SPECIES DIVERSITY OF FRUIT-FEEDING BUTTERFLIES (LEPIDOPTERA:

NYMPHALIDAE) IN SPATIAL, TEMPORAL AND VERTICAL DIMENSIONS, IN

KUBAH NATIONAL PARK, SARAWAK

Christharina S Gintoron

Master of Science

(Entomology)

2011

Faculty of Resource Science and Technology

DECLARATION

I hereby declare that no portion of the work referred to this thesis has been submitted in

support of an application for another degree or qualification to this or any other university or

institute of higher learning.

_________________________

(Christharina S Gintoron)

June 2011

SPECIES DIVERSITY OF FRUIT-FEEDING BUTTERFLIES (LEPIDOPTERA:

NYMPHALIDAE) IN SPATIAL, TEMPORAL AND VERTICAL DIMENSIONS, IN

KUBAH NATIONAL PARK, SARAWAK

CHRISTHARINA S GINTORON

A thesis submitted

in fulfillment of the requirements for the degree of

Master of Science

(Entomology)

Faculty of Resource Science and Technology

UNIVERSITI MALAYSIA SARAWAK

2011

Especially to Daddy and Mummy, thank you for everything

ACKNOWLEDGEMENT

First and foremost, I would like to say my gratefulness to God, for everything I have and

acquired especially during these three years of completing this study. Without any of His

guidance, blessings and grace, obviously I would not be able to accomplish this.

My sincere gratitude also goes to Prof. Dr. Fatimah Abang, for her endless advices, guidance

and support, which has encouraged me this whole time. Saying thank you would not ever be

enough! Also to Prof Dr. Mohd Tajuddin Abdullah, thank you for helping me with my thesis,

with the comments and suggestions. Huge appreciation to all staffs in Kubah National Park,

which may direct and indirectly, been helping me and Irene during our stay there. Thanks for

all the advices, safety guides and warm co-operations. I would also like to thank Sarawak

Forestry Department for granting the Research Permit (No. NCCD.907.4 (IV)-20 & 21) and

Park Permit (No: 22/2009 & 23/2009), and UNIMAS Zamalah Postgraduate scholarship

which had financially aided me during my postgraduate study.

Endless gratitude to Mr. Jalani Mortada, the best shooter ever, without you there would not

be 40 baited traps hung in Kubah National Park. You’re the best!! The same goes to

Muhamad Ikhwan Idris, I will never forget those times when we were still in the forest

checking the baited traps with Irene, and it’s getting dark and raining heavily. I have

promised you that we will sob over this during our graduation day right? Best memory ever!

Thanks also for the statistical analysis advices and software, journals, and for helping me

with my write-up. Bear-hug goes to Irene Christianus, for always being there for me (in the

forest!), spending our field-trips together with cooking and baking, and for always being

brave in the gloomy house. She’s the man!

Big thanks also for Mr. Raymond Atet, who never failed to deal with our transportation to

Kubah National Park, each and every month during our field-trips, and also to Mr. Wahap

Marni, for his companion and experience to survey for suitable tree replicates prior to the

field-sampling. The same goes to Mr. Besar Ketol and Mr. Dominic Nyuko, for being my

guarantor, thank you so much for having such confidence on me. Also to Zulkiflie from Kg.

Telaga Air, thank you for being such a fast learner and willingly helping us.

To my friends, Mohd Hanif Ridzuan and Aizat Japar, thank you so much for the lifts to and

from UNIMAS, you guys had just saved our life there! Also to Pang Sing Tyan, for your

concern while being a big sister to both of me and Irene, and visiting us back there. To Siti

Mariam Jamaludin, thank you for the short tutorial of R-program you have shared with me,

and for being such a good cook in Kubah last time. Now I know how to prepare ‘ayam

kurma’! For being very patient and helpful while reading and correcting my thesis drafts,

endless gratitude to Roberta Chaya Tingga Tawie and Mohd Fizl Sidq Ramji. You guys had

polished my write-up! Same goes to Eric Pui Yong Min and Ridwan Rahman, thank you for

the advices and comments for my thesis. Lastly, to all my friends, especially zoology

postgraduate students, thank you so much for your help and encouragement.

Especially to Johnnatan Farrel Dominic, thank you for always being there, cheering me up.

