Forest Ecology and Management 111 (1998) 127±135

Provenance and family variation in Sterculia apetala in Colombia

W.S. Dvoraka,*, H. UruenÄab, L.A. Morenob, J. Gofortha
a
Director and Forest Technician, respectively, CAMCORE Cooperative, College of Forest Resources, North Carolina State University,
Raleigh, NC 27695, USA
b
Forest Director and Research Forester, respectively, Pizano/Monterrey S.A., Apartado Aereo 6610, Cartagena, Colombia

Received 24 March 1998; accepted 27 April 1998

Abstract

Three provenances and 23 open-pollinated families of Sterculia apetala (Jacq.) Karst were established in a genetic ®eld test in
Zambrano, (Bolivar) northern Colombia (108 N lat.), where average annual rainfall is 920 mm. The provenances were
Tiquisate, Guatemala, Cofradia, Honduras, and northern (Atlantico and Bolivar), Colombia. Growth and quality traits were
measured over an 8-year period, and leaf initiation and wood speci®c gravity were assessed at age 10. The average height of S.
apetala was 7.6 m at 8 years of age. The local Colombian provenance had 11% better survival, produced 28% more volume,
and had better stem straightness than the best Central American provenance. Leaf initiation and leaf fall occurred in the
introduced Central American sources earlier than in the local Colombian provenance and their timing appeared partly out of
phase with the onset of the wet and dry seasons. Wood speci®c gravity averaged 0.208 at 10 years of age. The local Colombian
source had signi®cantly higher speci®c gravity than the introduced Central American sources, 0.216 vs. 0.204, respectively.
There was a weak positive phenotypic correlation (rˆ0.21) between height growth and wood speci®c gravity. Individual tree
heritability for growth traits and quality traits ranged between 0.10 and 0.23. Individual tree heritability for leaf initiation was
0.05 and for speci®c gravity 0.44. Conservation and breeding efforts for S. apetala will continue to be minimal until new
markets develop to utilize wood with low density. # 1998 Elsevier Science B.V.

Keywords: Dry zone; Provenance variation; Wood speci®c gravity

1. Introduction over 35 common names throughout its range

Sterculia apetala (Jacq.) Karst is found throughout
parts of southeastern Mexico, tropical Central and
South America (as far south as northern Bolivia),
and several islands in the Caribbean and the West
Indies (Timyan per. comm., Rojas and Sibelle, 1996;
TrivinÄo et al., 1990; Perez-ArbelaÂez, 1978; McCallan,
1943; Martorell, 1940). The species has been given
*Corresponding author.
(Table 1). Even though its natural distribution is wide-
spread, S. apetala is seldom the dominant tree species
in any one area. It occurs in both deciduous and semi-
deciduous dry zone forests where annual rainfall can
be as low as 920 mm with pronounced dry seasons of
4±6 months. It also occurs in riparian evergreen forests
in areas where precipitation can be well over 3000 mm
per year (TrivinÄo et al., 1990; Janzen, 1971). It is most
often found on sites below 400 m altitude and prefers
well-drained alluvial soils such as sandy loams. It can

0378-1127/98/$19.00 # 1998 Elsevier Science B.V. All rights reserved.

PII S0378-1127(98)00316-8
128 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135

Table 1
Common names used for Sterculia apetala in various countries

Country Common name(s) Source

Brazil chichaÂ, exixa, xixa Perez-ArbelaÂez, 1978

Regina Martins da Silva, 1998
Colombia algodoÂn, bobena, camajoÂn, TrivinÄo et al., 1990
camajoruÂ, camajondura, camajuruÂ, Escobar and RodrõÂguez, 1994
cau, gomo blanco, majao, pinÄon, Perez-ArbelaÂez, 1978
suaÂn tacazeiro, tun-tun, zapote,
zapote de monte, zapato, zapoterama
zapotillo
Costa Rica panama Janzen, 1971
Cuba camaguÈey, guana Escobar and RodrõÂguez, 1994
Ecuador sapote de montanÄa, Perez-ArbelaÂez, 1978
coco de monte Rojas and Sibelle, 1996
Guatemala castanÄo, mano de leoÂn Elmer GutierreÂz, 1997 (per. comm.)
Guyana (Brit) yahu Escobar and RodrõÂguez, 1994
Guyana (Fr) kola Escobar and RodrõÂguez, 1994
Haiti pistache des indes Joel Timyan, 1997 (per. comm.)
Honduras castanÄo, karst
Jamaica bastard mahogani Escobar and RodrõÂguez, 1994
Mexico bellota
Panama panamaÂ
Peru huayra caspi Rojas and Sibelle, 1996
Puerto Rico panamaÂ, anacaguitas, anacahuita Martorell, 1940
Surinam bofrohoedoe, kobehe, kroekroeamete Escobar and RodrõÂguez, 1994
Trinidad mahoe Escobar and RodrõÂguez, 1994
Venezuela cacaguÈillo, cacaguÈito, cacaõÂto Rojas and Sibelle, 1996
cacauito, cameruco, camojoco, Lamprecht, 1955
camoroco, camoruco, cumaruco, Escobar and RodrõÂguez, 1994
pato de danto, sun-suÂn,
yagrumo sun-suÂn, vara de indio,
zapato

