FOREST USE AND VERTICAL STRATIFICATION IN FRUIT-

FEEDING BUTTERFLIES OF SIBERUT, INDONESIA

Master thesis

by

Chung-Lim LUK
born 26 January 1983
in Hong Kong

Thesis submitted to the Faculty of Biology, Georg-August-Universität Göttingen, in

partial fulfilment of the requirements for the integrated bi-national degree

MASTER OF SCIENCE / MASTER OF INTERNATIONAL NATURE

CONSERVATION (M.SC. / M.I.N.C.)

of Georg-August-Universität Göttingen; Germany and Lincoln University, New Zealand

October 2009
Supervisor/Betreuer:
Examiner/Gutachter:
Date of Submission/Abgabedatum:
German Title/Deutscher Titel:
Dr. Matthias Waltert
Prof. Dr. Michael Mühlenberg
30. Oct 2009
Vertikale und horizontale Einnischung
bei fruchtsaugenden Schmetterlingen der
Insel Siberut, Indonesien
Acknowledgments

This study was part of the research supported by the Siberut Conservation Project (SCP).
SCP is managed by through a collaboration between the German Primate Centre (DPZ)
of Geog-August-Universität Goettingen and Bogor Agricultural University (IPB).
Funding of this project was kindly granted by SCP. Academic backstopping of this
research was carried out mainly by the Centre for Natural Conservation of Geog-August-
Universität.

First I would like to thank my supervisor Dr. Matthias Waltert, from the Centre for
Nature Conservation, for the continuous help, research ideas and academic support.

I would like to thank Prof. Keith Hodges and Dr. Thomas Ziegler from the Department of
Reproductive Biology at DPZ, giving me the great opportunity to experience the unique
Mentawai Forest, not least also for managing all the necessary documents and the
financial support.

On the Indonesia side I would like to thank Dr. Mudhammed Agil, for issuing all the
necessary documents in Indonesia. Thanks Pak Dodo and Aminah for the help of
shopping research equipment and for helping with financial issues in Padang.

For help in the field I would like to thank my guide Pak Gerson; special thanks also to
Dodo, Pak Tasan, Ai and “the Pungut angles” Lia, Ayu, for their “extra help”. Also I
would like to thank Dr. Christope Abegg and Pak Johan for managing many issues for me
in the project.

For identification I would like to thank the staff from The Indonesian Institute of
Sciences LIPI, especially Ibu Peggie, and Dr. R.I. Vane-Wright, from Natural History
Museum of United Kingdom.

I also would like to thank all the staff at the Centre for Nature Conservation, for
facilitating my time in Germany

Finally I would like to give my biggest thank to my mother, Tak-Mei Wong, for her love,
care and support. Without this generous and understanding lady my scientific career
would have ended already 3 years ago.
Table of Content
1 Introduction 1-3

2 Objectives 3

3 Study area and methods 4-11

3.1 Study area 4
3.2 Vegetation data 7
3.3 Butterfly trapping 7
3.4 Identification 8
3.5 Statistical analysis 9

4 Results 12-27
4.1 Vegetation structure 12
4.2 General abundance and diversity 13
4.3 Differences between forest types 15
4.3.1 Taxonomical differences between forest sites 18
4.3.2 Differences between forest sites at species level 20
4.4 Vertical stratification 21
4.4.1 Taxonomical differences among vertical height levels 24
4.4.2 Vertical stratification at species level 26

5 Discussion 28-37
5.1 General 28
5.2 Furit-feeding butterfly response to disturbance 29
5.3 Pattern of fruit-feeding buuterfly stratification 32
36

6 Conclusion 38-39

7 References 40-44

Appendices 45-48
Appendix 1 Distribution information and wing size of sampled butterfly
species 45
Appendix 2 Timetable of fruit-feeding butterfly sampling 46
Appendix 3 Photographic section 47
List of Table
Table Content Page

Table 1 Total number of trees enumerated, tree density and mean basal area,
of overstorey and understorey trees, in defined 10m x 10m quadrats
in natural and disturbed forest 12

Table 2 Individual numbers of Nymphalidae species trapped in three different

heights,
and understorey on forest types 14

Table 3 Total number of individuals, species richness, and diversity parameters of

fruit-feeding-butterflies trapped in two forest types 15

Table 4 Individual numbers of Nymphalidae trapped regarding to subfamily in

natural
forest site and disturbed forest site 18

Table 5 Species with abundances significantly different between disturbed and

natural
forest sites. Statistical test was done by using Mann-Whitney U test 20

Table 6 Numbers of individuals and species richness of fruit-feeding-butterflies

trapped
at three vertical height levels in Siberut, Indonesia 21

Table 7 Individual numbers of Nymphalidae trapped regarding to subfamily in the

three
vertical level: 1m, 15m and 30m 24

Table 8 Species considered as indicator regarding to different vertical level 26

List of Figure
Figure Content Page

Figure 1 Map of the location of the study area in Siberut (extracted from Hadi et
al. 2009),
and of the 14 trap locations in natural and disturbed forest 5

Figure 2 Cumulative number of Nymphalidae species against individual

captured at seven
natural forest and seven disturbed forest sites 16

Figure 3 Cumulative number of Nymphalidae species captured against trap days

(samples)
at seven natural forest and seven forest sites (including two vertical
trapping sites
in each habitat) 16

Figure 4 Multidimensional scaling of fruit-feeding butterfly communities at

understorey trap
of five natural forest and five disturbed forest sites 17

Figure 5 Proportional abundance of butterfly according to subfamily in natural

forest site
and disturbed forest site 19

Figure 6 Cumulative number of Nymphalidae species captured against trap days

at the three different vertical heights along the vertical strata of two
natural forest and disturbed forest sites
22

Figure 7 Cumulative number of Nymphalidae species captured against trap days

at the three
different vertical heights along the vertical strata of two natural forest
and disturbed forest sites 22

Figure 8 Multidimensional scaling of fruit-feeding butterfly communities at

three different trap heights along the vertical strata of two natural forest
and two disturbed forest sites 23

Figure 9 Proportional abundance of butterflies according to subfamily at three

vertical levels:
1m, 15m and 40m 25
1. Introduction

Tropical forests harbouring much of the Earth’s remaining biological diversity, but

are experiencing unprecedented rates of deforestation (Laurance 1999; Brooks et al.

2002). The rainforests of South-east Asia are among the most biologically diverse

areas in the world (Myers et al. 2000). Among the tropical regions South-east Asia

has the highest relative rate of net forest loss and degradation in the humid tropics

(Achard et al. 2002), and it is predicted that it could loose up to three-quarters of its

original forests and almost half of its species by 2100 (Brooks et al. 2002).

Butterflies (order: Lepidoptera) are among the best-studied insect groups in South-

east Asia in terms of taxonomy and biogeography (D’Abrera 1982, 1985, 1986; Aoki

et al. 1982; Tsukada et al. 1985; Tsukada 1991). They are highly sensitive to habitat

disturbance and have been used commonly as an indicator taxon for ecological

research (Kremen 1994; Koh & Sodhi 2004). However, the responses of insects in

general and butterflies in particular to disturbances and deforestation are still

relatively poorly known (Koh 2007). For Siberut Island or the Mentawai islands

example, no previous ecological studies on the responses of butterflies nor on those of

other arthropod communities to disturbances have yet been done.

