Biodiversity and Conservation 11: 521–541, 2002.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

Impact of logging on the richness and diversity of

forest butterflies in a tropical dry forest in Thailand

JABOURY GHAZOUL
T.H. Huxley School of Environment, Earth Sciences and Engineering, Imperial College of Science,
Technology and Medicine, Silwood Park, Ascot, Berks, SL5 7PY, UK (e-mail: j.ghazoul@ic.ac.uk;
fax: +44-207-5942308)

Received 20 December 2000; accepted in revised form 12 April 2001

Abstract. Studies on the impact of logging on tropical forest butterflies have been almost exclusively
conducted in moist forest habitats. This study considers the impacts of small-scale logging on butterfly
communities at three sites of varying disturbance intensity in a tropical dry forest in western Thailand.
Butterfly species richness was similar at all sites, but the abundance of butterflies and diversity of the but-
terfly community decreased with increased logging disturbance. The recorded decrease in diversity at the
relatively large sampling scale used lends further support to the hypothesis that disturbance effects are scale
dependent. Species abundance data for butterflies fitted a log-normal distribution at all sites, but also a log-
series distribution at the two disturbed sites. These analyses suggest a more complex butterfly community at
the undisturbed site, but also that log-series and log-normal distributions may not to be sufficiently sensitive
to be useful indicators of community changes following logging. Community ordination separates both the
butterfly species and transect samples into three distinct regions corresponding to the three study locations.
Ordination axes are correlated with tree density, understorey cover and understorey plant richness. Species
with the smallest geographic ranges tend to be the least abundant and occurred most frequently in the
undisturbed site. The observed diverging responses to disturbance among butterfly families diminishes the
value of butterfly communities as biodiversity indicators, and forest managers should perhaps focus on
restricted range species or of groups of recognized sensitive species for this purpose.

Key words: community structure, deciduous dipterocarp forest, disturbance, Lepidoptera, range size

Introduction

Tropical forest conservationists have the task of managing ecosystems that consist of
a huge diversity of organisms that vary widely in abundance, distribution and ecologi-
cal function. Human land-use must be integrated with conservation, and it is therefore
essential to understand how biological communities respond to anthropogenic distur-
bances. Concern about conversion of primary tropical forests to secondary growth has
instigated a number of studies on the responses of plant and vertebrate components of
forest communities to disturbance (reviewed in Ghazoul and Simmons 2001). As yet
relatively few studies have focussed on invertebrates, despite their dominance of the
animal community in species richness, abundance and biomass (Wilson 1987; Stork
1988). Insects have been shown to be sensitive to changes in vegetation composition
522

and the physical attributes of the environment (Erhardt 1985; Gardner et al. 1995;
Wood and Gillman 1998) resulting mostly in a decrease in insect diversity (Holloway
1987; Holloway et al. 1992). However some studies have shown that disturbance
results in negligible change (Wolda 1987), or even an increase in community diversity
(Raguso and Llorente-Bousquets 1990; Hill et al. 1995; Hamer et al. 1997; Spitzer
et al. 1997). Increases in species richness are often due to invasion of disturbed ar-
eas by generalist and widely distributed species, and the resulting homogenisation
of the world’s biological communities is an important threat to global biodiversity
conservation. Loss of invertebrate diversity may have further indirect effects, acting
through the disruption of ecological processes mediated by invertebrates (Ghazoul
and Hill 2001). Studies of human impacts on biodiversity should therefore place em-
phasis on both species distributions and their ecological interactions. If conservation
strategies are to be integrated with social and economic factors it is further necessary
to construct reliable generalisations about impacts of anthropogenic disturbances on
forest invertebrate communities. Currently there are few data to permit generalisation
about human impacts on invertebrate biodiversity, although butterflies are among the
best studied group in this respect. However, the addition of recent studies on butter-
flies may soon allow for generalisations necessary to inform forest management and
ecologically sustainable development (Ghazoul and Hellier 2000).
Tropical butterfly assemblages are highly diverse, with many endemic species,
most of which are dependent to some extent on closed-canopy forest (Collins and
Morris 1985; Sutton and Collins 1991). Compared with other invertebrate groups,
the high visibility of butterflies, together with their relatively well-known taxonomy,
has resulted in this group receiving a reasonable amount of attention throughout the
tropics. Additionally, the species richness of butterflies has been correlated with the
richness of vertebrates and vegetational types (Erhardt 1985; Thomas and Mallorie
1985; Viejo 1989; but see Lawton et al. 1998). Butterflies are therefore appropriate
subjects for the study of logging impacts on biotic communities and have been wide-
ly used to assess patterns of tropical insect diversity in forest conservation studies
(Brown 1991; DeVries et al. 1997; and references therein).
This study investigates logging impacts on the structure of a forest butterfly com-
munity. The research describes the impact of logging on the richness and diversity
of dry forest butterflies in Thailand, and places it in the context of similar studies
conducted across the tropics.

