nobilis, accounted for over 60% of the
stems under each softwood type. Finally, the number
of species and stem counts were highest within the
sapling class (85% in the pines and 96% in the
cypress), with few tree-size individuals observed.
3.3. Natural regeneration under softwood planta-
tions and natural f orest sites (unlogged and logged)
Sapling densities appear to be higher under the
natural forest than the softwood plantations and the
25-year-old logged natural forest sites (Table 2). In
the sapling class, stems under the pines (2168 ha- t )
were 70% as numerous as those in the natural forest
(3077 stems ha- t ), while the cypress contained only
36% (1111 stems ha - t ) as many as the natural
forest. The real disparity between these densities is
even greater however, as plantation counts represent
stems between 0.5 and 5 cm dbh, while the mini-
mum that Kasenene (1987) recorded was 2 cm.
When considering only saplings between 2 and 5 cm
dbh within the softwood stands, the pines support
only 43% of the number of saplings found in the
natural forest, and the figure for cypress drops to
11%. By comparison, the heavily logged K15 com-
partment supported 25% as many stems as the natu-
ral forest, and the lightly cut K 14 exhibited 49% the
number of saplings. In addition, the logged forests
appear to support higher numbers of commercial
species (17 and 19 respectively), and a higher per-
cent of their stems in the commercial category (65%
for K15 and 59% for K14) compared with the pines
(11 commercial species accounting for 12% of all
stems) and the cypress (13 species accounting for
22% of all sapling stems). The undisturbed natural
forest contained 18 commercial species in the sapling
class, accounting for slightly over one-third of the
stems observed (Table 2). Several species which
were either absent or found in very low densities
within the plantations compared to the natural forest
sites, included Chrysphyllum, Lovoa, Mimusops,
Parinari, Strombosia, Bosquiea, Leptoncychia, and
Uvariopsis (the former five are of a commercial
grade, the latter three considered as noncommercial).
In general, the logged forests appear to be closer in
sapling composition to the natural forest conditions,
when compared to the regeneration developing under
the plantations.
Within the pole class ( 5- 13 cm dbh), the pines
contained 21% of the number of stems ha - I of the
natural forest (1316 ha - I ) , and the cypress a mere
1.4% (Table 2). The heavily logged forest (K15)
supported 18% of the number of poles in the natural
forest, while the lightly harvested K14 exhibited
32% as many. These latter two sites supported 14
and 16 commercial tree species respectively, which
is similar to the 16 observed by Kasenene in the
unlogged natural forest, while the pines supported
four of these species and the cypress one. The num-
ber of commercial stems in K15 represented 58% of
the number in the unlogged natural forest, while the
density in KI 4 was 1.4 times that of the natural
forest. The density of commercial poles in the pine
plantation was 14% of that found in the natural
forest, whereas the figure for cypress was only 0.5%.
Very few tree size stems (13 cm dbh and larger)
were found developing in the plantations. These
areas exhibited less than 7% of the natural forest
stocking levels observed by Kasenene (1987), and
less than 10% of the stem count recorded by C.
Chapman and L. Chapman (unpublished data, 1994).
222 R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215-226
Table 3
Soil properties within the top 15 cm of soil under exotic soft wood
plantations of Cupressus lusitanica (CL) and Pinus caribaea
(PC), and natural forest (NF) in the Kibale National Park, Uganda,
Sept ember 1994. The P- val ue shows the si gni fi cance level of a
one-way ANOVA test for di fferences in soil property concent ra-
tions among t he three forest types. Val ues followed by different
letters are si gni fi cant l y di fferent at t he P ~< 0.05 level
Soil property CL PC NF P- val ue
( n = 12) ( n = 12) ( n = 12)
pH 5.6 5.6 6.1 0. 086
Mg (kg h a - l ) 949a 660b 714ab 0.037
P (kg ha -1 ) 267 311 196 0.216
K (kg ha ~) 466 266 336 0. 252
Ca (kg h a - ~ ) 5071 3434 5556 0. 052
Cu ( ppm) 7. 9a 4. I b 5.8b 0. 000
Mn ( ppm) 95. 6a 54.9a 199.7b 0. 000
Zn ( ppm) 8. 7a 4. 4b 7.8ab 0.035
B (ppm) 0. 6a 0. 4a 1.2b 0.003
The number of commercial trees within the softwood
groups was negligible (Table 2).