not tolerate soils that are constantly saturated with
water.
Through most of its range, Sterculia apetala is
usually a medium-sized tree that grows to 24±30 m
height. However, the tree has been reported to reach
50 m height and 2 m diameter on fertile soils in old-
growth evergreen forests in west central Venezuela
where rainfall averages 1900 mm per year (Lam-
precht, 1955). Mature S. apetala has a smooth green-
ish-white bark, a wide spreading crown when grown
free, and a ¯uted stem near the base of the tree. When
cut, mature trees do not produce coppice shoots
(Lamprecht, 1955). However, trees <5 years of age
have been observed to produce basal sprouts after
being felled.
Sterculia apetala is monecious. The seeds of S.
apetala trees ripen during the dry season (February
through March) in most locations in tropical America.
In its native environment in evergreen forests, S.
apetala trees produce their ®rst seed crops when trees
are 20±30-years old or 15±25 m tall (Janzen, 1971).
It appears to take 1 year from the onset of open
pollination to the time that the seed is ready to harvest
(Janzen, 1971, GutieÂrrez per. comm.). The quantity of
seeds produced may vary greatly from year to year.
The seeds of S. apetala have high starch content
relative to many other tropical broadleaf species in
Middle America (TrivinÄo et al., 1990).
Seed eating insects attack the pods as the seeds are
about to mature and also after the pods fall to the
ground, destroying most of the crop (Janzen, 1971;
Derr, 1980). Seeds have also been known to be eaten
by deer, squirrels, parrots, monkeys and other animals
(Janzen, 1971). In several locations in tropical Amer-
 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135
ica and the Caribbean, S. apetala seeds are collected
by farmers and either eaten raw or roasted (Herrera-
Isla, 1993; Perez-ArbelaÂez, 1978). In Cuba, the ¯ow-
ers are sometimes cooked and used as a decongestant
(Herrera-Isla, 1993).
Sterculia apetala is only planted occasionally in
tropical America, usually in gardens as ornamentals or
along the sides of roads (Herrera-Isla, 1993; Martorell,
1940). Farmers in Guatemala and Honduras some-
times leave large S. apetala trees in pastures as shade
for cattle. However, farmers in Costa Rica actively
remove trees when these are near to cotton ®elds
because of the cotton stainer (Dysdercus spp. ) insect
problems associated with the trees and the resulting
damage that the insects can in¯ict on the crops (Jan-
zen, 1971). However, the primary reason S. apetala
has not generated much local interest as a species for
afforestation or reforestation is that its yellow-brown
wood is generally described as spongy and soft, and is
not very durable. When planed, it produces a fuzzy
grain. It has been utilized locally for bed boards,
interior molding, ceiling boards, wood boxes, and
canoes. Wood from mature trees has also been used
for doors, window frames, door frames, forms for
cement walls, crude furniture, plywood and particle
board. Generally, however, the wood is mostly used
for indoor purposes (molding) where weathering is not
a concern. There is also interest to use it as cores in
plywood to produce light weight boards.
Sterculia apetala is not in danger of extinction
because of its widespread distribution. However, many
small populations throughout its range are being
Table 2
Location information for the provenance collection sites of Sterculia apetala in Guatemala, Honduras, and Colombia, and the test
establishment site at Zambrano, Colombia
Provenance Department Country
Tiquisate Escuintla Guatemala
Cofradia Cortes Honduras
Northerna Colombia Atlantico/Colombia Colombia
Zambrano Bolivar Colombia
a
Seed collections were made across several locations in the departments of Atlantico and Bolivar. Latitude, longitude, altitude and
precipitation information presented represent an average value for the departments where S. apetala occurs.
129
destroyed, usually to plant agricultural crops or clear
land for pastures. In efforts to better determine the
potential of S. apetala as a plantation species and
encourage its use locally, the Central America and
Mexico Coniferous Resources Cooperative (CAM-
CORE), North Carolina State University, the National
Forest Seed Bank, (BANSEFOR), Guatemala, and the
National School of Forest Science (ESNACIFOR),
Honduras, in collaboration with Pizano/Monterrey
S.A., a forest company in Colombia, established a
genetic ®eld test of the species in Colombia (Dvorak
and Donahue, 1992). Provenance and family variation
in growth and quality traits were studied over an 8-
year period and wood speci®c gravity was assessed at
age 10. This paper reports on the results from this
study, and makes recommendations about the breed-
ing and conservation strategies that should be imple-
mented. The potential of S. apetala as a plantation
species in the tropics is also discussed.
2. Materials and methods
Seeds of Sterculia apetala were collected by CAM-
CORE and its honorary members BANSEFOR and
ESNACIFOR in Guatemala and Honduras, respec-
tively, in March, 1986. The collection in Guatemala
was near Tiquisate, in the southwestern part of the