Methods

Species and study site

The dry deciduous forest (DDF) in Thailand is dominated by four tree species belong-
ing to two genera, Shorea and Dipterocarpus (Dipterocarpaceae). Prior to a national
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logging ban in 1989 Shorea siamensis Miq. and S. obtusa Wall. ex Bl. were extracted
commercially from many forests in Thailand. Since then illegal small-scale extraction
of Shorea trees for domestic use continues throughout the country, but particularly
where villages occur near protected forest areas where substantial numbers of trees
remain.
This study was conducted during the early-mid dry season from 16th October to
12th December 1995 at three sites of differing disturbance intensity in DDF at
Huay Kha Khaeng Wildlife Sanctuary (HKK), Uthai Thani Province, in western
Thailand (15◦ 36 N 99◦ 20 E). HKK has a total area of 2575 km2 of which 381
km2 is DDF most of which occurs in a single 260 km2 block on the eastern
edge of the sanctuary (Nakhasathien and Stewart-Cox 1990) where the study was
conducted. Anthropogenic disturbance over two decades has created a gradient of
increasing disturbance intensity from the sanctuary interior to its boundary. Three
sites were selected to represent this gradient. The undisturbed study site (labeled
henceforth as U) was a 6 × 3 km area located within the much larger undisturbed
forest region where human disturbance of any kind is strictly prohibited and
adequately enforced. Undisturbed forest begins at about 9 km within the sanctuary
boundary and extends towards its interior, and site U was located between 14–20
km along a restricted access dirt road starting at the wildlife sanctuary boundary.
Shorea trees had been extracted in small numbers from a moderately-disturbed
forest site (M) in the vicinity of the wildlife sanctuary headquarters where a small
village was located prior to its relocation away from the area in 1990. Since then
this moderately-disturbed forest area, which is located 6–9 km within the park
boundary, has been subject to the same strict controls as the undisturbed site.
Observations were made through the entire extent of moderately-disturbed forest
which covered an area of approximately 12 km2 . Highly disturbed forest exists in
a 3–4 km-wide buffer zone that extends along the length of the north-eastern
boundary of the sanctuary for some 15 km. Prior to the creation of the buffer zone
in 1990 this area was subject to widespread extraction of S. siamensis trees.
The disturbed site (D) lies outside the sanctuary but within this buffer zone and
observations of butterflies were made in an approximately 10 km2 area within this
disturbed region of forest. Sites M and U were roughly equidistant, by about 7 km,
from the edge of the buffer zone.
All sites are located within the same forest basin with similar elevations and
topography. The soil is homogeneous throughout this basin (Nakhasathien and
Stewart-Cox 1990) and no significant differences between sites were apparent in
depth, pH, organic content, or water, potassium and phosphorous availability,
although nitrogen did occur at a significantly higher concentration at D (J. Ghazoul,
unpublished data). Grazing and small-wood collection are prohibited in the sanctuary
but limited grazing is permitted in the buffer zone and may contribute to the relatively
high soil nitrogen content in this area. Soil resources were therefore assumed to be
equally available among sites.
524

Tree density and site vegetation characteristics

Tree densities were determined within 20 randomly situated 9-m radius plots at each
site by counting all trees exceeding 10 cm DBH (diameter at breast height). Under-
storey plant cover and species richness were recorded in 6 m radius plots (113 m2 )
at the start, middle and end of each butterfly census transect described below. An
estimate of percentage understorey vegetation cover was made by taking the mean
of independent estimates made by two observers. One species Eupatorium odorata
(Compositae) accounted for much of the cover in the disturbed site and its cover was
also separately estimated. Where this species was represented by fewer than 10 plants
in a plot it was scored as having 1% cover for each individual plant present. In plots
where 10 or more individuals occurred it was given a percentage cover estimate that
started from 10%.

Butterfly census techniques

Prior to butterfly censusing three observers spent 15 days in the field (between 16th
October and 5th November) constructing a reference collection (now housed at Royal
Forest Department, Bangkok) and familiarising themselves with the local butterfly
fauna. Individual workers were already familiar with the butterfly species of the re-
gion through earlier work in Vietnam and Thailand. Species were identified using
Pinratana (1979, 1983, 1988, 1992). Survey methods were similar to those described
by Pollard (1977) and Spitzer et al. (1993). Butterflies were surveyed for 30 min along
twenty 500 m transects at each site, each transect being used once only. Transects
were located within the forest to avoid existing tracks and paths as far as possible.
Within a site transects were mostly parallel to each other and adjacent transects were
separated from each other by a minimum distance of 200 m and up to about 600 m.
Using binoculars (Leica 8 × 32 mm and 10 × 42 mm), two observers identified to
species all butterflies except for Lycenaidae and Hesperiidae that were seen within an
estimated 20 m horizontally to their side of the transect. This distance was determined
to be the farthest that butterflies could be identified with certainty (using binoculars)
at all sites. The distance is greater than most other studies on forest butterflies due to
the relatively open forest structure typical of dry deciduous dipterocarp forests during
the dry season. Butterflies that could not be clearly identified on the wing were caught
in a kite net for closer examination. Unidentified butterflies that could not be caught
were listed as unidentified. Surveys were conducted between 9 and 12 h (during which
time butterfly activity appeared to peak) on days with constant sunlight during a 5-
week period from 6th November to 12th December. As with other similar studies
the Lyceanidae and Hesperiidae were excluded from the surveys due to difficulty
of identifying them to species from field observations. These families accounted for
39% of the numbers of all recorded butterflies, so this study does not represent a
complete investigation of the whole butterfly community. Species belonging to the
 525

genus Neptis (Nymphalidae) were difficult to distinguish on the wing and all of these
were caught for identification.