3.4. Soil properties of the softwood plantations and
adjacent natural forest sites
The concentrations of Mg, Cu, and Zn were lower
in the pine plantation soils than in the cypress (Table
3, all P ~< 0.037), with all other elements tested,
except P, showing a similar trend. Compared with
the natural forest, the plantations contained lower
concentrations of Mn and B ( P ~< 0.003), and the pH
of the soil may be undergoing some acidification
~ = 5.6 under the softwoods vs. 6.1 in the natural
forest) as a result of the coniferous species (Table 3,
P = 0.086).
4. Di scussi on
4.1. Natural regeneration development under the
softwood plantations --pines vs. cypress
The greater natural regeneration development ob-
served beneath the pines, compared with the cypress
stands, appears to have little association with the
physical location a n d / o r soils characterizing these
plantations. All plantations occupied similar topo-
graphic sites, and no portion of the stands sampled
was more than 150 m from the natural forest. All
other factors constant (an unknown postulation), seed
recruitment should have been equal across all areas
investigated. Nor did our cursory soil appraisal sug-
gest that the chemical properties in the soils were
responsible for the higher advanced regeneration ob-
served under the pines. With the exception of P (P
appeared slightly higher in the cypress compared to
the pine soils, yet the difference was not significant),
all other elements examined showed a trend towards
lower concentrations under the pines. No elements
appeared to fall within a range that one might con-
sider limiting to plant development (Sanchez, 1976;
Proctor, 1989).
The higher levels of advanced regeneration below
the pines, as evidenced by larger stems and greater
numbers of individuals and species per unit area,
appear to be linked to the growth and development
of this conifer. Under the site conditions at KNP, P.
caribaea grows faster, taller, and accumulates more
biomass per unit area than C. lusitanica (Ugandan
Forest Department, unpublished data). These growth
characteristics, coupled with the high stocking rates
used at the time of planting (2.7 m centers), suggest
that at a young age the pines would have rapidly
closed their canopy and shaded out the herbaceous
competitors that formerly dominated the grassland
planting sites. Hence a seedbed free of competitors
was probably created, yet sunlight sufficient to en-
courage the germination and establishment of many
native stems was still lacking in pole-size stands.
Through time, as this low shade-tolerant species
grew in height, its high stocking levels lead to the
shedding of lower branches and the development of
long bole lengths absent of live branches. This high
canopy condition, coupled with: (a) a low mean live
crown ratio; (b) the open crown structure characteris-
tic of P. caribaea (few branches and twigs in com-
parison with cypress, and a much less dense foliage
pattern); (c) limited competition from herbaceous
species, appears to have created the light regime and
growing space requirements conducive to the estab-
lishment of native woody species. Today we observe
the development of advanced regeneration, and the
continued recruitment of new individuals and species,
under the declining shade of the pine overstorey (the
latter process due to increasing stem mortality).