country (Table 2). The area had once been a semi-
deciduous forest but most of the high value trees in the
area were cut long ago for the development of agri-
culture. A relatively large stand of S. apetala occurs in
Latitude/Longitude Altitude Annual precipitation
(m) (mm)
148 170 N 50±70 2094
918 230 W
158 240 N 50±75 1370
888 020 W
108 380 N 20±70 1000±
748 550 W 1400
98 450 N 50±70 920
748 490 W
130 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135
the area with a density of approximately three trees per
km2 and 19 mother trees were sampled. In Honduras,
seed collections were made at Valle de Sula (Cofradia)
in a semi-deciduous forest. The Cofradia forest was
comprised of small stands of widely scattered trees.
Much of the forest area in the region had been con-
verted to pasture land. Seeds were collected from 20
trees. During the same period, Pizano/Monterrey
Forestal S.A. made a collection of 11 S. apetala trees
across a 200 km transect in the departments of Atlan-
tico and Bolivar in the dry-zone deciduous forests of
northern Colombia. The terrain of the collection area
ranged from ¯at to rolling hills. The annual precipita-
tion in Atlantico is 1400 mm but falls to <1000 mm
in Bolivar (Table 2). The stands sampled were com-
prised of small patches of relatively intact forests of S.
apetala with 3±5 trees per km2. Three of the 11 trees
selected were in Atlantico and the rest were in Bolivar.
Because of the small sample size and the fact that the
trees were chosen over a large area, no attempt was
made to separate the collection into an `Atlantico' and
`Bolivar' provenance. Instead, the 11 tree collections
from Atlantico and Bolivar were grouped together
and called the `Northern Colombian' provenance in
this paper. The provenances selected for the study
represented moderate-to-low rainfall areas in semi-
deciduous and deciduous forest where Sterculia
apetala occurs naturally (Table 2).
The seeds from the 38 open-pollinated families
collected in Guatemala and Honduras were sent
directly to Colombia and were germinated with the
11 Colombian S. apetala families. However, only 10
of the Guatemalan families and two of the Honduran
families produced enough seedlings to be included in
the provenance/progeny trial which was established
near Zambrano, (Bolivar) in the northern part of the
country. The test site was located on a small ridge
above the Magdalena River in an area with 920 mm
of annual precipitation. This represents the extreme
lowest rainfall limit where S. apetala naturally occurs
(Table 2). In years of drought, annual precipitation
may be only 600 mm. The soil at the test site was a
well-drained sandy clay loam derived from ¯uvial
sediments and is classi®ed as an aridic haplustalf.
The A (top) horizon was 34 cm in depth with soil
pH 7.5, the B horizon was 63 cm in depth with soil pH
7.7. A plow pan was present in the soil at 20 cm depth
that limited root growth and development especially
during the dry season. Sorghum (Sorghum vulgare)
was previously planted on the site followed by Car-
ibbean pine (Pinus caribaea var. hondurensis) which
failed to survive. Most likely, this area along the
Magdalena River had been under cultivation periodi-
cally for hundreds of years because of the high fertility
of alluvial soils for growing crops and the importance
of the river as the primary mode of transportation for
indigenous people in the region.
The 23 open-pollinated families (10 from Guate-
mala, 2 from Honduras and 11 from Colombia) were
planted in a compact family design, with 9 replications
and 6 tree family row plots. In this test design, families
are grouped together by provenance, and their position
within the block is randomized from one replication to
the other (Table 2). The 1242 trees were outplanted in
April, 1987 when seedlings were 40±50 cm tall. The
trial spacing was 33 m.
The test was measured at 1, 3, 5, and 8 years for a
number of adaptability, growth and quality traits.
Speci®cally, survival was assessed at all four measure-
ment periods. Height and diameter at breast height
(DBH) was measured at ages 3, 5, and 8 years.
Individual tree volume (outside bark) was calculated
at ages 5 and 8 years using a general formula for
juvenile trees derived by Ladrach and Mazuera (1978)
and had the form:
Volume ˆ 0:00003 height  DBH2 (1)
where height was recorded in m and DBH is measured
in cm. Stem straightness and branch diameter were
assessed at 5 and 8 years of age using a subjective 1±3
scale. The lowest score (1) was considered the poorest
and the highest score (3) was considered the best.
During a visit to the ®eld site after the 8th-year
assessment, it was noticed that there were great dif-
ferences between provenances with regards to degree
of leaf initiation at the onset of the rainy season. Since
timing of leaf initiation was thought to be related
to degree of adaptability, this trait was assessed
on February 13, 1996, when the study was nearly
10-years old. A subjective grading scale (1±4) was
used:
1. no new leaves initiated
2. some new leaves initiated
3. many new leaves initiated
4. all new leaves initiated
 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135 131