You have been motivating me always, thank you! To my parents Mr. Saurin Gintoron and

Mrs. Samitha Sompony, this is for you. Without your love and support, undoubtedly I would

not be as what I am now. I love you so much. Also to my sisters Claressa and Lizie, brothers

Benzon and Leo Xandro, I love you guys too. Regards!

Abstract

The overall diversity and distributions of the fruit-feeding butterflies in spatial, temporal and

vertical dimensions were investigated in the mixed-dipterocarp forest of Kubah National

Park, Sarawak, by utilising baited-traps. Fourteen continuous days monthly from 17th

May to

17th

November 2009 of field sampling had resulted to 665 individuals of nymphalids,

representing 49 species and four subfamilies. Bassarona dunya was listed as the most

abundant nymphalid species in this study. Forest edge was found to be sustaining the highest

number of species among the four forest habitats, yet the highest diversity was revealed at the

secondary forest (H’= 3.033, 1-D= 0.933, α= 13.370). In contrast, primary forest sustained

the highest taxonomic diversity of nymphalids, as in the distantly related species. Being

practically ‘sun-lovers’ explained the presence of most nymphalid species at the forest edge,

while with the intermediate disturbance observed in the secondary forest, enormous

microhabitats are offered here as both climax and pioneer tree species coexist together, thus

supporting a diverse nymphalids assemblage. On the other hand, the effects of different

monthly replicates on the nymphalids distribution was observed, as well as accentuating the

influence of rainfall to these forest butterflies. Undeniably, heavy rainfall disrupted the

nymphalids’ occurrence pattern, as apart from being synonym with the sun-rays, pupal and

larval stages of the nymphalids are badly affected. Vertical study on these nymphalids

elucidated a prominent preference towards the ground level, correlated with the presence of

rotting fruits on the ground. Phenomenon such as heliophobia was also hypothesised for B.

dunya, due to its dominance uniquely at the lower level. Larger-forewing nymphalids with

slender body occupied the lower strata, for better maneuverability through the herbaceous

plants. Conversely, in order to acquire strong and rapid flight in the forest canopy, smaller-

forewing yet with stouter-body characterised the nymphalids sampled here.

Keywords: Species Diversity, fruit-feeding butterflies, Nymphalidae, Kubah National Park

Kepelbagaian Spesies Rama-rama Pemakan Buah (Lepidoptera: Nymphalidae) dalam

Dimensi Ruang, Masa dan Menegak di Taman Negara Kubah, Sarawak.

Abstrak

Kepelbagaian dan taburan keseluruhan rama-rama pemakan buah dalam dimensi ruang,

masa dan menegak telah dikaji di hutan dipterokap Taman Negara Kubah, Sarawak, dengan

menggunakan perangkap buah. Kerja lapangan selama 14 hari berturut-turut untuk setiap

bulan dari 17 Mei hingga 17 November 2009 telah merekodkan 665 individu daripada 49

spesies dan empat subfamili. Bassarona dunya telah direkodkan sebagai spesies rama-rama

dengan bilangan individu tertinggi di dalam kajian ini. Jumlah spesies tertinggi direkodkan

di kawasan pinggir hutan, manakala jumlah kepelbagaian tertinggi adalah di kawasan hutan

sekunder (H’= 3.033, 1-D= 0.933, α= 13.370). Sebaliknya, kawasan hutan primer

merekodkan kepelbagaian taksonomi tertinggi, iaitu kumpulan rama-rama dengan

kepelbagaian genus dan subfamili tertinggi. Rekod jumlah spesies tertinggi di kawasan

pinggir hutan adalah disebabkan ciri-ciri rama-rama pemakan buah ini yang tertarik kepada

cahaya matahari, manakala tahap pertengahan degradasi di hutan sekunder yang

membolehkan spesies pokok pionir dan klimaks hidup bersama membolehkan lebih banyak

mikrohabitat tersedia untuk rama-rama ini. Kesan daripada pelbagai replikasi bulan

terhadap taburan rama-rama pemakan buah juga direkodkan, selain daripada kesan

pengaruh taburan hujan terhadap kumpulan rama-rama ini. Hujan lebat didapati

mengganggu kehadiran rama-rama pemakan buah disebabkan lebih tertarik kepada cahaya

matahari, di samping boleh mempengaruhi pupa dan larva rama-rama ini. Kajian taburan