R~4. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215-226 223
In contrast to the above scenario, cypress devel-
opment during the past 25 years appears to have
been less suited as a nurse crop to the establishment
of native species. Cypress seedlings develop slower
than their pine counterparts, and during the early
stages of stand development herbaceous competitors
may have suppressed stem growth. This condition
could have lead to high seedling mortality, thereby
slowing full canopy closure by the conifers until late
in the rotation, which in turn allowed herbaceous
competitors to dominate the site much longer than
under the pines. Possibly further retarding the estab-
lishment of native species was the low height and
very dense crown characteristics of cypress, which
serve to greatly limit both the direct and indirect
light reaching the forest floor. Dense shade still
persists across many areas of the cypress stands
today (the mean canopy cover values presented in
Table 1 are somewhat misleading, as the advanced
regeneration contributes significantly to our readings
of the shade beneath the pines, while in the cypress
stands almost all of the canopy closure is due to this
conifer species), with most of the advanced regenera-
tion limited to areas in and adjacent to sites of
cypress mortality. The latter areas are generally small,
often less than 30 m across, and are usually a
function of windthrow or elephant damage. Thus,
extended periods of herbaceous competition fol-
lowed by dense shading under the cypress, appears
to have limited the germination and establishment of
native tree species. These conditions contrast greatly
from those observed under stands of P. caribaea.
The nurse-crop characteristics discussed above
may only explain part of the reason that we observed
greater levels of advanced regeneration under the
pines. Seed dispersal mechanisms may have con-
tributed to the higher density and diversity of species
observed beneath the pines. Pine plantations in KNP
are utilized by a variety of wildlife species (Keith et
al., unpublished data), including elephants (Loxo-
donta africana), red colobus ( Colobus badius), red-
tail monkeys (Cercopithecus ascanius), chimpanzees
( Pan troglodytes), duiker (Cephalophus harveyi and
C. moniticola), bushbucks (Tragelaphus scriptus),
civet cats (Viverra civetta and Nandinia binotata),
and genets (Genetta sp). Many of these mammals, in
conjunction with birds, are probably responsible for
dispersing most of the seeds into these plantation
sites (C. Chapman and L. Chapman, unpublished
data, 1994); a condition common in many tropical
forests (Estrada and Fleming, 1986). Furthermore,
with the exception of elephant sign (bark stripping
on the cypress), almost all of our animal sightings
and sign observations were limited to the pines. The
large and tow density branching pattern of this species
appears to facilitate usage by primates, while the
denser and more diverse understorey growth helps to
attract terrestrial fauna, perhaps explaining our
greater number of encounters with these animals in
the pines compared to cypress stands.
4.2. Regeneration development under the softwood
plantations compared with unlogged and logged ar-
eas o f the natural f orest
Twent y-fi ve years following their establishment,
unmanaged plantations of P. caribaea and C. lusi-
tanica do not support native species in the same
density as found in the surrounding natural forest
(levels in the pines approximate those in logged-over
areas of this forest-type). The level of species rich-
ness developing beneath these conifers however, ap-
pears to approach the number recorded by Kasenene
(1987) and C. Chapman and L. Chapman (unpub-
lished data, 1994) for areas of surrounding natural
forest. Do the plantations, and in particular pine,
given its more advanced regeneration development
in comparison with cypress, offer managers a tool to
rapidly rehabilitate (and possibly restore) degraded
areas of natural forest?
A closer look at the species composition suggests
several deficiencies in the composition of the ad-
vanced regeneration beneath the plantations. To be-
gin, while several commercial species are well repre-
sented under the softwoods, several others are no-
tably missing or rare (Chrysophyllum, Lovoa,
Mimusops, Newtonia, Parinari, and Strombosia).
This condition is exasperated as one moves upwards
from the sapling class into the pole- and tree-size
stem classes. Additionally, in a mature state, these
same species are important producers of fruits con-
sumed by native wildlife (personal observation). The
absence of these species within the developing stands
suggests that the future overstorey could be impover-
ished in its commercial and ecological values.