Geographic distribution of butterfly subspecies

The geographic range of butterfly subspecies were derived from Pinratana (1979,
1983, 1988, 1992) where distributions are given as countries or regions within which
the butterflies are known to occur. The Oriental region was divided into seven geopo-
litical regions corresponding broadly to differences in climate and biogeography: (1)
Burma and Thailand excluding the peninsula, (2) Peninsular Thailand, Malaysia and
Singapore, (3) Indochina and South China, (4) China (excluding South China) and
Japan, (5) Indonesia and the Philippines, (6) North India, Sikkim, Nepal and Assam,
and (7) India (excluding North India), Sri Lanka and the Andaman Islands. Butterflies
were assigned an additive score of 1 for each region in which they occur, as listed by
Pinratana. Species occuring in only part of a geographical region were to be scored
as occupying it. Although the potential maximum score is 7, all the butterfly species
that occurred in more than four regions actually occurred in at least part of all seven
regions. Thus for purposes of data analysis and to avoid classes devoid of data, the
maximum score was set at 5. The geographic ranges of butterfly subspecies were thus
scored on a scale of 1–5 with a score of 1 for those species confined to Burma and
Thailand excluding the peninsula, and a maximum score of 5 reflecting a range that
included five or more of the geographic regions listed above.

Statistical analysis

To overcome some of the difficulties associated with pseudo-replication within the

sampling regime (which were unavoidable given the practicalities of the study), a
Mantel test (Mantel Nonparametric Test Calculator 2.0, Liedloff 1999) was used to
compare transect matrices of butterfly compositional similarity and geographical dis-
tance between transect pairs within D, M and U and across the whole data set. If the
null hypothesis, that there is no effect of distance on butterfly compositional similarity
between pairs of transects, is upheld within but not between sites then this would sup-
port the contention that any butterfly compositional differences result from inherent
differences between sites and are not generated by distance alone.
The butterfly assemblages from each site were examined by fitting species abun-
dance data to log-series and log-normal distributions (May 1975; Magurran 1988;
Krebs 1989). EstimateS 5.0.1 computer package (Colwell 1997) was used to com-
pute randomised species accumulation curves and to statistically estimate true species
richness. The effect of sample order can be removed by randomizing many times and
obtaining an average value which may be used to produce a smooth species accumu-
lation curve. In this way the particular order that observations were collected will not
affect a comparison of the data set. EstimatesS selects a single sample (transect) at
526

random and computes the richness estimators based on that sample, then randomly se-
lects a second sample for which the estimators are recalculated. Fifty randomisations
were completed for each data set. Cumulative species richness curves were drawn
using mean and standard deviation for each sample. A jack-knife procedure was used
to calculate mean and standard deviation values of Simpson’s index of diversity for
butterfly assemblages at each site. Simpson’s index was favoured over other indices
as it is particularly sensitive to changes in the more abundant species of a commu-
nity (Krebs 1989) for which rigorous data are relatively easy to obtain. This index
also has minimal bias when sample sizes are small (Lande 1996) and when species
accumulation curves intersect (Lande et al. 2000). Analyses were performed with
STATISTICA Release 5 (StatSoft Ltd). Ordination of the butterfly communities was
conducted using detrended correspondence analysis (DECORANA).

Results

Tree density and site vegetation characteristics

Disturbance by extraction of trees resulted in decreased tree density and an associat-

ed increase in ground cover by herbaceous plants (Table 1). At M and D there was
much variation in herbaceous understorey cover while at U understorey cover was
uniformly low (Table 1).

Butterfly species abundance and community structure

The Mantel test results showed a strong effect of distance on transect butterfly com-
position across forest types (g = 7.99, R = 0.36, P < 0.01) but not within forest
types (Undisturbed, g = 1.05, R = 0.11, P = NS; Moderately disturbed, g = 0.44,
R = 0.05, P = NS; Disturbed, g = 0.01, R = 0.01, P = NS), indicating (al-
though not proving) that inherent differences between the sites account for butterfly

Table 1. Tree density and understorey plant cover at three sites differing in disturbance impact.

Understorey ground E. odorata

Tree density Understorey cover (%) (% cover)
(/254 m2 ) Understorey species/plot mean ± SE (inter- mean ± SE
Site Mean ± SE (CV) species mean ± SE (median) quartile range) (median)

U 11.6 ± 0.8 (53) 14 3.8 ± 0.2 (3.7) 12.3 ± 0.6 (2.4) 0.08 ± 0.08 (0)
M 10.1 ± 1.0 (77) 17 2.8 ± 0.3 (3.0) 12.3 ± 1.8 (4.0) 0.33 ± 0.26 (0)
D 7.3 ± 0.8 (85) 10 3.0 ± 0.2 (3.0) 42.4 ± 4.8 (43.9) 30.8 ± 2.8 (30)
F(2, 57) = 6.4 H(2, 60) = 12.0 H(2, 60) = 25.7 H(2, 60) = 161.5
P = 0.003 P = 0.0025 P < 0.001 P < 0.001
Kruskal–Wallis test applied to untransformed data. At each site data were collected from 20 transects and
from three plots per transect. Coefficient of variation (CV as a %) for tree density values permits direct
comparison of data variability at the sites.
 527