Table 3 heritability (h2i ) was calculated for all traits except

Traits assessed in the field trial for S. apetala at Zambrano, survival using standard procedures suggested by Fal-
Colombia
coner (1981). It was assumed that coef®cient of
Trait Age (years) Abbreviation relationship for the open-pollinated material collected
Survival 1 Sur1 was 0.25 even though some stands were fragmented.
Survival 3 Sur3 The degree and importance of inbreeding in S. apetala
Survival 5 Sur5 has never been assessed to our knowledge.
Survival 8 Sur8
Height 3 Ht3
3. Results
Height 5 Ht5
Height 8 Ht8
Diameter 3 DBH3 3.1. Survival
Diameter 5 DBH5
Diameter 8 DBH8 Survival averaged 87% at the end of 8 years
Volume 5 Vol5
(Table 4). The Cofradia, Honduras provenance had
Volume 8 Vol8
Stem straightness 5 St5 signi®cantly poorer survival (50%) than the Tiquisate,
Stem straightness 8 St8 Guatemala (84%) source which in turn had lower
Branch diameter 5 Bd5 survival that the local Colombian material (95%).
Branch diameter 8 Bd8 With the exception of trees from the Cofradia source,
Leaf initiation 10 Leaf init
which continued to suffer mortality throughout the life
Wood specific gravity 10 Specific gravity
of the trial, survival in the other provenances remained
relatively constant after the ®rst year in the ®eld.
At age 10, bark-to-bark wood cores were extracted
from 4 trees per family per replication (36 cores per 3.2. Growth traits
family) using a 6-mm increment borer. Cores were
sent to North Carolina State University, partitioned at Sterculia apetala trees averaged 1.4 m height
the pith and unextracted speci®c gravity was calcu- growth per year for the ®rst three years but this growth
lated using the standard water displacement method rate declined to <0.75 m per year after age 5 (Table 4).
(Zobel and Talbert, 1984). The two speci®c gravity The mean height of trees in the trial was 7.6 m at 8
readings per tree were averaged to derive a whole tree years of age with the best source from northern
value. A summary of the traits assessed in the study Colombia 8.3 m tall. The volume production of the
are listed in Table 3. Colombia material was 28% greater than the best
Analyses of variance were conducted on the data source from Central America, Cofradia, Honduras.
using the general linear model (GLM) procedure of There were signi®cant differences in height, DBH,
the Statistical Analysis System (SAS, 1989). Because and volume among families at age 8. The best family
of the unbalanced nature of the provenance informa- for height from the northern Colombian source was
tion, least squares means were calculated for the 2.9 m taller than the worst family in the test from
provenance and family effects prior to analyses. Sta- Guatemala and produced 37% more volume than the
tistical differences between provenance means were test average.
determined using pairwise comparisons based on a
studentized maximum modulus and Sidak's uncorre- 3.3. Quality traits
lated t inequality that accommodates unequal sample
size (SAS, 1989). Comparisons among family means There were no signi®cant differences among the
were assessed using the Dunnett's one tailed t-test three provenances for stem form at age 5, but trees
(SAS, 1989). Phenotypic correlation between growth from the Central American provenance were notice-
traits and speci®c gravity were conducted using the ably poorer in bole quality by age 8. The branch
CORR procedure of SAS (1989). diameter scores indicated that the local seed source
Variance components were estimated using the from Colombia had slightly larger branch diameter
VARCOMP procedure (SAS, 1989). Individual tree than the Central American sources. This can be
132 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135