224 R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215-226
A direct comparison between the noncommercial
species mix under the plantations and the natural
forest sites is difficult, as Kasenene (1987) only lists
the total number of species he observed and the four
most common species within this category. Hence,
we turned to C. Chapman and L. Chapman' s listing
of noncommercial tree species (unpublished data,
1994), which indicated a slightly richer diversity in
the natural forest (their sample area was 2.6 times
larger than the area we sampled, which might help to
explain their higher number of species). Of greater
interest however, is the low percentage of overlap
between their list of natural forest trees and ours
derived from all size-classes within the plantations
(roughly one-third overlap between them). This raises
two possibilities: (1) that the species mix below the
plantations is impoverished in species capable of
growing into the tree size-class (shrubs and small
trees that seldom reach 10 cm dbh dominate); or (2)
that the species below the plantations simply com-
prise a small proportion of the total species composi-
tion comprising the highly heterogeneous natural
forest in KNP (J.M. Kasenene, personal communica-
tion), and therefore the plantation species we ob-
served are representative of a natural forest under-
storey community. Lacking Kasenene' s (1987) data
to clarify this discrepancy, we believe that both
possibilities contribute to the plantation response.
Approximately 50% of those species found under the
plantations but not recorded by C. Chapman and L.
Chapman (unpublished data, 1994) are capable of
exceeding 10 cm dbh, supporting both of the above
points.
Following 25 years of development, the advanced
regeneration beneath the plantations (especially the
cypress stands) is relatively impoverished compared
with the levels of tree diversity and stocking charac-
teristic of both unlogged and logged natural forest
sites. This state reflects the influence of time
(succession is young under the plantations compared
with the natural forest), differences in seed dispersal,
site conditions (plantations in general occupied ups-
lope sites while the logged and uniogged sites cover
a wider topographic gradient), etc. The conditions
beneath the pines however, suggest that a diverse
community of native species may eventually exploit
this environment through time. The dominance of the
fast-growing, light-canopied, N2-fixing Albizzia and
Milletia genera, could help to nurture the further
development of a well-structured, well-represented
secondary forest through time (Mitra and Sheldon,
1993). With the application of silvicultural treat-
ments, there may be management opportunities to
further enrich the quality of the advanced regenera-
tion beneath softwood plantations.
4.3. Silvicultural treatments to encourage the devel-
opment of native pl ant species beneath exotic soft-
wood pl ant at i ons--areas requiring additional re-
search
Resource managers faced with the challenge of
encouraging the regeneration of diverse tropical
hardwood stands, be it following timber harvesting
operations or when attempting to rehabilitate de-
graded forest lands, may find plantations of exotic
conifers a useful tool to foster the establishment and
development of native tree species. Unmanaged 25-
year-old stands of pine in the KNP appeared to
support the early stages of a diverse native forest. A
variety of mechanisms appeared to have interacted to
create these conditions (suppression of fire and
herbaceous competition, facilitation of seed dis-
persers across the sites, etc.). However, the current
species composition and the negative impacts of
unsupervised clear-felling operations on this ad-
vanced regeneration (personal observation) suggest
that without silvicultural practices to encourage the
establishment and release of these woody species,
that the successional process could lead to an ecolog-
ically and economically impoverished forest (at least
within the short-run). What treatments might be
available to managers to expedite the development
and quality of this process?
