Coleoptera:Tenebrionidae) from Malaysia

UNIVERSITI PUTRA MALAYSIA

TAXONOMIC REVISION OF STINOCHIINAE, TENEBRIONINAE AND DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) FROM MALAYSIA

DARYA KAREEM HWAYYIZ

FP 2017 6

TAXONOMIC REVISION OF STINOCHIINAE, TENEBRIONINAE AND DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) FROM MALAYSIA UPM

DARYA KAREEM HWAYYIZ

March 2017

i COPYRIGHT

All material contained within the thesis, including without limitation to text, logos, icons, photographs and all other artwork, is copyright material of University Putra Malaysia unless otherwise stated. Use may be made of any material contained within the thesis for non-commercial purposes from the copyright holder. Commercial use of material may only be made with the express, prior, written permission of University Putra Malaysia.

UPM

DEDICATION

I would like to dedicate to my beloved husband who helped me all the time during my study. Similarly, many thanks go to my father, brothers and sisters who supported me with prayers and endured the pain of being away for four years. In last, I also would like to dedicate my friends, who helped me with all what they got and these are the people whom I will never forget.

UPM

iii

Abstract of thesis presented to the Senate of Universiti Putra Malaysia in fulfillment of the requirement for the Degree of Doctor of Philosophy

TAXONOMIC REVISION OF STINOCHIINAE, TENEBRIONINAE AND DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) FROM MALAYSIA

DARYA KAREEM HWAYYIZ

March 2017

UPM Chairman : Associate Professor Nur Azura Adam, PhD Faculty : Agriculture

This study established the current taxonomic status of 30 species of darkling beetles (Coleoptera: Tenebrionidae) in Malaysia. A total of 392 specimens were examined. Thirty species has been identified belonging to three subfamilies, nine tribes and 17 genera with four potential new species and one potential new genus. A total number of 16 species were recorded for the first time in Malaysia namely Amarygmus metallicus, A. ovoides, Derosphaerus aeruginosus, Gonocephalum hispidocostatum, Leprocaulus rotundicollis, Promethis semisulcata, P. coracina, P. punctulator, P. opaca, Pseudonautes vagevittatus, Pseudoblaps javana, Strongylium erythrocephalum, S. orientale, S. various, Uloma sextuberosa and U. excise. A total of 42 internal and external morphological characteristic have been described for each species. Eleven structures for each species were illustrated based on their specific state namely antenna, labrum, labium, mandible, maxilla, pronotum, fore-leg, mid-leg, hind-leg, elytra, aedeagus. Taxonomic key to genera of subfamily Stinochiinae, genera of subfamily Tenebrioninae and the species of the genera Promethis, Strongylium, Eucyrtus, Gauromaia, Amargymus, Uloma, Tribolium species has been successfully developed. The distribution of Tenebrionidae species in Malaysia were shown in the maps for each species. The most common species is Tribolium castanum, Tribolium confusum, some species of the genus Promethis, the less common species is UlomaCOPYRIGHT excisa, and rare species is Strongylium varians. A comprehensive checklist of Malaysian Tenebrionidae was developed for the first time with the total number of 207 species belonging to 64 genera. The highest number of the species has been found in Borneo followed by Penang and Malacca. The less number of the species have been © found in Perak. The taxonomical effort produced from this study will be helpful for enhancing the species discovery while maintaining species inventories of Malaysian Tenebrionidae.

Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai memenuhi keperluan untuk Ijazah Doktor Falsafah

SEMAKAN TAKSONOMI STINOCHIINAE, TENEBRIONINAE DAN DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) DARI MALAYSIA

Oleh

DARYA KAREEM HWAYYIZ Mac 2017 UPM Pengerusi : Profesor Madya Nur Azura Adam, PhD Fakulti : Pertanian