dissimilarities across sites. Distance effects, as might be expected from a non-inde-

pendent sampling regime, can therefore be ruled out.
Ninety-six percent of all butterflies seen (excluding Lyceneidae and Hesperiidae)
were identified in the field, and there was no significant difference across sites in
unidentified butterflies as a proportion of the total recorded (Kruskall–Wallis test:
H(2,60) = 0.28, NS). Owing to their small and equal proportions across sites, un-
identified butterflies are excluded from subsequent analyses and discussion. In total
1936 butterflies from 60 transects were observed and identified. The greatest number
occurred at the undisturbed site (740 individuals, 38.2% of total) while smaller and
equal numbers were recorded at sites M (597, 30.9%) and D (599, 31.0%) (see Table 2
for statistical information). From a total of 53 butterfly species, 39 were observed at
U, 40 at M and 37 at D (see Table 3) and there was no difference in the number of
species recorded in per transects across sites (Table 2). At all sites cumulative butterfly
species richness curves leveled off rapidly after the fifth transect (Figure 1) indicating
similar sampling effectiveness despite differing forest architectural structure at each
location. Cumulative species richness over the 20 transects was consistently lower at
D than other sites (Figure 1), although the asymptotes show that new species were
still being recorded later at D (Figure 1).
Sites M and D had long tails of rare species (Figure 2), and 22 species (41.5%
of the total) were recorded less than 10 times at all sites. Fifteen percent of butterfly
species at U each accounted for less than 0.5% of the total sample, while the corre-
sponding figures at M and D were 30 and 35% of butterfly species respectively. The
commonest species Eurema hecabe was recorded 245 times accounting for 12.7% of
all individuals at all sites. Analysis of variance on jack-knifed values for Simpson’s
index produced significant differences between sites (F2, 56 = 69.16, P < 0.001;

Table 2. Diversity index scores for butterfly communities at sites U, M and D based on 20 transect surveys
at each site.

Moderately
Undisturbed disturbed Disturbed

Individuals/transect (F(2, 57) = 5.45, P < 0.01) 37.0 ± 3.8 29.9 ± 2.4 29.1 ± 1.8
Species/transect (F(2, 57) = 1.05, P = NS) 14.3 ± 0.8 13.0 ± 0.8 13.0 ± 0.4
Total species recorded 39 40 37
Simpson’s D (Jack-knife pseudo-values) 0.958 ± 0.005 a 0.898 ± 0.01 b 0.937 ± 0.005 a
(mean ± SE; F(2, 57) = 17.27, P < 0.01)
Q-statistic 13.1 10.0 8.1
Log-normal (λ) 81.8 75.3 64.8
Fits of models
Log-normal Yes Yes Yes
Log-series No Yes Yes
Simpson’s index, D, is calculated using Pielou’s formula, quoted in Krebs (1989), for finite populations.
Berger–Parker index of dominance = nmax /n, where nmax is the number of individuals recorded in the
most abundant species, and n is the total number of individuals. ANOVA is used to compare values across
sites.
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Table 3. Pooled results from 20 butterfly transects at each site and geographic range category (refer to
text for details).

Moderately Potential
Species Undisturbed disturbed Disturbed Range vagrants

Pieridae
A. paulina adamsoni (Moore) 8 0 3 1
C. pomona pomona (Fabricius) 45 48 61 4
C. scylla cornelia (Fabricius) 5 0 1 3
D. pasithoe thyra (Fruhstorfer) 0 1 1 3 V
D. descombesi descombesi (Boisduval) 37 33 18 2
D. acalis pyramus (Godart) 5 0 5 2
E. brigitta fruhstorferi (Moore) 64 27 6 3
E. hecabe contubernalis (L.) 74 99 72 4
Hebomoia glaucippe glaucippe (L.) 0 4 1 3
Ixias pyrene verna (Druce) 8 8 8 2
Leptosia nina nina (Fabricius) 0 5 2 3
Papilionidae
A. adamsoni (Grose-Smith) 14 35 35 1
A. zaleucus (Hewitson) 5 0 0 2
T. aecus (Felder) 2 6 16 3
Papilio demoleus malayanus (Wallace) 4 6 4 3
Nymphalidae
Athyma asura asura (Moore) 0 1 0 2 V
A. perius perius (L.) 10 6 9 5
A. seleneophora latifascia (Talbot) 0 3 0 2 V
Cethosia cyane euanthes (Fruhstorfer) 9 9 11 2
Ch. marmax marmax (Westwood) 6 0 0 1
Chersonesia risa risa (Doubleday) 10 0 0 3
Cirrochroa tyche mithila (Moore) 3 2 7 3
Eu. aconthea garuda (Moore) 0 1 1 3 V
Eu. lepidea lepidea (Butler) 0 2 0 2 V
Eu. pardalis jadeitina (Fruhstorfer) 11 5 21 2
Hypolimnas misippus misippus (L.) 24 1 0 5
Limenitis daraxa daraxa (Doubleday) 0 2 0 2 V
Moduza procris procris (Cramer) 0 0 1 2 V
N. clinia susrata (Moore) 1 2 0 3 V
N. hylas (L.) 62 36 64 5
Phaedyma columella martabana (Moore) 15 20 41 2
Pantaporia sandaka davidsoni (Eliot) 4 10 2 3
Polyura athamus samatha (Moore) 5 1 3 2
P. schreiberi assamensis (Rothschild) 0 5 0 2
Pr. almana almana (L.) 18 10 4 4
Pr. atlites atlites (L.) 26 34 22 3
Pr. lemonias lemonias (L.) 20 12 40 4
Pr. iphita iphita (Cramer) 12 0 0 3
Pr. orithya ocyale (Hübner) 0 9 0 3
Vindula erota erota (Fabricius) 0 0 5 2
Yoma sabine vasuki (Doherty) 1 0 0 2 V
Danaidae
Euploea core godartii (Lucas) 6 15 34 2
E. mulciber mulciber (Cramer) 5 7 2 4
Da. chrysippus chrysippus (L.) 0 0 1 5 V
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Table 3. Continued.