Siberut and Mentawai are geographically located within Sundaland, which is one of

the 34 ‘biodiversity hotspots’ (Mittermeier et al., 2000), regions which not only

having high levels of biodiversity and endemic species but are also undergoing

immense habitat loss. Siberut is one of the largest islands of the Mentawai Group of

Islands, which remained separated from Sumatra for more than 500,000 years

(Verstappen 1975). Due to this period of biogeographical separation from the

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 1
mainland, the Mentawai Islands consist of relatively high level of endemic species. In

mammals for example, seventeen out of thirty-four species, including all four primate

species, found on Siberut are known to be endemic (WWF 1982). More recent

ecological studies on the floral and fauna of Siberut and Mentawai were primarily

focused on the mega-fauna, especially the four endemic primate species. There has

been no previous extensive ecological study on butterflies or insects published. The

only publication on the butterfly community of Mentawai is mainly focused on

taxonomy and identification (e.g. Aoki et al. 1982, Tsukada et al. 1985 and Tsukada

1991), however the butterfly fauna of the island had been recorded as early as in

Hagen (1893).

Many authors had been describing the changes or differences in the butterfly

community between disturbed and undisturbed sites at understorey level (e.g., for

Asia: Hill et al. 1995; Hamer et al. 1997; Beck and Schulze 2000; Hamer and Hill

2000; Willott et al. 2000; Dumbrell & Hill 2005 for Africa: Kremen 1992, 1994; and

for the Neotropics: Lawton et al. 1998; Barlow et al. 2007). However, as butterflies

are aerial organisms and research is often ground-based, most of these studies did not

achieve to describe the ‘real’ assemblages of butterflies, for which a 3-dimensionally

designed research would be more appropriate, especially when examining differences

in community structure between habitat types.

The vertical structure of tropical rain forests can be described as having distinctly

different vegetation layers (Pomeroy & Service 1986, Schulze et al. 2001; Whitmore

1993). At different vertical heights different abiotic conditions and biotic resources

can be observed (e.g. light, food), which provide a gradient of microhabitat within a

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 2
locality. The analysis of vertical distribution patterns and niche breadths of animals

along such vertical gradients could provide one of the keys to the understanding of

processes underlying species composition in animal communities of multi-layered

forest habitats. Studies on vertical stratification of butterflies have also been

previously conducted in tropical regions (e.g. Fermon et al. 2005; Schulze et al. 2001;

DeVries et al. 1997; Hill et al. 2001).

2 Objectives

In South-east Asia, all fruit-feeding butterflies belong to the Nymphalid subfamilies

Satyrinae, Morphinae, Nymphalinae and Charaxinae (Corbet & Pendlebury 1992).

The aim of this study is to (1) describe the general patterns of vertical stratification in

frugivorous butterflies in Siberut and (2) to use them as an indicator group to analyse

the impact of human disturbance on the forest. This is done by comparing the vertical

distributions, abundance, species richness and diversity of Nymphalidae between

natural and disturbed forests; furthermore, the study intends to (3) increase our

knowledge about the frugivorous butterfly fauna of Siberut island.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 3
3 Methods

3.1 Study area

Siberut is a tropical island, lying at the northern most extremity of the Mentawai

group of islands, at a distance of approximately 130 km off the west coast of Sumatra.

Due to its separation from Sumatra for more than 500,000 years (Verstappen 1975).,

the biodiversity of the Mentawai is quite distinct from the mainland and is

characterised by a high degree of endemism, especially when considering its size

(Whitten 1982a).

Siberut has a total surface area of 4,030 km² and an estimated population of 25,000

people. The natural vegetation in the island is primarily composed of tropical moist

broadleaf forest of the Dipterocarpaceae family (Hadi et al. 2009). However, the

environment is now facing the impact of extensive logging and agricultural farming

(Fuentes 1996/1997). Apart from priamate study, there have been no extensive studies

so far on the ecology of plant or animal communities in Siberut and the Mentawai

islands. Til date no ecological study about butterfly or arthropods can be found. The

only scientific literature about butterflies are on faunistics and taxonomy (Aoki et al.

1982, Tsukada et al. 1985 and Tsukada 1991)

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 4
Figure 1, Map of the location of the study area in Siberut (extracted from Hadi et al. 2009), and of the
14 trap locations in natural and disturbed forest.

This research was based on work carried out at the field station of the Siberut

Conservation Project (SCP). SCP is run by a collaboration between the German

Primate Center (DPZ) and Bogor Agricultural University (IPB). The field station is

located in the northern part of the island, in the Peleonan forest (1o01’34’S,

98o50’16’E; elevation: 8–180 m above sea level). The Peleonan forest is subject to a

long-term international research and conservation program and consists of

approximately 5000 hectares of lowland, mostly primary, lowland forest, but also

coastal and peat swamp forest and tiny local farms (Quinten 2008). Most of the study

area is hilly with elevation measurements ranging from 2-182 m above sea level. The

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 5
forest area is drained by numerous small to medium sized creeks and rivers (Hadi et al.

2009). In total 93 species of trees were recorded in the region, with the tallest tree

sampled was 56m, with 73% of the trees were at the height class of 6 -20m.

Euphorbiaceae, Myrtaceae, Lauraceae and Moraceae were being the most common

tree families found in the site. At species level, species of the genera Mallotus and

Knema, as well as Baccaurea sumatrana were most dominant in all dbh classes .

(Hadi et al. 2009).

Currently, there is an agreement with local people and the Indonesian officials that the

Peleonan forest is being protected for research purpose. At a larger spatial scale, the

forest in Siberut is now facing the threat of commercial logging and establishment of

oil palm plantations. Large areas of the natural forest were already logged and cleared

for wood resources, it is estimated that about 7,000 m³ of timber is exported monthly

(Person 2003). As operations expanded, this Figure is likely to increase in the near

future (Person 2003).

Our trapping sites were situated along the transact, within 2.0km Northeast of the field

station (Figure 1). For the disturbed site chosen, the main cause of anthropological

disturbance at the research site was farming by local people, with banana and cocoa

being the major crops.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 6
3.2 Vegetation data

At each of the fourteen trapping sites, 10 x 10 m vegetation plots were used to assess

the vegetation. Plots were centred around each site. In each quadrat, the diameter at

breast height (dbh) of all trees with a dbh equal to and greater than 5 cm were

recorded. Trees with a dbh of 5-10 cm were defined as understorey trees, and trees

>10cm dbh as overstorey trees. Dbh was converted to basal area [m²/ha], calculated

separately for understorey and overstorey trees to be used as an index of disturbance.

3.3 Butterfly trapping

All trapping activities were conducted in between 6th June and 17th July, 2009.

Cylindrical gauze-traps (Rydon 1964) were used, in which a standard portion of

mashed, fermenting bananas was placed inside the trap to bait butterflies.