menegak menunjukkan kehadiran rama-rama pemakan buah yang lebih banyak di paras

permukaan tanah, sesuai dengan penemuan banyak buah-buahan yang jatuh di paras yang

sama. Fenomena seperti heliofobia juga diandaikan untuk B. dunya, dengan dominasi penuh

spesies ini di paras permukaan tanah. Rama-rama dengan kepak lebih besar dan badan

memanjang direkodkan di paras permukaan tanah, untuk lebih mudah terbang di antara

tumbuhan herba di paras ini. Manakala, untuk terbang dengan lebih efisien di paras kanopi,

rama-rama di paras ini mempunyai kepak yang lebih kecil dan badan yang membujur.

Kata Kunci: Kelimpahan spesies, rama-rama pemakan buah, Nymphalidae, Taman Negara

Kubah

Table of Contents

Declaration ii

Dedication iii

Acknowledgements iv

Abstract vi

Abstrak viii

Table of Contents x

List of Figures xvi

List of Tables xx

Abbreviations xxv

CHAPTER ONE

General Introduction 1

CHAPTER TWO

Literature Review 5

2.1 Tropical lowland mixed-dipterocarp forest 5

2.1.1 Species composition and tree diversity in a mixed-dipterocarp forest 6

2.1.2 Characteristics of tropical climate in the Bornean rainforest 9

2.1.3 Natural disturbances as a component of tropical diversity 10

2.2 Species diversity in different spatial, temporal and vertical distributions 11

2.2.1 Spatial dimension in the tropical rainforest 12

2.2.2 Temporal dimension in the tropical rainforest 15

2.2.3 Vertical dimension in the tropical rainforest 16

2.2.4 The association of spatial, temporal and vertical dimensions in

determining the species distribution in tropical rainforest

20

2.3 The selection of fruit-feeding butterflies (Lepidoptera: Nymphalidae) 22

2.3.1 Taxonomic classification of the fruit-feeding butterflies 22

2.3.2 Distinctive features of the fruit-feeding butterflies 23

2.3.3 The importance of fruit-feeding butterflies (Lepidoptera:

Nymphalidae) for conservational purposes in tropical rainforest

24

CHAPTER THREE

General Methodology

3.1 Introduction 26

3.2 Study area 26

3.3 Study community 30

3.4 Field sampling 30

CHAPTER FOUR

Overall Diversity and Abundance of the Fruit-Feeding Butterflies

(Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak

4.1 Introduction 32

4.2 Materials and methods

4.2.1 Study area 35

4.2.2 Study community 35

4.2.3 Field sampling 35

4.2.4 Statistical analysis 35

4.3 Results

4.3.1 Overall species diversity and abundance of the fruit-feeding

butterflies

38

4.3.2 Rank-abundance distribution of the fruit-feeding butterflies 44

4.3.3 Taxonomic diversity of the fruit-feeding butterflies across primary,

secondary, heath and forest edge

45

4.4 Discussion

4.4.1 Overall species diversity and abundance of the fruit-feeding

butterflies

50

4.4.2 The segregation of spatial, temporal and vertical dimensions in

determining the fruit-feeding butterflies’ distribution

51

4.4.3 Rank-abundance distribution of the fruit-feeding butterflies 53

4.4.4 Taxonomic diversity of the fruit-feeding butterflies in different

forest habitats

54

4.5 Conclusion 58

CHAPTER FIVE

Habitat Preference of Fruit-feeding Butterflies (Lepidoptera: Nymphalidae)