Moderately Potential
Species Undisturbed disturbed Disturbed Range vagrants

Da. genutia genutia (Cramer) 7 3 38 5

Parantica aglea melanoides (Moore) 5 11 4 3
Pa. melaneus plataniston (Cramer) 0 3 3 3 V
Tirumala septentrionis septentrionis (Butler) 6 0 0 4
Satyrinae
L. verma stenopa (Fruhstorfer) 25 4 0 1
Melanitis phedima abdullae (Distant) 67 25 17 5
Mycalesis intermedia (Moore) 15 29 2 3
My. perseus (F.) 87 57 34 5
Y. similis (Elmes and Edwards) 9 0 0 1
Potential vagrants represent species that were recorded very infrequently (<0.5% of the sample) at all three
sites.

Figure 1. Randomised cumulative species richness curves for butterflies observed along transects at three
sites differing in disturbance intensity. Values calculated from 50 repetitions at each site with means and
standard error bars shown.

Table 2) with highest diversity at U. All samples fit a log-normal distribution, but log-
series models only fit sites M and D. Differences in species abundance distributions
across sites may affect the validity of diversity indices that use the proportional abun-
dance of species in the sample (e.g. Simpson’s). Therefore, also presented in Table
2 is the Q-statistic, which is a measure of the inter-quartile slope of the cumulative
species abundance curve and provides an index of diversity at the three sites with-
out weighting towards very abundant or very rare species. A similarly high diversity
530

Figure 2. Butterfly species from three sites differing in disturbance intensity ranked according to their
proportional abundance in disturbed and undisturbed forest.

score for the butterfly community at site U is obtained using this statistic (Table 2)
confirming high butterfly diversity at U relative to sites M and D.
The responses to disturbance by individual species were not widely predictable by
higher taxonomic grouping, except, perhaps, for the Satyrinae where all five species
showed a marked decrease in abundance with increasing disturbance.
Five butterfly species appear to be restricted to the undisturbed forest and were
not recorded elsewhere (Table 4). Only one species was confined to M and no species
were recorded exclusively from D. Of the species recorded from more than a single
site, five show a significant predominance at the undisturbed site, while another five
have significantly higher abundance at the disturbed site, and only a single species
predominates at M. Thus 10 of the 39 species included in these analyses occur either
exclusively at U or predominate at this site.

Geographic range and species abundance

The least abundant species at HKK tend to be those with the smaller geographic
ranges (Figure 3; Kruskal–Wallis test comparing average ranked subspecies abun-
dances, by frequency of occurrence in all transects, across the five geographic range
 531

Table 4. Species listed in the first part of this table are those that occur entirely in a single site.

Site Total n Occurrence (%) χ2

Species occurring in a single site

Che. risa (Nym) U 10
Pr. iphita (Nym) U 12
Y. similis (Sat) U 9
Ti. septentrionis (Dan) U 6
Ch. marmax (Nym) U 6
Pr. orithya (Nym) M 9
Species predominant in one site
H. misippus (Nym) U 24 96 29.5
L. verma (Sat) U 29 86 24.3
E. brigitta (Pie) U 95 71 29.1
M. phedima (Sat) U 109 61 24.2
My. perseus (Sat) U 178 49 12.9
My. intermedia (Sat) M 46 63 12.2
Da. genutia (Dan) D 48 79 30.3
T. aecus (Pap) D 24 67 8.0
Eu. core godarti (Dan) D 55 62 13.4
Pr. lemonias (Nym) D 72 56 10.7
Ph. columella (Nym) D 76 54 9.7
Family association is abbreviated after the species name. To be included in this analysis it was arbi-
trarily decided that a species must have been recorded at least six times. In the second part of the table
species are listed that have greater than expected abundance recorded at one site (expected abundance
at any site is one-third of the total abundance across all sites; χ 2 critical value = 7.88, at P = 0.005
(Dunn–Sidak adjusted for multiple tests) with 1 d.f.). Species with at least 20 recorded individuals
were included in this analysis.

classes: H(4, 53) = 9.93, P = 0.042), though this tendency is not observed among
species that have range size categories of 1–3. To avoid problems associated with
clumped distributions the analysis was repeated by substituting total butterfly abun-
dance with the number of transects from which each species was recorded. Similar
result were obtained (Kruskal–Wallis test: H(4, 53) = 12.64, P = 0.013). At U, five
species, Appias paulina, Atrophaneura adamsoni, Charaxes marmax, Lethe verma
and Ypthima similis, belonged to the smallest geographic range category (Burma and
Thailand excluding the peninsula), but all except A. adamsoni declined significantly
or disappeared completely at M and D (see Table 3). Butterfly species associations,
characterized by their position on the first DECORANA ordination axis, were not cor-
related with range size (Spearman rank correlation: n = 53, r = 0.001, t51 = 0.007,
P = NS) indicating that restricted range species are not closely associated with each
other, at least at the scale and location of this study.