All traps were hand-made. Each trap was 30cm in diameter and 60cm long, and

comprised a 30x30cm plastic cardboard base. Each trap was stabilised by 2 rings

(30cm diameter), with 60 cm long white tissue connecting the rings to produce the

cylindrical shape with the bottom open. Nylon strings were used to connected the

bottom of the cylinder and the plastic cardboard, to produce a 3cm gap to allow the

butterflies to enter. The bait was placed in the middle of the cardboard on a small

plate.

Fourteen trapping sites were selected of which seven were in primary forest and seven

were situated in disturbed forest. At four of these sites (two primary forest and two

disturbed forest sites), traps were established at different vertical heights. At each of

these four trapping sites, a tree taller than 30m was chosen in order to suspend traps at

different heights. Traps were installed at ground level (1m), in the midstorey (15 m),

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 7
and the canopy (30 m), resulting in a total of 2 x 2 x 3 = 12 traps. Traps were placed

for 32 non-rainy days and were checked for butterflies and rebaited every 24hours. A

total of 384 trap-days (12 traps x 32 days) were spent to collect fruit-feeding

butterflies.

Out of the other 10 trapping sites, five were in natural forest and fiver in disturbed

forest. At these 10 sites, no traps were placed in the midstorey or canopy, but only at

understorey level (1m):

Traps were left for 16 non-rainy days and were checked, sampled and rebaited every

24hours, with a total of 160 trap-days spent at the 10 understorey sampling sites (5 x 2

x 16).

3.4 Butterfly identification

For field identification, most specimens trapped were preserved and were given a

“specimen ID”. The preserved specimen was identified later in the laboratory of the

Indonesian Institute of Sciences (LIPI). At least three individuals of each species were

preserved for later identification. Trapped butterflies of known species identity, were

identified and released. The identification was mainly based on the publication of

Aoki et al. (1982), Tsukada et al. (1985) and Tsukada (1991)

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 8
3.5 Statistical analysis

Differences in capture frequencies between forest sites were analysed using Mann-

Whitney U-statistics. For the vertical stratification of butterfly community, Kruskal-

Wallis was used to test for differences among the three levels (1m, 15 m, 30m). The

abundance regarding to ‘general’ (the whole nymphalid community), subfamily, and

species were tested regarding the factors habitat type and vertical level. All statistical

tests were done using the software SPSS 13.0 for Windows

The diversity of the butterfly community in each forest type was analysed using

EstimateS 7.52. The species richness estimators, Abundance Based (ACE) and

Incidence Based (ICE) were calculated to estimate the size of local species pool. The

diversity parameters Fisher’s alpha index, Simpson index, Shannon (or Shannon-

Weaver) index and evenness were calculated:

The Shannon or Shannon-Weaver index is calculated as:

Where:

ni is he number of individuals in species i; the abundance of species i.

S is the total number of species.

N is the total number of all individuals

pi is the relative abundance of each species, calculated as the proportion of individuals

of a given species to the total number of individuals in the community

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 9
Evenness is calculated as:

Where:

H' is the number derived from the Shannon diversity index and

H' max is the maximum value of H'

The Simpson’s index is calculated as:

where

S is the number of species

N is the total percentage cover or total number of organisms and n is the percentage

cover of a species or number of organisms of a species.

Note that D therefore ranges from 0 to 1, with 1 representing infinite diversity and 0

representing no diversity.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 10
Fisher’s alpha diversity index is calculated as:

α = N (1 –x ) x

S/N = (1 – x)/ X – ln (x)

Where:

N is the total number of all individuals

S is the species richness

x is estimated by the second solution

Species composition among different vertical strata and between forest types were

grouped using a matrix of dissimilarity (1 - Sorensen Index similarity index) and then

were analysed using multidimensional scaling.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 11
4 Results

4.1 Vegetation

The mean density and basal area of overstorey and understorey trees in natural and

disturbed forest are shown in Table 1. No significant difference was found in basal

area between the forest types, but the standard deviation was quite high (all > 35%)

and basal area of overstorey and understorey trees in natural forest was considerably

higher than in disturbed forest (>50% higher in figure). The density of overstory tree

in natural forest was significantly higher than disturbed forest (Z=-2.47, p=0.013),

however no significant difference was found in the density of understorey trees

between forest type (Table 1).

Table 1. Total number of trees enumerated, tree density and mean basal area, of overstorey and

understorey trees, in defined 10m x 10m quadrats in natural and disturbed forest. Numbers in brackets

represent standard deviation. Statistics from Mann-Whitney U test.

Natural Disturbed
(n=7 plots) (n=7 plots) Z p value
Overstorey trees enumerated 44 32 - -
Understorey trees enumerated 25 19 - -
Overstorey Tree Density [Ind/ha] 628.6 (75.6) 357.1 (190.2) -2.47 0.013
Understorey Tree density [Ind/ha] 457.1 (207.0) 271.4 (197.6) -1.55 0.122 (NS)
Total density [Ind/ha] 1085.7 (177.3) 628.6 (111.3) -3.09 0.002
Overstorey basal area (m²/ha) 49.1 (18.1) 34.5 (20.7) -1.087 0.277 (NS)
Understorey basal area (m²/ha) 1.82 (0.99) 1.12 (0.64) -1.22 0.224 (NS)
Total BA [m²/ha] 51.0 (18.5) 35.6 (20.8) -1.087 0.277 (NS)

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 12
4.2 Overall abundance and diversity

A total of 244 fruit-feeding butterfly individuals belonging to 5 subfamilies

(Biblidinae, Charaxinae, Morphinae, Nymphalinae and Satyrinae), 14 genera, and 20

species were trapped in both natural and disturbed forests (Table 2).

The Satyrinae showed the highest abundance which contributed to168 individuals, or

69% of total trapped individuals. The Biblidinae were the second most abundant

subfamily with 39 individuals (16%), followed by the Morphinae, and the Charaxinae.

The Nymphalinae showed the least abundance with only 10 individuals recorded (4%).

Species abundance

The most common species found were Neorina lowii (n=62), representing about a

quarter of all individuals, followed by Mycalesis maianeas (n=41), Dichorragia

nesimachus (n=34) and Mycalesis oresis (n=32).

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 13
 Table 2. Individual numbers of Nymphalidae species trapped in three different heights, and understorey

on two forest types in Pungut, Siberut, Inodnesia. Trapping was done for 32 rainless days with two

replicates for vertical traps, and 16 rainless days, with five replicates for understorey traps. Trapping

was being held in between June and July, 2009.

The species with an “E” meaning the species is an endemic species to Mentawai Islands.