Table 4
Provenance least squares means for the traits assessed in the S. apetala field trial at Zambrano, Colombia from 1 to 10 years of age

Provenance Sur1 Sur3 Sur5 Sur8 Ht3 Ht5 Ht8 DBH3 DBH5 DBH8
(%) (%) (%) (%) (m) (m) (m) (cm) (cm) (cm)
Cofradia, Honduras 75 66 61 50 4.5 5.4 7.6 9.1 11.0 13.7
Tiquisate, Guatemala 89 87 86 84 4.2 5.0 6.7 9.2 11.4 13.3
Northern Colombia 97 97 97 95 5.2 6.2 8.3 10.5 12.8 15.2
Test mean 92 90 90 87 4.8 5.6 7.6 9.9 12.1 14.3

Provenance Vol5 Vol8 St5 St8 Bd5 Bd8 Leaf Specific

init gravity
(m3) (m3)
Cofradia, Honduras 0.0227 0.0477 2.5 2.3 2.4 2.1 3.1 0.204
Tiquisate, Guatemala 0.0211 0.0383 2.5 2.1 2.0 2.0 2.9 0.203
Northern Colombia 0.0328 0.0611 2.6 2.5 2.0 1.9 1.3 0.216
Test mean 0.0273 0.0509 2.5 2.3 2.0 2.0 2.0 0.208

attributed to the fact that trees from Colombia were
also the largest in the test. Family differences for stem
straightness and branch diameter at 5 and 8 years of
age were signi®cant. At 8 years of age, family stem
straightness scores ranged from 1.9 to 2.6 and branch
diameter scores varied from 1.6 to 2.5.
3.4. Leaf initiation
There were signi®cant differences in leaf initiation
prior to the onset of the rainy season between the
northern Colombia provenance and the Central Amer-
ican sources in 1996 (Table 4). Only 10% of the trees
from the northern Colombia provenance had either
initiated most or all of their leaf growth by mid-
February compared with 78% of the trees from the
Tiquisate, Guatemala and 65% of the trees from
Cofradia, Honduras. Fifty-three percent of all the
variation in this trait was at the provenance level.
There were signi®cant family differences
(Pr>Fˆ0.04) for leaf initiation as expected but within
provenance values were relatively consistent. For the
10 open-pollinated families from Guatemala, most of
the trees had already completely leafed-out (scores
from 2.4 to 3.0) by mid-February while trees from the
11 families from northern Colombia were just begin-
ning leaf initiation (scores from 1.0 to 1.9).
ni®cantly higher speci®c gravity (0.216) than the
Central American sources (0.204). Family differences
in speci®c gravity were also found to be signi®cant.
Values for the 10 families from Tiquisate, Guatemala
ranged from 0.191 to 0.219 and for the 11 families
from northern Colombia from 0.204 to 0.235. The
trees with the highest wood speci®c gravity in the test
had values that ranged between 0.250 and 0.262.
There were weak positive phenotypic correlations
between height at 8 years and speci®c gravity at 10
years (rˆ0.21) as well as volume at 8 years and
speci®c gravity at 10 years (rˆ0.13).
3.6. Individual tree heritability (h2i )
Individual tree h2i for the growth and quality traits
did not change much from 3 to 8 years of age
(Table 5). Individual tree h2i for height and volume
were 0.11 and 0.15, respectively, at age 8 and were
under moderate additive genetic control. The h2i for
height was always lower than that for DBH at ages 3,
5, and 8 years of age. The h2i for branch diameter
(0.21) was higher than that for stem straightness (0.11)
at 8 years of age. The individual tree h2i for leaf
initiation at 10 years of age was 0.05 and not under
strong additive control. The h2i for wood speci®c
gravity at 10 years of age was 0.44.