Kajian ini telah memperihalkan status semasa taksonomi 30 spesies kumbang Tenebrionidae (Coleoptera) di Malaysia. Sejumlah 392 spesimen telah diperiksa. Sebanyak 30 spesies telah diperihal tergolong dalam tiga subfamili, sembilan tribus dan 17 genus dengan potensi tiga spesies baru dan satu genus baru. Sejumlah 16 spesies telah direkodkan buat kali pertama di Malaysia iaitu Amarygmus metallicus, A. ovoides, Derosphaerus aeruginosus, Gonocephalum hispidocostatum, Leprocaulus rotundicollis, Promethis semisulcata, P. coracina, P. punctulator, P. opaca, Pseudonautes vagevittatus, Pseudoblaps javana, Strongylium erythrocephalum, S. orientale, S. various, Uloma sextuberosa and U. excise. Sebanyak 42 ciri morfologi dalaman dan luaran telah diperihalkan bagi setiap spesies. Sebanyak 11 struktur bagi setiap spesies telah diilustrasi berdasar kepada ciri spesifiknya iaitu antena, labrum, labium, mandibel, maksila, pronotum, kaki hadapan, kaki tengah, kaki belakang, elitra dan aedegus. Taburan spesies Tenebrionidae di Malaysia ditunjukkan dalam peta. Spesies paling umum ialah Tribolium castanum, Tribolium confusum dan sebahagian spesies di bawah genus Promethis manakala spesies kurang umum ialah Uloma excisa, manakala spesies langka ialah Strongylium varians. Satu senarai semak yang komprehensif telah dibangunkan buat pertama kalinya untuk spesies Tenebrionidae di Malaysia di mana jumlah spesies Tenebrionidae Malaysia adalah sebanyak 207 spesiesCOPYRIGHT terdiri daripada 64 genera. Jumlah spesies terbanyak di jumpai di Borneo diikuti oleh Pula Pinang dan Melaka. Perak mempunyai bilangan spesies terendah. Usaha kajian taksonomi daripada kajian ini membantu dalam meningkatkan penemuan spesies di samping menyelenggara inventori spesies Tenebrionidae di © Malaysia.

ii ACKNOWLEDGEMENTS

All praise and gratitude for the beloved Allah; who is the compassionate and the most Merciful. This work would have been impossible without the continuous support and supervision of my supervisor, Associate Professor Dr.Nur Azura Adam. All steps taken on the way to finishing this thesis were under her direct guidance and also the other members of the supervisory committee, Dr. Lau Wei Hong and Dr. Izfa Riza and also thankful for the laboratory assistant, Mr. Hishamuddin Zainuddin, who endured with great patience in all my laboratory tasks.

I would like to express my sincere gratitude to Dr. Julio Ferrer who supported me spiritually and scientifically. I thank the individuals who helped my field sampling Dr. Azman Sulaiman and Mr. Audi and thanks to all workers in Kinabalu park who they gave me the chance to check my specimen and loaning me some of theirUPM specimen.

iii UPM

COPYRIGHT © This thesis was submitted to the Senate of the Universiti Putra Malaysia and has been accepted as fulfillment of the requirement for the degree of Doctor of Philosophy. The members of the Supervisory Committee were as follows:

Nur Azura Adam, PhD Associate Professor Faculty of Agriculture Universiti Putra Malaysia (Chairman)

Lau Wei Hong, PhD Senior Lecturer Faculty of Agriculture Universiti Putra Malaysia (Member) UPM

Izfa Riza Hazmi, PhD Senior Lecturer School of Environmental and Natural Resources Sciences Faculty of Science and Technology Universiti Kebangsaan Malaysia (Member)

ROBIAH BINTI YUNUS, PhD Professor and Dean School of Graduate Studies Universiti Putra Malaysia COPYRIGHTDate: ©

v Declaration by graduate student

I hereby confirm that:  this thesis is my original work;  quotations, illustrations and citations have been duly referenced;  this thesis has not been submitted previously or concurrently for any other degree at any institutions;  intellectual property from the thesis and copyright of thesis are fully-owned by Universiti Putra Malaysia, as according to the Universiti Putra Malaysia (Research) Rules 2012;  written permission must be obtained from supervisor and the office of Deputy Vice-Chancellor (Research and innovation) before thesis is published (in the form of written, printed or in electronic form) including books, journals, modules, proceedings, popular writings, seminar papers, manuscripts, posters, reports, lecture notes, learning modules or any other materials as stated in the Universiti Putra Malaysia (Research) Rules 2012; UPM  there is no plagiarism or data falsification/fabrication in the thesis, and scholarly integrity is upheld as according to the Universiti Putra Malaysia (Graduate Studies) Rules 2003 (Revision 2012-2013) and the Universiti Putra Malaysia (Research) Rules 2012. The thesis has undergone plagiarism detection software

Signature: ______Date: ______

Name and Matric No: Darya Kareem Hwayyiz , GS35191

Declaration by Members of Supervisory Committee

This is to confirm that:  the research conducted and the writing of this thesis was under our supervision;  supervision responsibilities as stated in the Universiti Putra Malaysia (Graduate Studies) Rules 2003 (Revision 2012-2013) were adhered to.