Mycalesis maianeas 16 9 12 4 41

Mycalesis oresis 2 Natural forest 1 12Disturbed forest 1

Natural Dist16
urbed 32

Neorina lowii 27
1m 151m 30m 6
1m 154m 30m 13understorey
11 T62
otal

Subfamily Total
Species 57 18 11 49 13 9 46 41 244

Biblidinae Dichorragia nesimachus 3 12 5 2 5 5 2 34

Lexias dirtea 2 3 5

Charaxinae Charaxes bernardus 1 1 2

Prothoe franck 5 1 3 1 10

Morphinae Amathuxidia amythaon 2 1 5 1 2 1 12

Discophora spp 1 1

Zeuxidia amethystus 2 2

Nymphalinae Bassarona dunya 2 2 4

Bassarona teuta 2 1 3

Lebadea Martha 1 1

Tanaecia spp. 1 1

Tanaecia visandra E 1 1

Satyrinae Elymnias nelsoni E 1 2 1 1 2 7

Elymnias panthera 2 1 1 4 2 10

Elymnias spp. 1 2 1 4

Melanitis phedima 8 1 1 1 11

Melanitis zitenius 1 1

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 14
4.3 Differences between forest types

Overall, numbers of trapped individuals were similar between natural forest (n=132)

and disturbed forest (n=112) (Table 3, Chi-square test, p=0.200, χ²=1.639, df=1).

Table 3: Total number of individuals, species richness, and diversity parameters of fruit-feeding-

butterflies trapped in two forest types in Siberut, Indonesia, between June and July, 2009. Diversity

indices and estimators of total species richness (ACE, ICE) were calculated with 272 samples ((vertical

traps: 2 reps x 3 levels x 32 days) + (understorey traps: 5 traps x 16 days)).

Natural forest Disturbed forest χ² p-value

Individuals 132 112 1.639 0.20

Species richness 15 15

Fisher's alpha Index 4.35 4.65

Simpson Index 5.52 7.83

Shannon Index 2.02 2.25

Evenness (E) 0.75 0.81

ACE 16.75 18.22

ICE 16.72 17.68

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 15
 16

14

12

Species 10

8 natural
6
disturbed
4

0
0 30 60 90 120
Individuals

Figure 2. Cumulative number of Nymphalidae species against individual captured at seven natural

forest and seven disturbed forest sites ((vertical traps: 2 reps x 3 levels x 32 days) + (understorey traps

5 traps x 16 days)) in Siberut, Indonesia.

16

14

12

10
Species

6 natural
4 disturbed
2

0
0 5 10 15 20 25 30 35
Samples

Figure 3. Cumulative number of Nymphalidae species captured against trap days (samples) at seven

natural forest and seven forest sites (including two vertical trapping sites in each habitat) in Siberut,

Indonesia.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 16
Figure 4. Multidimensional scaling of fruit-feeding butterfly communities at understorey trap of five

natural forest (N) and five disturbed forest (D) sites. Scaling is based on Sorensen similarity values.

Sites of the same habitat type are connected by lines.

Mainly due to the three common natural forest species Dichorragia nesimachus,

Mycalesis maianeas, and Neorina lowii (Table 2), disturbed forest showed slightly

higher values in all diversity and species richness indices. However, total fruit-feeding

butterfly species richness was similar in natural (15 species) and disturbed forest (16

species). This was also reflected in the Evenness index, indicating the distribution of

of individuals to species was more even in disturbed than in natural forest (0.81 vs

0.75). The two species richness estimators, ACE and ICE, indicated species pools of

around 18 in natural or 17 in disturbed forest.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 17
4.3.1 Taxonomical differences between forest sites

Among the five subfamilies recorded in this study, three, namely Biblidinae,

Charaxinae and Satyrinae, were found in very highly significantly different

abundances between natural and disturbed forest. Biblidinae (Table 4, n=39, Z=-3.67,

p<0.001) and Charaxinae (n=12, Z=-2.966, p<0.001) showed a higher abundance in

natural forest than in disturbed forest. Satyrinae (n=168, Z=-4.856, p<0.001) showed

opposite results, having a higher abundance in disturbed forest (Table 4). Furthermore,

Nymphalidae tended to be more common in natural forest, but this was only nearly

significant (Table 4). A MANOVA with the two-dimensional scores resulting from

multi-dimensional scaling did not reveal a significant difference in species

composition of understorey traps between habitat types (F = 1.788; df = 2, 7; p=0.236,

Figure 4).

Table 4. Individual numbers of Nymphalidae trapped regarding to subfamily in natural forest site and

disturbed forest site in Pungut, Siberut, Indonesia, between June and July, 2009. Statistical test was

done by using Mann-Whitney U test, which the p=0.05 is defined as significance level.

No. of spp. natural forest disturbed forest total Z p-value

Biblidinae 2 30 9 39 -3.637 <0.001
Charaxinae 2 8 4 12 -2.966 0.003
Morphinae 3 7 8 15 -0.382 0.702 (NS)
Nymphalinae 5 8 2 10 -1.936 0.053(NS)
Satyrinae 8 79 89 168 -4.856 <0.001
Total 20 132 122 244

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 18
 100%

80%

60%
Satyrinae

Nymphalinae
40%
Morphinae

Charaxinae
20%
Biblidinae

0%
Natural (n = 132) Disturbed (n= 112)

Figure 5. Proportional abundance of butterfly according to subfamily in natural forest site and disturbed

forest site, in Pungut, Siberut, Inodnesia.

In general, Satyrinae was the dominant subfamily, which contributed 60.0% (n=79)

and 67.4%(n=89) to butterflies found in natural and disturbed forest respectively

(Figure 5). Bilblidinae were the second most abundant subfamily in both forest types,

contributing to 22.3% (n=30) of the natural forest butterflies, but represented only

6.8% (n=9) of the individuals in disturbed forest.The other three (less abundant)

subfamilies Charaxinae, Nymphalinae and Morphinae, represented the remaining

individuals (total 17.5% in natural forest and 12.5% in disturbed forest) of the

Nyphalid count.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 19
4.3.2 Differences between forest sites at species level

Of the 20 Nymphalidae species sampled, only four species were found to have a

significantly different abundance between forest types (Mann-Whitney U test for

species with at least 7 individuals, Table 5). Results wer highly significant (p<0.001)

for all four species: Dichorragia nesimachus (Z=-3.037), Mycalesis maianeas (Z=-

4.231) and Neorina lowii showed significantly higher numbers of individuals in

natural forest (n=25, n= 28 & n=41, respectively) than in disturbed forest (n=9, n=13

& n=21, respectively), whereas Mycalesis oresis (Z=-5.259) showed the opposite

pattern (Table 5).

Table 5: Species with abundances significantly different between disturbed and natural forest sites.

Statistical test was done by using Mann-Whitney U test, with 0.05 as significance level. Only species

with total abundance number >7 were tested.

Natural forest Disturbed forest Total p-value Z

Dichorragia nesimachus 25 9 34 <0.001 -3.037

Mycalesis maianeas 28 13 41 <0.001 -4.231

Mycalesis oresis 4 28 32 <0.001 -3.439

Neorina lowii 41 21 62 <0.001 -5.259

Given that these distribution patterns could be potentially biased due to the

combination of possible shifts in vertical stratifiation and the inclusion of understorey

trapping sites, an analysis was done excluding the data from the understorey trapping

sites: Distributions tested by a Chi-Square test were similar and also significant for

Dichorragia nesimachus (χ²=6.259, df=1, p=0.01), Mycalesis oresis (χ²=5.400, df=1,

p=0.020) and Neorina lowii (8.526, df=1, p=0.004). For Mycalesis maianeas, the

distribution was also similar but not significant at the 5% level (1.960, df=1, p=0.162).

