Forest Ecology and Management 148 (2001) 287±289

Short communication

The effectiveness of pruning in mitigating

Hypsipyla grandella attack on young mahogany
(Swietenia macrophylla King) trees
Jonathan P. Cornelius*
Tropical Agricultural Research and Higher Education Center (CATIE), Turrialba 7170, Costa Rica
Received 21 March 2000; accepted 4 June 2000

Abstract

The planting of mahogany (Swietenia macrophylla King) is severely limited by the attack of the shoot-borer Hypsipyla
grandella. Pruning is sometimes advocated as one means of controlling and/or mitigating shoot-borer attack. However, there
appears to be no published experimental support for such recommendations. The present short communication reports on the
results of an experimental study in which trees in 13 pairs were subjected to contrasting pruned/control (unpruned) treatments.
Twenty-nine months after planting, pruned trees had signi®cantly better values for form traits (p between 0.012 and 0.09) with
no apparent difference in growth traits. It is concluded that pruning can mitigate the Hypsipyla problem. The generality of the
results is brie¯y discussed. # 2001 Elsevier Science B.V. All rights reserved.

Keywords: Meliaceae; Pest-control; Shoot-borer

1. Introduction However, published empirical evidence for the ef®-

Fast growth, adaptability, good form and high value
are desirable properties of plantation forestry species.
Mahogany (Swietenia spp.) ful®ls these and other such
characteristics. That this potential has not been rea-
lized is largely due to the `shoot-borer problem'.
Larvae of Hypsipyla grandella (Lepidoptera: Pyrali-
dae) bore into the leading shoots of young trees,
reducing height increment and, more importantly,
causing forks of the main stem that greatly reduce
commercial value (Newton et al., 1993).
Pruning of attacked leaders and/or of the codomi-
nant resprouts that attacks may cause is sometimes
advocated as a solution to the Hypsipyla problem.
*
Tel.: ‡506-558-2625; fax: ‡506-556-1533.
E-mail address: jcorneli@catie.ac.cr (J.P. Cornelius).
cacy of pruning seems to be restricted to Chable's
(1967) report of positive experiences in Honduras and
two further, unpublished, reports of favorable results
described by Mayhew and Newton (1998). The latter
authors recommend that `rather than concentrating on
shoots that have been attacked, pruning should seek
to encourage those shoots which are displaying
vigorous vertical growth, by removal of those which
are not'.
However, it is not axiomatic that removal of com-
peting leaders will lead to restoration of one dominant
leader, except during the period immediately follow-
ing pruning. Removal of competing leaders weakens
the correlative inhibition imposed by the uppermost
shoot, and may stimulate the sprouting of adventitious
buds below the pruning point. Equally importantly,
there may be other, unforeseen, practical problems. It

0378-1127/01/$ ± see front matter # 2001 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 0 ) 0 0 5 3 1 - 4
288 J.P. Cornelius / Forest Ecology and Management 148 (2001) 287±289
seems justi®ed, therefore, to test experimentally the
hypothesis that pruning can effectively control or
mitigate Hypsipyla attack. In the present note, I report
on such an experiment.
2. Materials and methods
In July 1996, 48 three-month old Swietenia macro-
phylla seedlings of Upala, northern Costa Rica pro-
venance, were planted in the CATIE estate (Turrialba,
Costa Rica: 600 m a.s.l., premontane wet forest
life-zone (Holdridge, 1967; Tropical Science Center,
1969), mean annual precipitation 2544 mm (S.D.
ˆ551 mm), mean of 1.8 (S.D.ˆ2.2) dry (<80 mm)
months per year (MIRENEM, 1988)). The average
spacing was 3 m2.5 m. Twenty-eight attacked trees
in 14 pairs were selected for inclusion in the experi-
ment during the 4 weeks following the ®rst attack in
May 1997. The pairing criterion was tree height and
the difference in total height of the two members of
any one pair was within the range 1±10 cm with two
exceptions (differences of 28 and 31 cm). The mean of
the difference between pairs was 4.0 cm (S.D.ˆ12.92)
(overall mean of the 28 trees was 102 cm). One tree of
each attacked pair was randomly chosen as a `treat-
ment' tree, and other designated a `control' tree. In
October 1997, utility workers killed three trees in
three different pairs. Two of these trees were control
trees. They were replaced by randomly selecting, from
the non-experimental trees, an attacked tree whose
height at experimental initiation had been comparable
to that of the corresponding treatment tree. The lost
treatment tree could not be replaced, as the non-
experimental trees had not been pruned. Conse-
quently, the ®nal number of pairs fell to 13.
The pruning regime applied to the `treatment' trees
had two components. Periodically (see below), each of
the 26 experimental trees was inspected for Hypsipyla
attack. When Hypsipyla larvae were present on a
treatment tree, a `protection pruning' was done: the
seedling was cut back with secateurs to just below the
lower limit of the larval tunnel. The aim of the
`protection pruning' was to free the tree of Hypsipyla
infestation. Subsequently, in those trees which
responded to decapitation by the production of more
than one resprout, a `form pruning', consisting of the
removal by pruning of all shoots except one, was
carried out. In order to minimize renewed attack on
the selected shoot, the form pruning was delayed until
the latter had lost its initial succulence and appeared
partially ligni®ed and, therefore, less attractive to the
pest. Inspections were made fortnightly, except during
the period of peak attack (see below, Section 3), when
trees were visited every 1±3 days.
Grass competition was controlled through clean
weeding with a sharpened spade of a circle of radius
0.7±1.0 m around each tree, with mowing of interven-
ing areas. Fertilizer (10±30±10 NPK) was applied
to the base of each tree in January 1997 (50 g per
tree), June 1997 (50 g per tree) and January 1998
(100 g per tree).
At age 29 months, the following variables were
measured: total height, height to ®rst fork (equal to
total height in unforked trees), dbh, basal diameter,
number of codominant leaders (ˆ1 in the case of
unforked trees). In addition, an experienced ®eld
worker without knowledge of the experimental design
compared the overall quality of the two trees of each
pair, taking into account both form and growth. One
tree of each pair was booked as inferior, the other
superior (there were no `draws'). The presence of
treatment effects on the variables was evaluated using
the Wilcoxon signed-rank test, except for the quality
comparison, which was analyzed using the sign test
(Snedecor and Cochran, 1980; Ramsey and Schafer,
1997).
3. Results
The trees were attacked from 27 May 1997 through
to mid-July 1997. Intensity of attack was not recorded
by tree, but varied considerably from aborted attacks
(presumably due to predation) to multiple attacks on
both succulent and woody tissue. No attacks were seen
between 19 July and 31 August 1997. There were
sporadic attacks in September and October 1997, but
none during the rest of 1997 or during 1998. All the
planted trees were alive at the end of the experimental
period (December 1998), except those killed by the
utility workers.
There is little or no statistical evidence of negative
or positive effects of pruning on height, dbh or basal
diameter (Table 1). For the other variables, there are
suggestive (`quality comparison', pˆ0.09) to strong
 J.P. Cornelius / Forest Ecology and Management 148 (2001) 287±289 289