Community ordination

Community ordination of the transects by butterfly species abundance using DEC-

ORANA revealed a clear differentiation along the first ordination factor of butterfly
532

Figure 3. Box plots illustrating the association between butterfly species abundance and geographic range.

species between D and the two less disturbed sites M and U. The second factor load-
ing axis further separated M from U (Figure 4a). Axis 1 correlated with all three
environmental variables (Spearman rank order correlations: understorey cover, R =
0.53, P < 0.0001; understorey plant species richness, R = −0.33, P = 0.009; tree
density, R = −0.4, P = 0.0013). The second axis correlated only with understorey
plant species richness (R = 0.37, P < 0.004). There appears to be no taxonomic
pattern underlying the ordination diagram of butterfly species (Figure 4b). Qualitative
comparison of transect and species ordinations (Figure 4a,b) identifies certain species
(labeled on Figure 4b) that are most closely associated with each of the three sites.
As expected, these species correspond closely with those listed in Table 4.

Discussion

Constraints on the design of the sampling regime limit the inferences that might be
drawn on the general impact of disturbance by logging on dry forest butterfly commu-
nities. However, while it is not possible to prove that differences in butterfly compo-
sition across sites are a direct consequence of their disturbance histories, the Mantel
test does reveal inherent differences between sites. Disturbance history remains the
most obvious explanation for these differences. Based on this analysis, the discussion
proceeds on the implication of the Mantel test results that the sampling transects
represent true replicates for disturbance regime at HKK.
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Figure 4. (a) DECORANA ordination showing clear differentiation of the butterfly communities across
the three sites. Axis 1 is correlated with understorey cover, understorey plant species richness and tree den-
sity. Axis 2 is correlated with understorey plant species richness. (b) DECORANA ordination of butterfly
species showing no taxonomic pattern underlying the ordination diagram of butterfly species. Comparison
with Figure 4(a) allows identification of species (labeled) that are most closely associated with each of the
three sites.
534

Species richness of butterflies at HKK (a total of 53 species recorded during the

period of study) appears low when compared to some tropical moist forest sites where
butterflies were recorded at disturbed and undisturbed sites over broadly similar du-
rations and using similar techniques (e.g. Spitzer et al. 1993, 117 species; Spitzer
et al. 1997, 64 species; Wood and Gillman 1998, 80 species), but higher than other
disturbed or insular tropical moist forests (e.g. Hamer et al. 1997, 50 species; Hill
et al. 1995, 41 species). Compared to wet tropical sites, seasonally dry forests such as
HKK are thought to be relatively species poor in plants, vertebrates, and invertebrates
(Nanakorn 1993). Indeed, within Thailand the DDF is species poor even compared
to other dry forest formations (Nanakorn 1993). The total number of butterfly species
(excluding Lyceneidae and Hesperiidae) expected in western Thailand in all habitat
types is 297 (from Pinratana 1979, 1983, 1988, 1992), and the sample reported here
comprises 18% of that. The cumulative species richness curves at all three sites were
approaching an asymptote by the 20th transect, thus 53 species can be taken to be a
good estimate of the total number of species occurring in dry deciduous dipterocarp
forests at HKK at this time of year. A 5 week sampling period minimized multiple-site
comparison errors resulting from seasonal changes in the composition of the butterfly
fauna, but also limited the study’s capacity to estimate total butterfly richness in HKK,
and undoubtedly the species count would have been higher had survey period been
extended, particularly into the wet season.
Although log-normal and log-series models do not necessarily reflect biological
conditions, log-series models imply a structurally simplified habitat whose biological
communities are dominated by a single limiting resource, the largest fraction of which
is taken by few very abundant species (May 1975; Putman 1994; Hill and Hamer
1998). Forest structural complexity and a diversity of vegetation forms have certainly
been shown to be correlated with animal and insect species diversity (James and
Wamer 1982; Erdelen 1984; Gardner et al. 1995). The increased dominance of the
forest understorey at D by a single plant species (Eu. odorata, Table 1) is likely to
favour few butterfly species and may have contributed to the shift from a wholly log-
normal distribution at U to log-series distributions as observed at M and D. Hill et al.
(1995) suggested that species abundance distributions of butterflies might be used
as indicators of forest disturbance, but in this case log-normal and log-series mod-
els alone are not sufficiently sensitive to be useful disturbance indicators for sites at
HKK, as significant changes in diversity were observed across sites yet a log-normal
distribution could be fit to butterflies at all sites.
Spitzer et al. (1993), Hill et al. (1995), Hamer et al. (1997) and Wood and Gillman
(1998) using similar censusing methods recorded higher butterfly species number,
diversity, and abundance in ruderal and disturbed sites than in closed-canopy undis-
turbed sites. On the other hand, Willott et al. (2000) obtained results similar to those
reported from HKK in that diversity declined with increasing disturbance. Hamer
and Hill (2000) point out that contrasting responses of butterfly communities to dis-
turbance are a function of the scale of the sampling regime, such that a decrease in
 535