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 20
4.4 Vertical stratification

Abundance was generally highest at trap height 1m which amounted to 67.5%, or 106

out of the total of 157 individuals sampled at the four main sites where vertical

stratification was studied (Table 6). Regarding to the butterflies trapped at 15m and

30m, 31 (19.7%) and 20(12.7%) individuals were found respectively, with no

significant difference between these two levels (Table 6).

Table 6: Numbers of individuals and species richness of fruit-feeding-butterflies trapped at three

vertical height levels in Siberut, Indonesia, between June and July, 2009. Statistical test was done by

using the Kruskal-Wallis. Different letters indicate significant differences based on Mann-Whitney U

statistics. All test with p=0.05 as significance level.

1m 15m 30m p-value

Individuals Total 106 a 31 b 20 b <0.001

Natural Forest 57 18 11

Disturbed Forest 49 13 9

Species Total 14 8 10

Natural Forest 7 5 6

Disturbed Forest 12 6 5

The distribution of numbers of individuals per height level (Table 6) did not differ

between natural and disturbed forest (χ2=0.352, df=2, p=0.839). Likewise, the

distribution of the numbers of species did not differ significantly between height

levels. Furthermore, there was no difference in vertical height distributions between

natural and disturbed forest, although disturbed forest seemingly had an elevated

number of species at 1m level compared to natural forest.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 21
 16

14

12
Species 10

6
1m
4
15m
2 30m
0
0 5 10 15 20 25 30 35
Trap days

Figure 6. Cumulative number of Nymphalidae species captured against trap days at the three different

vertical heights along the vertical strata of two natural forest and disturbed forest sites in Siberut,

Indonesia.

16

14

12

10
Species

6 1m
4 15m
2
30m

0
0 30 60 90 120
Individuals

Figure 7. Cumulative number of Nymphalidae species against number of individuals captured at the

three different vertical heights along the vertical strata of two natural forest and disturbed forest sites in

Siberut, Indonesia.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 22
Figure 8. Multidimensional scaling of fruit-feeding butterfly communities at three different trap heights

along the vertical strata of two natural forest (N) and two disturbed forest (D) sites. Scaling is based on

Sorensen similarity values. Sites of similar trap heights are connected by lines.

Out of the 20 fruit-feeding butterfly species sampled in the study, 70% or 14 species

were found at the lowest level. At the midstorey level, only 8 species were found,

while 10 species were recorded at the canopy level (Table 6, Figure 6). A species

accumulation curve indicated that species sampling at 1m level was fairly complete,

reaching a plateau but that at 15m and especially at 30m level species were still added

at the end of the study period (Figure 7).

Figure 8 shows results from the.multidimensional scaling of fruit-feeding butterfly

communities at three different trap heights along the vertical strata of natural forest

and disturbed forest sites. The fruit-feeding butterfly communities of both natural

forest and disturbed forest along the stratum did not show a very clear distinction,
Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 23
when examining results visually. Also, a MANOVA of the two-dimensional scores

from multi-dimensional scaling showed no significant differences in species

composition among the vertical stratum (F=2.03; df=4, 10; p=0.165) and between

habitat type (F = 1.75; df = 2, 5; p00.265), based on the four vertical trapping sites.

4.4.1 Taxonomical differences among vertical height levels

Of the five subfamilies, only Biblidinae and Satyrinae showed significantly different

vertical distributions (Table 7; Kruskal-Wallis test, p<0.021 and <0.001, respectively).

While most Bibilidinae (2 spp.) were trapped at midstorey levels, Satyrinae were

mainly trapped at understorey level (Table 7).

Table 7. Individual numbers of Nymphalidae trapped regarding to subfamily in the three vertical level:

1m, 15m and 30m, in Pungut, Siberut, Inodnesia, between June and July, 2009. Statistical test was

done using Kruskal-Wallis Test, which the p=0.05 is defined as significance level.

No. of
spp. 1m 15m 30m total p-value
Biblidinae 2b 7 17 a 5a 29 0.021
Charaxinae 2 6 1 1 8 0.211
Morphinae 3 6 3 1 10 0.061
Nymphalinae 5 2 3 2 7 0.051
Satyrinae 8a 85 7b 11 b 103 <0.001
Total 20 106 31 20 157

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 24
 100%

80%

60% Satyrinae

Nymphalinae
40% Morphinae

Charaxinae
20%
Biblidinae

0%
1m (n = 106) 15m (n = 31) 30m (n = 20)

Figure 9. Proportional abundance of butterflies according to subfamily at three vertical levels: 1m, 15m

and 40m, in Pungut, Siberut, Inodnesia.

None of the subfamilies had highest numbers of individuals at height 30m. For

Charaxinae, Mophinae, and Satyrinae, highest number of individuals were found at

1m, which contributed to 75%, 60% and 82.5% respectively (Figure 9). Two of the

subfamilies were found to be the most abundant at the mid-storey level:

At this vertical level, Biblidinae showed 58% and Nymphalinae showed 42% of the

total abundance as compared to the remaining three subfamilies.

At the lowest (understorey) level, Satyrinae were the dominant subfamily,

contributing to 80.2% (n=85) of all fruit-feeding butterfly individuals sampled, with

the remaining 20% represented by the remaining four sub-families (Figure 4). At the

mid-storey level, Biblidinae were the most abundant contributing to 54.8% (n=17) of

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 25
 the total 31 individuals sampled. Satyrinae were second in abundance (22.6%, n=7),

followed by Morphinae and Nymphalinae (both 9.7%. n = 3). At the highest (canopy)

level, Satyrinae and Biblidinae were the two most dominant subfamilies, with 55.0%

(n = 11) and 25.0% (n = 5) of individuals (Figure 4).

4.4.2 Vertical stratification at species level

For species with a total count higher than 10, Kruskal-Wallis test was done to test if

there was any difference in abundance amongst the three vertical levels. Only two

species showed significant results (Table 8). Dichorragia nesimachus showed

significant higher abundance in 15m (n=17) and 30m (n=5) than in 1m (n=5)

(p=0.009). Although at 1m and 30m the species had the same number of individuals,

due to the reason of statistics and the butterfly sampled in different locality (i.e.

different combination of sample trapped in different replicate, but both height level

had five individuals in total), a significant higher result was found at 30m than at 1m.

Table 8 Species considered as indicator regarding to different vertical level. Statistical test was done by

using Kruskal-Wallis test, with 0.05 as significant level. Only species with total abundance number > 9

were tested. Abundance and median are based on total butterfly sampled in 32

p-
1m 15m 30m Total value
abundance median range abundance median range abundance median range
Dichorragia
nesimachus 5b 1.5 1-2 17 a 2.5 2-10 5a 0 0-5 27 0.009
Mycalesis
maianeas 25 a 5.5 1-13 0b / / 0b / / 25 0.005
Neorina lowii 33 a 5 0-23 5b 1.5 0-2 0 / / 38 <0.001

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 26
For Neorina lowii, a significantly higher abundance was found at 1m than at 15m

(n=33 & n=5 respectively) (Mann-Whitney U test, p<0.001). At level 30m no

individuals of this species were trapped. All twenty-five individuals of Mycalesis

maianeas collected were found at level 1m, being significantly different from the

other vertical levels (p=0.005, Table 8).