3.5. Wood specific gravity 4. Discussion

The average speci®c gravity for S. apetala at age 10 The magnitude of provenance and family variation
was 0.208. The northern Colombia source had sig- exhibited by S. apetala were quite large and not
 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135
Table 5
Individual tree heritability (h2i ) for growth and quality traits, leaf fall and specific gravity for S. apetala from 3 to 10 years of age
Age (years) Height DBH Volume Stem form
3 0.10 0.19 Ð Ð
5 0.15 0.20 0.13 0.13
8 0.11 0.23 0.15 0.11
10 Ð Ð Ð Ð
noticeably different from a number of other tropical
forest species that have been studied. The local land
race material from Atlantico and Bolivar, Colombia,
were more competitive in survival and growth than
introduced sources from Central America. The poor
performance of the Central American sources com-
pared to the Colombian provenance may be partially
explained by examination of differences in timing of
leaf initiation and leaf fall. The majority of trees from
Guatemala and Honduras initiated their new leaf
growth in February, 1996 during a month where
average rainfall was only 22 mm and 2 months before
the heavy rains begin in May. In 1996, the majority of
trees from northern Colombia did not initiate new leaf
growth until mid-March. In years when rainfalls in late
January and February are atypically above normal,
like those in 1997 where 92 mm fell in a 2-week
period, the Central American and Colombia sources
initiated their new leaf growth at the same time (data
not shown). Furthermore, our ®eld observation indi-
cate that the Central American sources drop their
leaves earlier than do the Colombian sources. There-
fore, it appears that leaf initiation and leaf fall for the
introduced Central American sources are slightly out
of phase with the timing of the wet and dry seasons at
Zambrano. Possibly as a result of this only one open-
pollinated family from the Central American prove-
nances ranked in the top 50 percentile in volume at 8
years of age.
The heritability values derived for the traits
assessed in this study suggest that S. apetala does
not differ much from other tropical species in terms of
the magnitude of additive genetic variance. Heritabil-
ity estimates for growth traits in S. apetala established
in Colombia were similar to those found for Cordia
alliodora (R. & P.) Oken planted in Costa Rica by
Boshier and Henson (1997) and for several of the
tropical pines like P. caribaea var. hondurensis and P.
tecunumanii (Schwd.) Eguiluz and Perry (Hodge and
133
Branch diameter Leaf init Specific gravity
Ð Ð Ð
0.18 Ð Ð
0.21 Ð Ð
Ð 0.05 0.44
Dvorak, 1998a, b). If inbreeding depression is a
problem in S. apetala, then the heritability values
presented in this paper are over estimated. Based on
the individual tree heritability results and the family
rankings across ages in this study, early selection of
trees at age 5 appears to be about as ef®cient as
selection at age 8.
The advantages of planting S. apetala in northern
Colombia are its rapid early growth, its excellent
adaptability to dry sites (if the right provenance is
chosen), and its ability to produce a 3±4 m clear basal
log even in climates that are marginal for tree growth
of most forest species. However, it also has several
disadvantages. First, its volume productivity is poor in
the dry zones of northern Colombia. The average
productivity of the best source from northern Colom-
bia was only 8 m3/ha/year at 8 years of age. Exotic
species like Gmelina arborea Roxb., even when
planted on sandy loams like those at Zambrano, will
out compete it. Second, the wood speci®c gravity of S.
apetala is low relative to other tree species at the same
age and its wood will have dif®culty in being accepted
by local markets on a large scale. This market situation
may change somewhat if the local plywood industry
expands and the demand for core wood increases.
Even though the wood speci®c gravity of S. apetala
will increase with age, the tree does not appear to
produce reasonably dense wood (0.400) before the
age of 40 years. A small sample of 40-year-old S.
apetala trees from a natural stand in Costa Rica had an
average speci®c gravity of 0.370 (Weinmann and
Williamson, 1989). Escobar and RodrõÂguez (1994)
report wood speci®c gravity of the species to be
0.430 but do not report at what age. Third, as men-
tioned previously, S. apetala carries with it a long
list of numerous insect problems. In addition to pod
and seed insects, it is often attacked by a host of
leaf miners and stem borers. For example, in this trial,
the trees were attacked by a defoliator, Arohips sp.
134 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135