We suggest that research is needed in several
areas of plantation management to maximize the
native forest rehabilitation-restoration potential as-
sociated with this nurse-crop practice. For example,
longterm studies on ways to reclaim grassland sites
to woodlands, or selectively logged sites to their
preharvesting conditions, are needed to compare nat-
ural forest regeneration development on areas en-
riched with conifer plantings versus those treated
with traditional silvicultural practices (no inputs,
plantings, weed control, etc.). An economic evalua-
tion of this process would be useful to assess whether
R, 4. Fimbel, C.C. Fi mbel / Forest Ecol ogy and Management 81 (1996) 215- 226 225
t he f i nal cash val ue of t he s of t wood st ems, coupl ed
wi t h t he benef i t of havi ng es t abl i s hed a qual i t y har d-
wood st and, woul d j us t i f y t he cost s as s oci at ed wi t h
r es t or i ng de gr a de d f or est l ands. Compl e me nt a r y st ud-
i es mer i t i ng i nves t i gat i on mi ght i ncl ude oppor t uni -
t i es f or si l vi cul t ur al pr act i ces t o encour age t he est ab-
l i s hment and de ve l opme nt of nat i ve speci es dur i ng
t he pl ant at i on r ot at i on, i ncl udi ng the eval uat i on of
t he r ol es of: ( a) t hi nni ng oper at i ons; (b) enr i chment
pl ant i ngs; (c) r esi dual ha r dwood st ems as bot h per ch
si t es and seed sour ces ( Cr uz, 1988); (d) wi l dl i f e
s peci es as seed di s per s er s and pl ant pr edat or s; (e)
har ves t i ng schedul es and t echni ques t o mi ni mi ze
da ma ge t o t he advanced r egener at i on, et c. In addi -
t i on, wor k needs t o be done t o subst ant i at e or di spel
f i ndi ngs such as t hose of St r uhsaker et al. (1989),
t hat exot i c coni f er pl ant at i ons ma y negat i vel y i mpact
adj acent nat i ve f or est speci es ( our st udy f ound l i t t l e
e vi de nc e t o s ugges t t hat t hese s of t wood speci es re-
t ar d t he de ve l opme nt of nat i ve speci es, howe ve r a
def i ni t i ve r es pons e t o t he poi nt s r ai sed by St r uhsaker
et al. wer e b e y o n d t he scope of t hi s st udy). The
l evel s o f i nves t i gat i on are near l y l i mi t l ess, and wi l l
var y wi t h o n e ' s obj ect i ves, be i t t ot al communi t y
r es t or at i on or s i mpl y t he encour agement of sec-
ondar y woody pl ant successi on.
We woul d l i ke t o cl os e by encour agi ng ot her s t o
r eas s es s t he t r adi t i onal r ol es of coni f er pl ant at i ons as
sour ces of f i ber a n d / o r t ool s t o cur b soi l er osi on,
and t o hel p eval uat e t he br oader pot ent i al o f t hese
pl ant i ngs t o pr ovi de e c onomi c r et ur ns whi l e hel pi ng
t o r ehabi l i t at e de gr a de d f or est communi t i es . Ma ybe
we r eal l y ' c a n have our cake and eat i t t oo' .
Acknowledgments
We wi sh to t hank Dr s. John Kas enene and Gi l ber t
I s a bi r ye - Ba s ut a of t he Maker er e Uni ver s i t y Bi ol ogi -
cal Fi e l d St at i on f or t hei r many const r uct i ve com-
ment s on t he exper i ment al des i gn and hel p wi t h
pl ant i dent i f i cat i ons. A ver y speci al t hanks is due t o
Dr s. Col i n and Laur en Cha pma n of the Uni ver s i t y of
Fl or i da f or t hei r many usef ul c omme nt s on al l phas es
of t he st udy i ncl udi ng t hi s manuscr i pt . We al so wi sh
t o r ecogni ze Ol i vi a Wa n y a n a Ma ga nyi and Dr. An-
t hony Kat ende o f t he Maker er e Uni ver s i t y her bar i um
f or t hei r as s i s t ance wi t h t he pl ant i dent i f i cat i ons, and
Pat r i ck our f i el d assi st ant for hi s many l ong hour s of
f i el d wor k. An a nonymous r eader al so pr ovi de d many
usef ul c omme nt s t o t he p a p e r ' s cont ent . Fi nal l y,
f undi ng f or t hi s pr oj ect was pr ovi de d by t he Wi l dl i f e
Cons er vat i on Soci et y ( WCS) and t he Uni t ed St at es
Ag e n c y f or I nt er nat i onal De ve l opme nt ( USAI D) .
References
Anderson, M., 1987. The effects of forest plantations on some
lowland soils. I. A second sampling of nutrient stock. Forestry,
60: 69-86.
Brown, S. and Lugo, A.E., 1994. Rehabilitation of tropical lands:
a key to sustaining development. Restort. Ecol., 2: 97-111.
Cruz, A., 1988. Avian resource use in a Caribbean pine plantation.