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 27
5 Discussion

5.1 General

An overall capture rate of 0.45 fruit-feeding butterflies per day per trap was recorded

in this study. This figure is relatively low when compared to other studies in tropical

regions. For example Devries and Walla (2001) found an average number of baited

nymphalids per day and per trap of 0.8 in floodplain forest in Ecuador. Hill et al.

(2001) had nymphalid capture rates of 0.9 individuals per trap day in an unlogged

forest plot in Sabah. A capture rate of 2.9 nymphalids was reported by Shahabuddin

and Terborgh (1999), from a set of forested islands in Venezuela. A very high capture

rate of 8.3 was reported by Fermon (2002), in secondary moist semi-deciduous forest

of Côte d’Ivoire.

A total of 244 fruit-feeding butterflies belonging to five sub-families (Biblidinae,

Charaxinae, Morphinae, Nymphalinae and Satyrinae) and 20 species were recorded

during this study (Table 2). The species richness recorded was relatively low when

compared to other vertical stratification studies on fruit feeding butterfly communities

in SE Asia region: Sulawesi (Fermon et al. 2005: 43 species) and Borneo (Schulze et

al. 2001: 53 species) in particular.

Out of the 20 species of Nymphalindae recorded in this studied, two were not

previously found in the Mentawai Islands (Discophora spp & Tanaecia spp.). Further

identification will be done to confirm the two unknown species. Tanaecia visandra

and Elymnias nelsoni, were the two species sampled in the study area which are

endemic to Mentawai. The 10% endemism (2 out of 20 species) recorded for the

Nymphalidae family from this study was lower than expected since flora and fauna in

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 28
Siberut have been know to show a high degree of endemism (example: 50% of

mammals are endemic, WWF1982). Due to Mentawai Island being originally

separated from Sumatra, about 500,000 years ago, all non-endemic Nymphalidae

butterfly individuals sampled in the study area are also found in the main island of

Sumatra. The status of endemism of butterfly at species and subspecies level can be

found in the references of Aoki et al. (1982), Tsukada et al. (1985) and Tsukada

(1991).

5.2 Fruit-feeding butterfly response to disturbance

Our vegetation analysis showed that tree densities in natural forest were significantly

higher than in disturbed forest (1185.7 vs 628.6 Ind/ha). In contrast, basal area was

not significantly different between natural and disturbed forest. However the figure of

basal area in natural (total: 51.0 m²/ha, dbh ≥5cm ) forest is general higher than in

disturbed area (35.6 m²/ha). The reason of non-significant result may due to the high

level of variance (Standard derivation >36%) (Table 1). The figures were however

consistent with previous vegetation surveys in Siberut (Hadi, in litt: 58.2m²/ha for dbh

class >= 10cm, also from Peleonan forest; or the Paitan forest: 47m²/ha, dbh>=15cm,

Whitten 1982) Sarawak (Mulu forest: 57m²/ha: Proctor, 1986). Whitten (1982) reports

basal areas from peatswamp forests median = 23 m²/ha, dbh ≥15cm). The basal areas

found in this study therefore might indicate that study sites are largely representative

for conditions of natural and disturbed forests of Siberut.

This study did not reveal major significant differences in nymphalid butterfly

abundance or species richness between natural forest and disturbed forest (Table 3).

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 29
Biblidinae and Charaxinae were the two subfamilies, which showed very significant

higher abundance in natural forest than in disturbed forest (Table 3) while Satyrinae

showed an opposite trend.

Only four species with individuals >7 were found having significant differences

between habitat types. Dichorragia nesimachus, Mycalesis maianeas and Neorina

lowii found significant higher abundance in natural forest. Mycalesis oresis found

opposite trend as significant higher abundance was found in disturbed forest.

These results are coherent to the findings of Fermon (2005), whose study on Sulawesi

can be considered as a template for this study, where the two different Mycalesis spp.

((M. horsfieldi & M. itys). showed significantly also showed higher abundances in

natural forest.

Dichorragia nesimachus was also recorded in Fermon (2005), showing a similar trend.

There was no similar species or genera of Neorina lowii found in Sulawesi in previous

scientific record.

Concerning diversity and species richness, similar results were found between habitats

(Table 3). Findings on the effect of tropical forest disturbance on butterfly species

richness and diversity among recent studies show mixed results. Butterflies in SE Asia

for example, resulted in no differences (Hill et al 2001, in Borneo), higher (Hill et al

1995, in Buru island, Indoesnesia; Beck & Schulze 2000 in Borneo) and lower

( Hamer et al. 1997, in Sumba island, Indonesia; Willott et al. 2000, in Borneo,

Fermon et al. 2005, Sulawesi) species diversity or/and richness in natural forest than

in disturbed forest.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 30
The spatial scale maybe one of the factors influencing the result of effect of habitat

disturbance on species richness, and this has been widely accepted by most

Lepidopterists. The trend suggested that a smaller spatial scale research tended to

result in a higher species diversity and richness found at disturbed sites, while at

larger spatial scales research showed opposite trend (Koh 2007, Hamer & Hill 2000,

Hill & Hamer 2004).

An example can be seen by combing the survey of Hamer & Hill (2000), and Hill &

Hamer (2004), about the impact of land use change on the butterfly community

sampled. Eleven of 12 studies conducted at spatial scales of < 1ha reported higher

species diversity in disturbed sites, while 10 of 15 studies at larger scale of >3.1ha

reported higher diversity in undisturbed sites. According to the definition of the

Hamer & Hill (2000), this study is categorised as large scale, and did not consistent to

the trend suggested.

Species richness and diversity is usually correlated with habitat diversity. We

expected that in natural forest, more microhabitat types (e.g. canopy, understorey, gap)

could be found, as disturbance produces a more homogeneous vegetation structure

compared to undisturbed forest (Hill et al. 1995, Hamer et al. 2003). So a higher

species diversity and richness should be found if the spatial sampling scale was large

enough to cover the entire habitat. However, disturbance is also known to create

opportunities and different niches for additional species, not found in undisturbed

forest (Connell 1978), by opening up areas of closed-canopy dense forest as well as

creating light gaps. So if spatial scales were small, a higher habitat heterogeneity

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 31
would be found in the sample area resulting in higher species diversity and richness

observation in disturbed site.

Evidence showed that climate might also be a factor influencing the result of effect of

habitat disturbance on species richness. Hamer et al. (2005) conducted a study of the

impact of logging on butterfly community in Bornean forest, where a higher species

diversity and richness was found in primary forest during the dry season, while an

opposite trend was recorded during the wet season. The cause was attributed to the

immature stages of the butterfly being affected by climatic variables. This suggested

that a temporal scale might also be an important factor to be considered while

conducting research on butterfly. The ecology of different taxonomic groups

responding to climate and seasonal abiotic variables would affect the sampling result

in ecological research.