in Kubah National Park, Sarawak

5.1 Introduction 59

5.2 Materials and methods

5.2.1 Study area 62

5.2.2 Study community 62

5.2.3 Field sampling 62

5.2.4 Statistical analysis 62

5.3 Results

5.3.1 Species diversity and abundance of the nymphalids in different

forest habitats

65

5.3.2 Patterns of species abundance with species-rank of the nymphalids,

and the distribution of rare as well as common species in different

forest habitats

72

5.3.3 Abundance-based similarity of the nymphalids assemblages

between different forest habitats

75

5.3.4 Interhabitat comparisons with the effects of rainfall to the

nymphalid assemblages in different forest habitats

76

5.4 Discussion 80

5.4.1 Specific associations of the nymphalids with different forest habitats

and environmental gradients

81

5.4.2 The relationship between geographical distribution of the

nymphalids and forest habitats

85

5.4.3 Similarity of the nymphalids assemblages and the interhabitat

comparisons in different forest habitats

87

5.4.4 Effects of rainfall distribution to the spatial dimension of the

frugivorous butterflies

89

5.5 Conclusion 91

CHAPTER SIX

Temporal Illumination with the Effects of Rainfall to the Fruit-feeding

Butterflies (Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak

6.1 Introduction 92

6.2 Materials and methods

6.2.1 Study area 94

6.2.2 Study community 94

6.2.3 Field sampling 94

6.2.4 Statistical analysis 95

6.3 Results

6.3.1 Overall diversity and abundance of the nymphalids in six monthly

replicates

97

6.3.2 Similarity of the nymphalids assemblages between six monthly

replicates

105

6.3.3 Effects of rainfall to the overall diversity and abundance of the

nymphalids

106

6.3.4 Abundance analyses as well as temporal patterns of the nymphalids

between seasonal replicates

110

6.4 Discussion

6.4.1 Overall diversity, abundance and similarity of the nymphalids in

relation with the rainfall distribution

112

6.4.2 Effects of rainfall and temperature to the temporal patterns of the

nymphalids

115

6.5 Conclusion 117

CHAPTER SEVEN

Vertical Distribution of Fruit-Feeding Butterflies (Lepidoptera:

Nymphalidae) in Kubah National Park, Sarawak

7.1 Introduction 118

7.2 Materials and methods

7.2.1 Study area 121

7.2.2 Study community 121

7.2.3 Field sampling 121

7.2.4 Statistical analysis 122

7.3 Results

7.3.1 Overall diversity and abundance of the nymphalids in both ground

and canopy strata

123

7.3.2 Vertical fluctuations of the nymphalids in six monthly replicates,

and in relation with the rainfall distribution

126

7.3.3 Vertical occurrence patterns of the nymphalids in different forest

habitats

129

7.3.4 The relationship between morphological parameters of the

nymphalids with strata heights

137

7.4 Discussion

7.4.1 Overall diversity and abundance of the nymphalids in both ground

and canopy strata

140

7.4.2 Vertical fluctuations of the nymphalids in six monthly replicates,

and in relation with the rainfall distribution

142

7.4.3 Specific trends of the nymphalids assemblages along the vertical

gradient

143

7.4.4 Vertical occurrence patterns of the nymphalids in different forest

habitats

147

7.4.5 The relationship between morphological parameters of the

nymphalids with strata heights

151

7.5 Conclusion 153

CHAPTER EIGHT

General Discussion and Conclusion

8.1 General discussion 155

8.2 General conclusion 160

REFERENCES 164

APPENDIX A 182

APPENDIX B 188

APPENDIX C 191

List of Figures

Figure 2.1 Forest zones on the main mountain, after Whitmore (1998). 8

Figure 2.2 Subdivisions of the butterflies (Lepidoptera: Rhopalocera) after Corbet and

Pendlebury (1992).

22

Figure 2.3 Seven of the subfamilies (bold) are represented in Sundaland, Ithomiinae is

confined to the Neotropical region, Tellervinae in the Papuan subregion

whereas Calinaginae is a Sino-Himalayan subfamily (Corbet &

Pendlebury, 1992).

23

Figure 3.1 Study area which is located in Kubah National Park, Kuching, Sarawak, in

southern part of Malaysian Borneo (Source: Modified after Google Map

2011).

27

Figure 3.2 (A) Study site in Kubah National Park, Sarawak with four sampling areas;

A. Primary forest, B. Secondary forest, C. Forest edge, D. Heath forest; (B)

Numbers designate individual replicate sampling units which represents

one canopy and one ground level trap (Note: Map is not up to scale)

(Source: Hazebroek & Abang (2000)).

28

Figure 4.1 Total abundance of fruit-feeding butterflies (Lepidoptera: Nymphalidae)

sampled in baited traps in Kubah National Park, Sarawak.

40

Figure 4.2 Percentage representation of species and individuals of the fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak.

40

Figure 4.3 Rank-abundance distributions of fruit-feeding butterflies (Lepidoptera:

Nymphalidae) sampled in Kubah National Park, Sarawak (Species rank are

based on Hamer et al., 2003).