J. Wildl. Manage., 52: 274-279.
Estrada, A. and Fleming, T.H., 1986. Frugivores and seed disper-
sal. Dr. W. Junk, Boston, MA.
Evans, J., 1992. Plantation Forestry in the Tropics, 2nd edn.
Clarendon Press, Oxford.
Hughes, C.E. and Styles, B.T., 1987. The benefits and potential
risks of woody legume introductions. Int. Tree Crops J . , 4:
209-247.
Jordan, C.F. and Farnworth, E.G., 1982. Natural vs. plantation
forests: a case study of land reclamation strategies for the
humid tropics. Environ. Manage., 6: 485-492.
Jurgensen, M.F., Frederick, D.J., Magwick, H.A.I. and Oliver,
G.R., 1986. Soil development under Pi nus radiata and Euca-
lyptus regnans plantations. N. Z. J. For. Sci., 16: 69-77.
Kasenene, J.M., 1987. The influence of mechanized selective
logging, felling intensity and gap-size on the regeneration of a
tropical moist forest in the Kibale Forest Reserve. Ph.D.
Thesis, Michigan State University, Ann Arbor, USA.
Kingston, B., 1967. Working plan for Kibale and Itwara Central
Forest Reserves. Forestry Department, Entebbe, Uganda.
Lieth, H. and Lohmann, M., 1993. Restoration of Tropical Forest
Ecosystems. Kluwer, London.
Lugo, A.E., 1992a. Tree plantations for rehabilitating damaged
forest lands in the tropics. In: M.K. Wali (Editor), Environ-
mental Rehabilitation, Vol. 2. SPB, The Hague, Netherlands,
pp. 247-255.
Lugo, A.E., 1992b. Comparison of tropical tree plantations with
secondary forests of similar age. Ecol. Monogr., 62: 1-41.
Lugo, A.E., Parrotta, J. and Brown, S., 1993. Loss in species
caused by tropical deforestation and their recovery through
management. Ambio, 22: 106-109.
Mitra, S.S. and Sheldon, F.H., 1993. Use of an exotic tree
plantation by Bomean lowland forest birds. Auk, 110: 529-
537.
Parrotta, J. A. , 1992. The role of plantation forests in rehabilitating
degraded tropical ecosystems. Agric. Ecosys. Environ., 41:
115-133.
Parrotta, J.A., 1993. Secondary forest regeneration on degraded
tropical lands. The role of plantations as 'foster ecosystems'.
226 R.A. Fimbel, C.C. Fimbel / Forest Ecology and Management 81 (1996) 215-226
In: H. Lieth and M. Lohmann (Editors), Restoration of Tropi-
cal Forest Ecosyst ems. Kluwer, Netherlands, pp. 63-73.
Proctor, J. (Editor), 1989. Mineral Nutrients in Tropical Forest
and Savanna Ecosyst ems. Blackwell, London.
Sanchez, P.A., 1976. Properties and Management of Soils in the
Tropics. John Wiley, New York.
Struhsaker, T.T., Kasenene, J.M., Gaither, J.C., Jr., Larsen, N.,
Musango, S. and Bancroft, R., 1989. Tree mortality in the
Kibale Forest, Uganda: a case study of dieback in a tropical
rain forest adjacent to exotic coni fer plantations. For. Ecol.
Manage., 29: 165-185.
Thompson, D.A., Wright, D.L. and Evelyn, O., 1986. Forest
resources in Jamaica. In: D.A. Thompson, P.K. Bretting and
M. Humphreys (Editors), Forest of Jamaica. The Jamaican
Society of Scientists and Technologists, Kingston, Jamaica,
pp. 81- 90.
Turnbull, J.W. (Editor), 1987. Australian acacias in developing
countries. Proc. No. 16, Australian Centre for International
Research, Canberra, Australia.
Yu Zou Yue and Pi Yong Feng, 1985. The path and its effects to
rehabilitating vegetation on tropical littoral eroded land in
Guangdong Province. Trop. Subtrop. Ecosys., 3: 97-108.

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