(Oletereutride) and by a stem borer Steirastroma
historionica (Cerambycidae). Other researchers in
Cuba and Puerto Rico also report on insect and fungal
infestations on the leaves of S. apetala (Herrera-Isla,
1993; Martorell, 1940).
Pizano/Monterrey Forestal has continued to
develop its breeding activities for S. apetala at a
low level even though the company mainly plants
Gmelina arborea and Bombacopsis quinata (Jacq.)
Dugand. The best trees of S. apetala have now been
selected by Pizano/CAMCORE using a best linear
predictor (BLP) selection index and the best 40 trees
(1.6%) will go into a ®rst-generation clonal production
orchard. Our observations of the S. apetala ®eld trial
indicate seed pods are formed as early as age 5 years in
northern Colombia, much earlier than Janzen (1971)
observed in a natural stand in a riparian evergreen
forest in Costa Rica. Work by Pizano/Monterrey
Forestal also indicates that 9-year-old trees of S.
apetala are easy to clone by simply cutting the top
portion of the main stem, or branches from the top
portion of main stem, and placing these in a rooting
bed in the nursery. Rooting percent is between 80 and
90. However, more research is needed to better study
the importance of plagiotropic effects in S. apetala.
Therefore, once the seed orchard material is crossed,
either naturally or arti®cially, and the most productive
progeny is selected for growth and wood speci®c
gravity, it may be possible to clonally mass produce
the best genotypes of S. apetala. Pizano/Monterrey
Forestal has also established a 20-ha pilot planting of
the species on its property with seeds from a local
collection.
Slow growing species like S. apetala that have little
apparent use seem destined to be forgotten by the
international forest community unless reasons for
actively conserving them are either intensi®ed and/
or new markets are developed (Dvorak, 1996). One of
the ®rst priorities for better conservation of S. apetala
is to more accurately de®ne the geographic extent of
its natural range and the conditions of the stands that
remain. Some reports of its natural distribution prob-
ably include introduced material planted for ornamen-
tal purposes or shade. Furthermore, published
information describes the range of S. apetala in only
broad general terms by country or by region and one is
never sure if S. apetala is being confused with the
many other species in the genus Sterculia that are
morphologically similar to it. More provenance col-
lections should be made and the material tested. One
question that needs to be addressed is, ``Is there
provenance, family or tree-to-tree variation in insect
resistance in S. apetala?'' Most important for the long
term conservation of S. apetala is the need for better
understanding of the medicinal uses of the species.
Furthermore, the development of new markets for its
soft wood of low durability is needed if small local tree
growers or the private sector are ever to become
interested in its establishment as a plantation species.
5. Conclusion
Sterculia apetala has a geographic distribution of
5000 km from southern Mexico to northern Bolivia.
A local provenance from northern Colombia was
superior in growth and stem form and had higher
wood speci®c gravity than introduced sources from
Central America when grown in a genetic test near
Zambrano, Colombia. The magnitude of heritability
exhibited by S. apetala for growth and wood quality
traits at 8±10 years of age was very similar to that of
several other tropical broadleaf and coniferous species
native to the region. The main advantage of planting
S. apetala is its ability to survive in sandy clay soils
in areas with annual rainfall amounts below 1000 mm.
Its disadvantages are its slow growth, low wood
density, and its propensity to be attacked by insects.
Acknowledgements
The authors would like to thank Ing. Ernesto Ponce,
formerly with ESNACIFOR, Honduras, and Forest
Technician Elmer GutieÂrrez, CAMCORE, Guatemala,
for their assistance in the seed collections in Central
America. Furthermore, the authors would like to thank
the Pizano/Monterey research staff for their assistance
in the establishment, maintenance, and measurement
of the trial over the last 10 years. This includes Paul
and Sheryl Bolstad, Mike Kane, Gabriel Castellar and
Alvaro Vallejo. The seed collections of S. apetala
were partially funded by the Of®ce of the Science
Advisor, Agency for International Development
(AID) USA. Their ®nancial support is greatly appre-
ciated. Finally, we would like to thank Gary Hodge,
 W.S. Dvorak et al. / Forest Ecology and Management 111 (1998) 127±135
Jose Luis Romero, and Bruce Zobel for their editorial
comments on the manuscript.
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