diversity is recorded where the scale of the sampling regime is large (transect lengths
exceed 100 m) and increased diversity is recorded where sampling is conducted on a
small scale. The results reported here support this pattern, although the recent study
by Willott et al. (2000) using 50 m transects, and not included in Hamer and Hill’s
review, does not.
The risk of underestimating species richness and butterfly abundance in the un-
disturbed forest canopy using the transect method, as pointed out by DeVries et al.
(1997), are likely to be less important at HKK where observations of butterflies in
both the understorey and canopy are relatively easy due to low tree density and stat-
ure, and an open canopy structure. This is supported by the asymptotic leveling of
the cumulative species curve at each of the sites after only a few transects (Figure 1),
as well as the absence of significant differences in the proportions of unidentified
butterflies in transects from the three sites. The problems of identifying butterflies to
species on transects (Walpole and Sheldon 1999, working in moist tropical forest) are
also minimized by an open forest structure, and were in any case largely overcome
at HKK by supplementing the transect method with kite netting. Declining diversity
with increasing disturbance at HKK is therefore unlikely to be a function of sampling
method but rather a reflection of a real result with underlying biological explanations.
The naturally open undisturbed forest formations at HKK appear to support butter-
fly species that are more commonly associated with ruderal habitats, e.g. Neptis spp.,
Precis lemonias, Pr. atlites, Catopsilia pomona and Euthalia pardalis, which may raise
any diversity measure. Widely ranging heliophilous generalist species typical of ruder-
al habitats are most likely to successfully establish in naturally open forest than in closed
canopy evergreen forest. Thus species such as C. pomona (range score 4), E. hecabe
(range score 4), Pr. atlites (range score 3), Pr. lemonias (range score 4), and Danaus gen-
utia (range score 5) that were recorded by Spitzer et al. (1993) only in ruderal and tran-
sitional habitats were relatively abundant in undisturbed forest at HKK. These species
contribute to the relatively high diversity and richness values in the undisturbed site at
HKK, but are unlikely to do so in undisturbed moist forest elsewhere. Transect ordina-
tion shows that butterfly community structure is, nevertheless, influenced by habitat
characteristics at HKK which are a function of habitat disturbance. Principal among
these are understorey cover and tree density, which are themselves closely correlated.
Widespread invasion of disturbed forest by a few understorey plant species, as occurred
at HKK, is a widely observed phenomenon in tropical forests that leads to a relative ho-
mogenisation of the forest habitat at small scales (Gardner et al. 1995). Dominant
invasives are likely to favour non-specialist butterfly species, that are best adapted to
benefit from novel resources, at the expense of habitat specialists.
Some forest species that are considered specialists of certain habitats in undis-
turbed forest may actually benefit from increased disturbance. Troides aecus and Atro-
phaneura adamsoni, for example, occur in forest gaps and along riparian corridors
where there is an abundance of vines upon which their caterpillars feed (Pinratana
1992). Disturbance that causes increased canopy openness and light penetration
536

increases the abundance of herbaceous growth and vines, and favours species nor-
mally frequenting treefall gaps and streams. Butterfly distributions are expected to
covary with the distribution of their host plants even at small scales within forest
stands (DeVries 1988; Beccaloni 1997), and changes in stratification and type of
forest vegetation may reflect differences in the composition of butterfly communi-
ties among sites at the generic and family level (Beccaloni 1997). Delias spp. for
example, are typically confined to the forest canopy (Spitzer et al. 1993) but it is
unclear whether their decline at D is linked directly to canopy loss or to reduced
abundance of their host plants (Loranthaceae). Increased density and variety of her-
baceous understorey plants (Table 1), that constitute the larval host plants of most
danaid butterflies, may account for the observed high abundance of many species of
this butterfly family in the disturbed site (see Table 3). However, the results presented
here give little support for constructing generalities at the family level of species
responses to disturbance. The possible exception may be the Satyrinae for which a
marked decline in abundance with increasing disturbance was consistently recorded
for all five species observed. The distribution of Satyrinae matched that of their larval
host plants, the bamboos and grasses, which occurred most frequently at U and (for
the bamboos) hardly at all at D (pers. observ.) where the understorey was dominated
by Eu. odorata. Changing abundance of grasses across sites in the wet season (pers.
observ.) may affect the temporal diversity and abundance of Satyrinae leading to their
increase in the disturbed site at other times of year. Adult Satyrinae have been noted
to be mostly restricted to undisturbed understorey forest habitats and are sensitive
to canopy opening (Spitzer et al. 1993, 1997; Hamer et al. 1997; Hill et al. 1998;
Wood and Gillman 1998), although Willott et al. (2000) reported the converse with
an increase in the abundance of several Satyrinae species in Bornean logged forest.
A potentially confounding variable is the occurrence of vagrants along sampling
transects. Without autecological studies it is not possible to identify vagrants with
certainty. Nevertheless, an impression of the contribution of vagrants to forest butter-
fly communities may be obtained by comparing the occurrence of species that were
recorded very infrequently. A total of eleven species accounted for less than 0.5%
of the total sample at all sites, but only two of these occurred at site U while eight
were recorded at M and five at D (see Table 3). Some or all of these species may
be vagrants and not permanently associated with forest habitats. More vagrants are
certainly expected at D given that this site is the closest to ruderal habitats. How-
ever, both M and U sites are equidistant from these habitats and no difference in
the frequency of vagrants should be expected on the basis of isolation alone. The
occurrence of vagrants is normally considered a sampling difficulty that confounds
descriptions of habitat-community associations, and it is possible that the long tails on
the rank abundance plots (Figure 2) for M and D are due to vagrants, though further
autecological studies are needed to confirm such a conclusion.
The conservation value of a biological community is determined not only by
its richness and diversity, but also by the rarity and endemicity of its constituent
 537