Table 1
Mean differences in six response variables between treatment (pruned) and control (unpruned) pairs in a pruning experiment of mahogany (S.
macrophylla) in Turrialba, Costa Rica, 29 months after planting

Variable Mean Mean differencea Ncontrol superior

b
Ntreament superior
b
Pc

Height 3.34 m 0.32 m (S.D.ˆ0.85) 5 7 0.21d

Height to first fork 2.88 m 0.12 m (S.D.ˆ0.13) 3 9 0.012d
Dbh 3.0 cm 0.3 cm (S.D.ˆ0.68) 4 9 0.18d
Basal diameter 5.4 cm 0.1 cm (S.D.ˆ1.45) 7 6 0.89d
Number of codominant leaders 1.3 0.5 (S.D.ˆ0.66) 0 6 0.02d
Quality comparison n.a. n.a. 3 10 0.09e
a
i.e. SYt Yc/n, where Yt is observed value of response variable on pruned tree and Yc is observed value of response variable on unpruned
tree.
b
Number of pairs in which control tree (column 4) or treatment tree (column 5) was superior (two columns sum to <13 when the 1
treatment/control pairs have the same mean).
c
Two-tailed probability of observed mean differences under null hypothesis.
d
Wilcoxon's signed-rank test.
e
Sign test.

(height to ®rst fork, pˆ0.012) indications of a bene- Acknowledgements

®cial effect of pruning. None of the pruned trees, as
compared to seven unpruned trees, were forked; the
observed differences in `quality comparison', as well
as number of leaders, re¯ect this.
4. Discussion and conclusions
The results suggest that, under conditions similar to
those obtaining during the experiment, trees subjected
to a pruning regime of the sort implemented will be
better formed and, therefore, more valuable than
unpruned trees. That is to say, pruning can mitigate
the Hypsipyla problem. The study offers some experi-
mental con®rmation of the experiences mentioned
previously.
Generalization to other conditions, particularly
other attack intensities, would be speculative. How-
ever, it seems reasonable to suggest that, under more
severe and prolonged attack, the superiority of pruned
trees might be greater. In the present case, the lightness
of attack and consequent excellent form of several
control trees was such as to preclude any inferiority of
these with respect to the respective paired pruned
trees. These conjectures could be tested through sam-
pling of other attack scenarios, using spatial or tem-
poral replication. Further studies should also seek to
clarify the economic viability of this and other pruning
regimes for this high-value species.
I thank Mr. Manuel Sojo, Mr. Oldemar Baeza
(CATIE) and the CATIE maintenance department
for their help in trial maintenance and measurement,
Mr. Carlos Navarro for supplying the plants, and Dr.
G. Galloway (CATIE), Dr. J.P. Dick, Dr. R.R.B.
Leakey, Dr. A.D. Watt (Institute of Terrestrial Ecol-
ogy, Penicuik, Scotland) and two anonymous
reviewers for their careful reviews and/or suggestions
of improvements to the manuscript.
References
Chable, A.C., 1967. Reforestation in the Republic of Honduras.
Ceiba 13 (2), 1±56.
Holdridge, 1967. Life Zone Ecology, Revised Edition. Tropical
Science Center, San JoseÂ, Costa Rica, 206 pp.
Mayhew, J.E., Newton, A.C., 1998. The Silviculture of Mahogany.
CABI Publishing, Wallingford, UK, 226 pp.
MIRENEM, 1988. Catastro de Las Series de Precipitaciones
Medidas en Costa Rica. Imprenta Nacional, San JoseÂ, Costa
Rica, 361 pp.
Newton, A.C., Baker, P., Ramnarine, S., MeseÂn, J.F., Leakey,
R.R.B., 1993. The mahogany shoot borer: prospects for control.
For. Ecol. Mgmt. 57, 301±328.
Ramsey, F.L., Schafer, D.W., 1997. The Statistical Sleuth. Duxbury
Press, Belmont, CA, 742 pp.
Snedecor, G.W., Cochran, W.G., 1980. Statistical Methods, 7th
Edition. Iowa State University Press, Iowa, 507 pp.
Tropical Science Center, 1969. Mapa EcoloÂgico de Costa Rica.
Tropical Science Center, San JoseÂ, Costa Rica.