45

Figure 4.4 Dendogram based on hierarchical agglomerative clustering (group-average

linking) of fruit-feeding butterflies (Lepidoptera: Nymphalidae) in four

different forest habitats with the Master-list (Bornean butterflies) in Kubah

National Park, Sarawak.

48

Figure 4.5

Measured values of ∆+ of 4 fruit-feeding butterflies’ assemblages in Kubah

49

National Park, Sarawak, plotted against the number of species as points on

the simulated 95% confidence funnel and theoretical 95% confidence

funnel.

Figure 5.1 Total species richness and abundance of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak.

66

Figure 5.2 (A) Species accumulation curve for the total community of fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in four different forest habitats of

Kubah National Park, Sarawak. (B) Individual rarefaction curve computed

by PAST (version 1.96) with standard errors converted to 95% confidence

interval (blue line).

69

Figure 5.3 Species abundance of fruit-feeding butterflies (Lepidoptera: Nymphalidae)

with species-rank after Hamer et al. (2003), for every forest habitats in

Kubah National Park, Sarawak.

74

Figure 5.4 Species overlap among three forest habitats in Kubah National Park,

Sarawak. For graphic clarity the heath forest was omitted in this analysis,

giving 45 total species. (A) Rare species, which are represented by n ≤ 3

individuals; (B) common species represented by n ≥ 4 individuals. (Each

comparison was carried out independently; e.g. rare species at the forest

edge could be common in the secondary forest).

75

Figure 5.5 Dendogram based on Bray-Curtis similarity index of fruit-feeding

butterflies sampled in four different forest types, in Kubah National Park,

Sarawak.

76

Figure 5.6 Interhabitat comparisons of observed richness, abundance and diversity of

fruit-feeding butterflies (Lepidoptera: Nymphalidae) as well as the

abundance of all subfamilies (except Charaxinae), were observed to be

dependent upon the timing of the seasonal replicate (monthly). P-values for

two-way repeated measure ANOVAs are shown for season (S) and habitat

(H).

79

Figure 6.1

Monthly variations for each subfamily (Lepidoptera: Nymphalidae), in

100

Kubah National Park, Sarawak; (A) number of individuals; (B) total

number of species.

Figure 6.2 Monthly variations of ten fruit-feeding butterfly species (Lepidoptera:

Nymphalidae) which were sampled in all six sampling occasions, in Kubah

National Park, Sarawak.

101

Figure 6.3 Dendogram based on Bray-Curtis similarity coefficient (UPGMA) of six

assemblages of fruit-feeding butterflies, in six continuous months, in

Kubah National Park, Sarawak.

106

Figure 6.4 Number of species and individuals of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) sampled, with the accumulated rainfall data,

in Kubah National Park, Sarawak (A. Primary forest, B. Secondary forest,

C. Forest Edge, D. Heath forest).

107

Figure 6.5 Number of species and individuals of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) with monthly mean temperature (°C),

monthly mean relative humidity (%), and monthly rainfall (mm) for six

months (May to Nov 2009) in Kubah National Park, Sarawak.

108

Figure 7.1 Species accumulation curves showing total species versus cumulative

individual abundance through time, in vertical dimensions.

126

Figure 7.2 Total species richness (A) as well as abundance (B) for the fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in each stratum throughout the six

months of sampling, in Kubah National Park, Sarawak.

127

Figure 7.3 Temporal variations in the total community of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) throughout the six sampling occasions; (A)

species richness, (B) abundance and (C) overall community, in Kubah

National Park, Sarawak.

128

Figure 7.4 Species overlap among primary, secondary and forest edge (heath forest

was omitted for better clarity) of fruit-feeding butterflies (Lepidoptera:

Nymphalidae) in Kubah National Park, Sarawak; (A) ground and (B)

canopy.

132

Figure 7.5

Total species richness and abundance of the fruit-feeding butterflies

134

(Lepidoptera: Nymphalidae) in; (A) four subfamilies; (B) and sustained in

four forest habitats, partitioned by vertical dimension, in Kubah National

Park, Sarawak.

Figure 7.6 Vertical patterns of the fruit-feeding butterflies (Lepidoptera:

Nymphalidae); (A) for subfamilies (all forest areas pooled together) and;

(B) in all forest habitats (all nymphalid subfamilies pooled together), in

Kubah National Park, Sarawak.