5.3 Pattern of fruit-feeding butterfly stratification

Only few similar studies on vertical stratification have been done in the past. In

general this research was aimed at examining the vertical stratification of nymphalid

butterflies by sampling and considering three different levels: ground, midstorey and

canopy. The result suggested that in general, the abundance was the highest in the

understorey level and significantly higher than at midstorey and canopy levels. In our

study, we could not establish different abundances between midstorey and canopy

levels. This is consistent to most similar study (Example: in Ecuadorian rainforest-

DeVries et al 1999, in North Borneo-Dumbrell & Hill 2005, in Sulawesi- Fermon et

al. 2005, etc).

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 32
The multivariate analysis and MANOVA showed that a stratification of distinct

communities cannot be confirmed (Figure 8). Concerning species diversity,

understorey revealed the highest species numbers (14), while only 8 and 10 species

were recorded at midstorey and canopy level respectively (Table 6, Figure 6 &7).

From examining the species accumulation curve (Figure 6 & 7), it is very likely more

species would be observed at the level of midstorey, and in particular overstorey level,

if the sample size was increased.

The most widely accepted explanation for the relatively higher abundance of

butterflies at understorey level is the availability of food resources like rotting fruits

(Schulze et al. 2001). Fallen rotting fruit on the forest ground provide a better amount

and diversity of favorable food resource to support a higher species diversity and

abundance of fruit-feeding butterfly individuals.

Our results – in line with previous research - showed that midstorey and canopy might

share relatively large number of similar fruit-feeding butterfly species. This is because

at the canopy and midstorey level, availability of fruit at the vertical position is not

stable hence affecting the niche width of the non-understorey fruit-feeder. The niche

shared by the fruit-feeder is more likely to be expanded at vertical level. Since the

food availability is scare, the fruit-feeder above ground level need to forage in

different strata in order to get sufficient food (Schulze et al. 2001).

Fermon et al. (2005) studied vertical structure of fruit-feeing butterfly in Sulawesi,

and found that species diversity was highest at midstorey level, suggesting that at this

level distinct vertical communities overlap. Schulze et al .(2001) found a quite similar

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 33
result of species diversity and richness which understorey and midstorey had the

highest value. Another factor was the difference in canopy height in this research

study as compared to previous studies of Fermon et al (2005) and Schulze et al. (2001)

were sampling was done at canopy level of 50m, while in this study sampling was

done at 30 m. The higher upper canopy level of the two studies compared to this study

also result in a larger spatial extent of what is considered “midstorey” and therefore in

a bigger niche width, because of the much larger space and probability of rotten food

available is higher. Moreover, the higher spatial ratio of midstorey to undstorey size

results in a higher chance of more species to be observed.

In the study of Fermon et al. (2005), all six subfamilies recorded showed significant

correlations between abundance and vertical height (Spearman rank correlation). The

abundance of Charaxinae and Apaturinae was positively correlated with vertical

height, while the other four subfamilies: Satyrinae, Morphinae, Biblidinae and

Nymphalinae showed a negative relationship with vertical height. In our study, where

we used Kruskal-Wallis to test the abundance of the five subfamilies sampled at three

different vertical levels. Charaxinae, Morphinae and Nymphalinae showed no

significant difference among vertical levels but Satyrinae were found being most

abundant at the lowest level, coherent to Fermon (2005) but Biblidinae showed least

abundance at the lowest level in contrast to the findings of Fermon (2005). However it

should be note that the Biblidinae in our study only consist of two species, out of

which one species Dichorragia nesimachus represents 87% of the whole subfamily

(details see Table 1), while the trapped species of Biblidinae in Fermon (2005)

consists of twelve species. In contrast to the results of Schuze et al. (2001) on the

vertical pattern of fruit-feeding butterfly in Bornean forest, the abundance and

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 34
diversity subfamily Satyrinae, Morphinae and Nymphalinae showed a negative

relationship with vertical height, while Charaxinae did not show a significant

difference in abundance among vertical levels.

Only two species showed significant differences in abundance among three vertical

levels. Dichorragia nesimachus was in general found in mid-storey to canopy level,

the selection of the vertical strata of the species was being very similar to the study of

Fermon (2005). Although no statistical difference in abundance could be found in the

Mycalesis maianeas among vertical levels, since it could be only sampled in the

understorey level with 25 individuals and statistical test cannot be performed. This

species could still be considered as an understorey species as this species showed a

similar trend in the studies of Fermon (2005). In previous studies, Charaxes spp. were

frequently found at canopy level, and were thus defined as canopy species (e.g.

Fermon 2002, Fermon et al. 2005, Schulze et al.2001). In the present study, only two

individuals were sampled, one found in each midstorey and canopy level, suggesting

Siberut’s Charaxinae having a behaviour similar to previous studies.

Disturbance is known to modify or influence the vertical stratification of butterfly

communities in tropical forests (e.g. DeVries 1988; Schulze et al . 2001; Fermon et al .

2005). Previous studies showed that the canopy species of natural forest maybe

present in the understorey of disturbed forest. The main suggested reason is that

canopy species are known to treat light gaps and forest edges as canopy (DeVeries

1988). The light gaps present in disturbed sites causing the light preferring canopy

species will be found in the lower stratum (Dichorragia nesimachus in this study

showed this pattern). When considering studies which sampled butterflies only at

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 35
understorey level, species richness or diversity in disturbed forest may have been

inflated to give a bias result. True canopy species in natural forest hence cannot be

sampled, but might be present in the understorey level of disturbed forest, resulting in

(biased) higher species diversity and richness in disturbed forest. In order to study

butterfly communities, a 3-dimentional sampling design should be considered to give

a better description of the community structure.

Fermon (2002) suggested that any species community should reach its minimum

richness and abundance at canopy, which means that more rare species should be

found at canopy level. In the study of Schulze (1995), the result suggested a similar

trend of rare species of fruit-feeding butterfly with respect to increasing stratum

height, but the study also found that nectar-feeding species had an inverse trend with

increasing number of rare species at lower stratum. We could come to a tentative

conclusion that in contrast to nectar-feeing butterflies, fruit –feeding butterfly

species have a lower pressure of resource limitation at lower strata.

5.4 Fruit-feeding butterflies as indicator

The result suggested that Dichorragia nesimachus was the best species to be used as

indicator species since it was responding most prominently to habitat change in our

study area. It was both a common species to be found and was highly sensitive to

disturbance. Concerning vertical stratification, the species was more likely to be found

in midstorey-canopy level in natural forest, but absent in canopy in disturbed forest

suggesting that it might be sensitive to light availability. Mycalesis maianeas,

Mycalesis oresis, and Neorina lowii were also good species in indicating disturbance

since the abundance of these significantly changed between habitats, and were all

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 36
common species in the region. Mycalesis maianeas and Neorina lowii found higher

abundance in natural forest, while Mycalesis oresis showed an opposite trend.