Figure 5. Species overlap among the three sites illustrated by a Venn diagram representing the number of
species recorded.

species, and the ability of species to maintain viable populations in the face of distur-
bance pressure. Thus while there is considerable species overlap among the three sites
(Figure 5) (the disturbed site alone contains 70% of the species encountered in all
three sites), this simple comparison does not include a measure of the rarity or geo-
graphic range of individual species. At HKK, the species recorded least frequently
tend to be those with small geographic ranges (Figure 3). A large species range has
been shown to correlate, at least for British butterflies, with a number of ecological
variables but particularly dispersal capacity and the utilisation of a wide variety of
ubiquitous hostplants, while a narrow range is associated with low migration indices
and sparse and limited hostplant resources (Dennis et al. 2000). Dominance of the
disturbed site forest understorey by Eu. odorata would therefore seem to general-
ly favour butterfly species with large geographic ranges. These data lend support
to the hypothesis that the rarest and most vulnerable species are also the species
with restricted distributions (Thomas 1991; Spitzer et al. 1993; Hill et al. 1995;
Hamer et al. 1997; but see Willott et al. 2000) and the loss of these species from
538

human-modified habitats will lead to increasing regional homogeneity of the butterfly

fauna.
This study has shown that for a dry forest site in western Thailand even moder-
ate anthropogenic disturbance is sufficient to cause a loss in overall diversity of the
dry season butterfly community (Table 2) and changes to its community composition
(Figure 4a). These changes are highly correlated with forest attributes such as plant
understorey cover and richness which in turn are a function of the reduced tree density
by logging. Although no single species, genus or family can be confidently used as
an indicator of disturbance impacts on community structure a set of species from
several families might be identified as a potentially useful indicator assemblage. The
use of overall butterfly diversity or abundance as a surrogate for diversity of other
taxa should be questioned, given that species responses even within butterfly fam-
ilies can be highly divergent. Likewise, while surrogate taxa might be informative
in some habitat types (e.g. rainforest) and at some sites, they may be misleading
in other habitats such as dry forests, where species associated with ruderal habitats
can persist in characteristically open forest formations. Thus, high diversity at U is
simply a reflection of the occurrence of ruderal species in the open and deciduous
forest habitat at HKK and diversity alone is insufficient to assess conservation value.
More important for conservation is the assessment of the status of butterflies that
are confined to certain forest habitats, and of butterflies that have restricted-range
distributions. If forest disturbance results in a loss of these species then both richness
and heterogeneity of the regional butterfly community will diminish, even though
diversity may increase at local scales (as observed by Spitzer et al. 1993, 1997; Hamer
et al. 1997; and Willott et al. 2000). Finally, this study and others like it, are strictly
relevant only to the areas and seasons in which they were conducted. The response
of butterfly community diversity to anthropogenic disturbance is dependent on the
type of forest formation and on the nature and history of the disturbance, and is a
function of the butterfly community itself which will vary with season. Generaliz-
ing from any single study is therefore inappropriate, but there is, nevertheless, an
urgent requirement for reliable generalizations in planning conservation strategies.
The accumulation of a small number of studies on the impacts of logging on butterfly
communities have highlighted the often divergent responses of butterfly families to
similar disturbance regimes in different geographic areas, as well as the sensitivity of
the response of whole community structure to local site conditions, habitat type and
the scale of assessment.

Acknowledgements

I am grateful for the permission, support and encouragement received from Mr Chat-
chewan Pitdamkhan, Mr Preecha and the staff of Huay Kha Khaeng Wildlife
Sanctuary throughout our stay at the Sanctuary. Dr Chaweewan Hutacharern and
 539

Kowit Chaisurisri of the Royal Forest Department and Jim Coles at the ASEAN
Forest Tree Seed Centre provided much support for this project. Field research was
conducted with the assistance of Katharine Liston and Andy Macgregor. Advice on
community ordination analyses was provided by Alan Morton. Francis Gilbert, Anne
Magurran and Jane Hill commented on an earlier draft of the manuscript. Permission
to conduct research in Thailand was granted by the National Research Council of
Thailand. This paper describes research conducted as part of the CIFOR Conserva-
tion of Forest Genetic Resources programme in collaboration with the Royal Forest
Department in Thailand and the Natural History Museum in London.

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