135

Figure 7.7 Total community curve (solid line) with the 95% confidence interval

(dashed line) of the fruit-feeding butterflies (Lepidoptera: Nymphalidae)

sampled, with species richness for all forest habitats and cumulative

species abundance for vertical dimensions (G: ground, C: canopy), in

Kubah National Park, Sarawak.

136

Figure 7.8 Dendogram of a cluster analysis showing the degree of abundance-based

similarity of fruit-feeding butterflies (Lepidoptera: Nymphalidae), over

four different forest habitats partitioned by vertical dimension, in Kubah

National Park, Sarawak.

137

List of Tables

Table 2.1 Listed synusiae of the tropical rainforest, after Richards (1996). 6

Table 2.2 Characters of structure and physiognomy used to define the principal

mountain forest formations, after Whitmore (1998). Text marked with

asterisk (*) refers to the forest formation applied for Kubah National Park,

Sarawak.

7

Table 2.3 List and descriptions of climbers as well as epiphytes which characterise

tropical rainforest, summarised from Richards (1996), Whitmore (1998),

Primack & Corlett (2005) and Freiberg (2001).

8

Table 2.4 Summarised methods of canopy access and entomological techniques in

the tropical rainforest, after Basset et al. (2003).

19

Table 2.5 Main characteristics of the fruit-feeding butterflies (Lepidoptera:

Nymphalidae) after Corbet & Pendlebury (1992).

23

Table 2.6 Summarised descriptions of the four subfamilies (Lepidoptera:

Nymphalidae) mostly sampled in the Bornean tropical rainforest after

Corbet & Pendlebury (1992).

24

Table 3.1 Habitat descriptions for study area in Kubah National Park, Sarawak, based

on Hazebroek & Abang (2000) and personal observations.

29

Table 3.2 Summary of all six sampling occasions with study period of each occasion

during May to November 2009, in Kubah National Park, Sarawak.

31

Table 4.1 Summarised trap allocations in every forest habitat in the study area, in

Kubah National Park, Sarawak.

35

Table 4.2 Summarised list of fruit-feeding butterflies (Lepidoptera: Nymphalidae)

collected by using baited traps in Kubah National Park, Sarawak; (a)

overall, (b) list of species with ≥ 20 individuals and (c) singletons.

38

Table 4.3 Diversity indices for fruit-feeding butterflies (Lepidoptera: Nymphalidae)

and for each subfamily, sampled in Kubah National Park, Sarawak.

42

Table 4.4 Species richness, abundance and diversity of the fruit-feeding butterflies

(Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak. (Each

sampling unit consists of two; ground and canopy, functional baited traps).

42

Table 4.5 (A) Species richness of the total community of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) partitioned by vertical position. Rare species

43

are those represented by < 4 individuals, and common species are those

represented by ≥ 5 individuals. (B) Individual abundance of the community

partitioned by vertical position.

Table 4.6 Distribution of the total species richness of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) among habitats, in Kubah National Park,

Sarawak.

43

Table 4.7 Chi-squared tests for homogeneity of species abundance distributions

among heights, habitats and months (p < 0.05) for the total community of

fruit-feeding butterflies (Lepidoptera: Nymphalidae) in Kubah National

Park, Sarawak (ns = not significant).

44

Table 4.8 Conventional and taxonomic diversity indices of the fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in four different forest habitats in

Kubah National Park, Sarawak. (Values in parentheses show the range for

95% confidence interval, derived from PAST v.1.96).

47

Table 5.1 Summarised trap allocations in every forest habitat in the study area, in

Kubah National Park, Sarawak.

62

Table 5.2 Species richness, abundance and diversity (Fisher’s α) of the fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in Kubah National Park, Sarawak.

(Each sampling unit consists of two; ground and canopy, functional baited

traps).

67

Table 5.3 Diversity indices for fruit-feeding butterflies (Lepidoptera: Nymphalidae),

sampled in four different forest types in Kubah National Park, Sarawak.

68

Table 5.4 Abundance of fruit-feeding butterflies (Lepidoptera: Nymphalidae) in

Kubah National Park, Sarawak; (a) species with ≥ 20 individuals; (b)

subfamilies. The abundance of each species/ subfamily is tested against the

null hypothesis of homogeneity of distribution between forest habitats.

Species with asterisks (*) depart significantly (p < 0.05) between forest

habitats (ns = not significant).