Although some butterfly species show clear response to the change of habitat (e.g.

Charaxes spp. in Schulze et al 2001 and in Fermon et al. 2005), the role of butterflies

as indicators of habitat change needs much detailed study. Lawton et al. (1998)

investigated species richness patterns across a tropical land-use gradient in Cameroon.

The habitats investigated ranged from highly modified grasslands to relatively intact

closed-canopy forest. The results showed that trends in species richness along the

disturbance gradient varied widely for different taxonomic groups, suggesting that

conservation monitoring and assessment based on indicator taxa may not be useful.

Simply counting the number of species provides no information on the composition of

the species sampled. If species richness alone is used as a measure of biological

conservation value, it may be misleading higher abundance or/and species richness

might be found in disturbances site, as it may favour wider dispersal of species (Akite

2008). The focus on well-defined assemblages rather than on selected species captures,

or species richness, gives more information, and more likely to reflect the process of

impacts of fragmentation on ecosystem structure and function (Kitching, 1994)

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 37
6 Conclusion

The forest of Siberut is now facing the threat of commercial logging and agricultural

farming. Studies on the impact of these anthropological changes on biodiversity are of

urgent needed to access conservation concern.

In our study, we compared the fruit feeding butterfly assemblages in natural

undisturbed forest, and disturbed forest modified by agricultural farming. Overall

species abundance and richness showed no significant differences between natural

and disturbed forest. Dichorragia nesimachus, Mycalesis maianeas, Mycalesis oresis

and Neorina lowii were the species which are sensitive to the change of forest type.

Concerning vertical stratification, the understorey showed significantly higher

abundance than the midstorey and canopy level. The species richness and diversity is

the highest at the understorey level but the trend of the indices at the midstorey level

and especially canopy level seem to be still rising. The vertical stratification of

species composition in natural and disturbed forest was not clear, however at family

level two (Biblidinae & Satyrinae) of the five families showed significant differences

in abundance among vertical heights. Satyrinae were dominant at the understorey

level, and compose more than 60% of total number of fruit-feeding butterfly sampled

and might explain the non-significant result of species composition in natural and

disturbed forest.

Butterflies are one of the most ideal taxonomic groups for investigation of habitat

change. The lack of consensus among studies makes it impossible to conclude to what

extent land use change affects butterfly species richness or diversity. The use of

species richness, diversity and species diversity as valuing disturbance impact needs

in-depth studies. Future community level studies should be designed carefully, so as

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 38
to be able to examine the true responses of the assemblage of butterfly communities to

anthropological disturbance of habitat.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 39
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Appendix 1 Distribution information and wing size of sampled butterfly species

Distribution information (based on Aoki et al. 1982, Tsukada et al. 1985 and Tsukada 1991) and wing size of sampled butterfly species.

wing size
family distribution remarks on distribution (mm)
Amathuxidia amythaon Satyridae 1, 2, 3, 5 also found in Philipines (Samar, Leyte, Panon, Mindanao & Negros), Malaya to Burma 43
Bassarona dunya Nymphalindae 1, 2, 3, 4, 5, 6, 29.5
Bassarona teuta Nymphalindae 1, 2, 3, 4, 5, 6, 7 also found in Ruteng (Indonesia), Thailand and Myanmar 23.5
Charaxes bernardus Nymphalindae 1, 2, 3, 5, 6, 8 also found in W. & S. China, Myanmar, Nepal 29
Dichorragia 1, 2, 3, 4, 5, 6, 8,
nesimachus Nymphalindae 10 also found in S. C. W. China, Japan S. Koera, Taiwan and Myanmar 30.5
Discophora sp. Satyridae / 25
Elymnias nelsoni Satyridae 1 23.5
Elymnias panthera Satyridae 1, 2, 3, 5, 6, 8 24.5
Elymnias sp 1 Satyridae 26,5
Lebadea martha Nymphalindae 1, 2, 3, 4, 5, 6, 8 also found in Indo-China region 17.5
Lexias dirtea Nymphalindae 1, 2, 3, 4, 5, 6 also found in Myanmar, Thailand 28
Melanitis phedima Satyridae 1, 2, 3, 5, 6, 10 also found in from W. Malay to India, Sri Lanka, Formosa, Japan, and S., C. & W. China 28
Melanitis zitenius Satyridae 1,2,3,4,5,6,7, 9 also found in S. India, and from W. Malay to S. Burma 31
Mycalesis maianeas Satyridae 1, 2, 3, 5 13
Mycalesis oresis Satyridae 1, 2, 3, 5 also found from Malaya up to Burma 14
Neorina lowii Satyridae 1, 2, 3, 4, 5 39
Prothoe franck Nymphalindae 1, 2, 3, 4, 5, 6, 8 also found in Ruteng (Indonesia) Indo-China region and Myanmar 26
Tanaecia spp. Nymphalindae / 32
Tanaecia visandra Nymphalindae 1 27
Zeuxidia amethystus Satyridae 1, 2, 3, 5 37
Where 1=Mentawai, 2=Sumatra, 3=Borneo, 4=Palawan, 5=W.Malay, 6=Java, 7=Lombok, 8=Bali, 9=Sumbawa, 10=Sulawesi

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 45
Appendix 2 Timetable of fruit-feeding butterfly sampling

The date of furit-feeding butterfly sampling of vertical traps (3 traps x 4 locations, 32

days), and undstorey traps (5 traps x 10 locations, 16 days) in research site of Siberut,
Indonesia. Each day represents 4 trap location of vertical trap and 5 trap locations of
understorey trap sampled. Traps were checked every 24 hours.

Month June July

Day 6 1
7 2
17 3
18 4
19 5
20 6
21 7
22 8
23 9
24 10
25 11
26 12
27 13
29 14
30 15
16
17
Days per month 15 17

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 46
Appendix 3 Photographic section

Dichorragia nesimachus Neorina lowii

Mycalesis maianeas Charaxes bernardus

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 47
The local guide with vertical tap Vertical trap at mid-storey level

Vertical trap at mid-storey level

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 48
Declaration
Hiermit versichere ich gemäß § 9 Abs. 5 der Prüfungsordnung für den integrierten
binationalen Master-Studiengang Internationaler Naturschutz (engl.: International
Nature Conservation) vom 16.08.2006, dass ich die vorliegende Arbeit selbstständig
verfasst und keine anderen als die angegebenen Hilfsmittel verwendet habe. Diese
Arbeit wurde nicht in der gleichen oder einer ähnlichen Form bereits einem anderen
Prüfungsausschuss vorgelegt und wurde bisher noch nicht veröffentlicht.

Hereby I affirm – according to § 9 section 5 of the examination regulations for the

integrated bi-national Master programme International Nature Conservation (deutsch:
Internationaler Natur-schutz) from 16.08.2006 – that I have penned the present thesis
autonomously and that I did not use any other resources than those specified above.
This work was not submitted previously in same or similar form to another
examination committee and was not yet published.

Chung-Lim LUK Forest use and vertical stratification in fruit-feeding butterflies of Siberut, Indonesia 49