71

Table 5.5 Similarity indices (upper diagonal for Morisita-Horn’s similarity index,

lower diagonal for Bray-Curtis similarity index) for the fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in four different forest habitats, in

76

Kubah National Park, Sarawak.

Table 5.6 Average Bray-Curtis dissimilarity scores (and ANOSIM p-values) for

between habitat comparisons of the community structure of all fruit-

feeding butterflies (Lepidoptera: Nymphalidae) and subfamilies.

Significant interhabitat comparisons are marked with asterisks accordingly

(p < 0.05).

78

Table 6.1 Summarised trap allocations in every forest habitat in the study area, in

Kubah National Park, Sarawak.

95

Table 6.2 Species richness and abundance of fruit-feeding butterflies (Lepidoptera:

Nymphalidae) in Kubah National Park, Sarawak; (a) for each respective

sampling occasions, and (b) for every subfamilies (upper part refer to

species number, whereas the lower part refer to individuals number with

Fisher’s-Alpha in parentheses). Bold values indicate the highest diversity.

98

Table 6.3 Abundance of fruit-feeding butterflies (Lepidoptera: Nymphalidae) with ≥

20 individuals, tested against the null hypothesis of homogeneity of

distribution between six sampling occasions, in Kubah National Park,

Sarawak.

102

Table 6.4 Species abundance distribution of 11 singletons (Lepidoptera:

Nymphalidae) sampled in Kubah National Park, Sarawak.

103

Table 6.5 Total number of species sampled every month during the field sampling,

with species representation (%) from the overall total species sampled.

104

Table 6.6 Diversity indices for fruit-feeding butterflies (Lepidoptera: Nymphalidae),

in six continuous months, sampled in Kubah National Park, Sarawak.

104

Table 6.7 Similarity indices between six monthly replicates; upper diagonal are the

Morisita-Horn’s similarity indices, whereas the lower diagonal are Bray-

Curtis similarity indices, of fruit-feeding butterflies in Kubah National

Park, Sarawak.

105

Table 6.8 p-values computed from the Mann-Whitney test, for the comparisons of

abundance between sampling occasions of fruit-feeding butterflies

(Lepidoptera: Nymphalidae) sampled in Kubah National Park, Sarawak.

Significant comparisons (p < 0.05) were marked with asterisks (ns = not

111

significant).

Table 6.9 Pairwise correlations across forest habitats between the six monthly

replicates (n = 4), based on fruit-feeding butterflies (Lepidoptera:

Nymphalidae) abundance and richness. Significant correlation is marked

with asterisks (p < 0.05).

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Table 7.1 Summarised trap allocations in every forest habitat in the study area, in

Kubah National Park, Sarawak.

121

Table 7.2 Vertical distribution for 20 most abundant (n ≥ 20) fruit-feeding

butterflies’ species (Lepidoptera: Nymphalidae), pooled across the four

forest habitats and 84-days of sampling, in Kubah National Park, Sarawak.

Species with asterisks depart significantly (p < 0.05) between forest strata

(ns = not significant).

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Table 7.3 Total number of individuals and species (Lepidoptera: Nymphalidae)

sampled in 40 baited traps (20 at the ground level, 20 at the canopy level)

over the 84 days of sampling, in Kubah National Park, Sarawak.

124

Table 7.4 Diversity indices for the fruit-feeding butterflies (Lepidoptera:

Nymphalidae) pooled over forest habitats and sampling occasions, in a

vertical distribution, sampled in Kubah National Park, Sarawak.

125

Table 7.5 Diversity indices for overall community and subfamilies of fruit-feeding

butterflies (Lepidoptera: Nymphalidae) in vertical distribution, sampled in

Kubah National Park, Sarawak.

125

Table 7.6 Species richness and abundance of the pooled fruit-feeding butterflies

(Lepidoptera: Nymphalidae) in all four different forest habitats, partitioned

by vertical position.

129

Table 7.7 Species richness, abundance and diversity of the fruit-feeding butterflies

(Lepidoptera: Nymphalidae) in vertical dimension, for all four forest

habitats in Kubah National Park, Sarawak.

131

Table 7.8 Spearman rank correlation between fruit-feeding butterflies’ (Lepidoptera:

Nymphalidae) abundance and species richness with trap heights in Kubah

131