Signature: Name of Chairman of Supervisory Committee: Associate Professor Dr. Nur Azura Adam

UPM

Signature: Name of Member of Supervisory Committee: Dr. Lau Wei Hong

Signature: Name of Member of Supervisory Committee: Dr. Izfa Riza Hazmi

vii

TABLE OF CONTENTS

Page

ABSTRACT i ABSTRAK ii ACKNOWLEDGEMENTS iii APPROVAL iv DECLARATION vi LIST OF TABLES x LIST OF FIGURES xi LIST OF ABBREVIATIONS xvii

CHAPTER

1 INTRODUCTION 1 1.1 Background of study 1 1.2 Significance of the study UPM2 1.3 Research Objective(s) 2 1.4 Outline of thesis 2

2 LITERATURE REVIEW 3 2.1 Taxonomic Status of family Tenebrionidae 3 2.2 Evolution of the Phylogenetic study 4 2.3 Family division 5 2.4 The economic importance of the family 5 2.5 Biological studies for the family 8 2.6 Distribution study 8 2.7 Distribution of the family 9 2.8 Molecular studies of the family 12

3 MATERIAL AND METHODS 13 3.1 Insect collection 13 3.2 Insect preservation 14 3.3 Morphological and Anatomical study 14 3.4 Image analysis 15 3.5 Identification 16 3.6 Identificationkey 16 3.7 Illustration 16 COPYRIGHT3.8 Checklistof Malaysian Tenebrionidae 17 4 RESULTS AND DISCUSSION 18 4.1 Identification and Key construction of subfamily 18 © Stinochiinae, Tenebrioninae and Diaperinae (Tenebrionidae) 4.2 Distribution of the species 19 4.3 Identification Key 23 4.3.1 Key to subfamily Stinochiinae 23 4.3.2 Key to subfamily Tenebrioninae 24 4.3.3 Subfamily Diaperinae 25 4.4 Description and illustration 26

viii Promethis semisulcata (Faimaire, 1882) 26 Promethis coracina (Knoch, 1801) 33 Promethis punctulator (Fairmaire, 1883) 37 Promethis opaca Carter, 1914. 41 Rophobas asperatus (Fairmaire, 1882) 45 Eucyrtus pretiosus Lacordaire , 1859 49 Eucyrtus anthracinus Fairmaire, 1882 54 Asbolodes humerosus Fairmaire. 1893 60 Pseudonautes vagevittatus Fairmaire,1893 64 Gauromaia annulipes (Pic 1921) 68 Gauromaia haagi Fairmaire, 1896 72 Camptobrachys sp.n. 77 New genuse 82 Strongylium Kirby, 1818 86 Strongylium erythrocephalum (Fabricius 1801) 91 Strongylium orientaie Fairmaire, 1893 95 Strongylium n.sp. UPM99 Strongylium forcipicolle Fairmaire 1900 103 Leprocaulus rotundicollis Pic, 1922 107 Derosphaerus aeruginosus (Fabricius, 1787) 112 Pseudoblaps javana Wiedemann, 1819 116 Amarygmus metallicus ( Perty ,1831) 120 Amarygmus aeneolus Fairmaire, 1893 124 Gonocephalum hispidocostatum (Fairmaire, 1883) 128 Uloma sextuberosa Kaszab, 1980 132 Uloma excise (Gebien, 1913) 137 Alphitobius diaperinus (Panzer, 1797) 141 Tribolium confusum (Jacquelin, 1868) 146 Tribolium castanum (Herbst, 1797) 151 Hemiceraspn 203 155 4.5 Checklist of Malaysian Tenebrionidae 161

5 SUMMARY, CONCLUSIONS AND RECOMMENDATIONS 167 FOR FUTURE RESEARCH 5.1 Summary 167 5.2 Conclusion and recommendation for future research 167

REFERENCES 169 APPENDICES 179 BIODATACOPYRIGHT OF STUDENT 185 LIST OF PUBLICATIONS 186 ©

LIST OF TABLES

Table Page

2.1 Tenebrionidae economic important 7

2.2 IndoMalayan groups 11

4.1 The collected genera and the species of Subfamilies Stinochiinae, 18 Tenebrioninae and Diaperinae (Tenebrionidae).

4.2 Less common and most common species location in the study states 22 with their GPS coordinates (Appendice).

4.3 Number of the species belonging to each genus 162

4.4 List of Malaysian Tenebrionidae (Coleoptera) and occurrenceUPM of 163 species in each Malaysian state.

LIST OF FIGURES

Figure Page

2.1 The life cycle of Darkling beetles 8

3.1 Collection of insets through a. light traps and b. pitfall traps 13

3.2 Collection boxes of the insect 14

3.3 Examination of morphological characters of tenebrionidae under 15 Dinolite

3.4 Promethis semisulcata showing a. Dorsal view b. Ventral view 17

4.1 Distribution of each species of family Tenebrionidae in MalUPMaysia. 20

4.2 Head of Promethis semisulcata (a) dorsal side (b) ventral side 28

4.3 Antenna and mouth part of Promethis semisulcata (a) Antennae. 29 (b) Labrum. (c) Labium. (d) Maxilla. (e). Mandible.

4.4 Thorax of Promethis semisulcata (a) Pronotum. (b)Fore leg. 30 (c)Midleg. (d)Hindleg

4.5 Elytra and Abdomen of Promethis semisulcata (a) Elytra. (b) 31 Abdomen.

4.6 Male and Female genitalia of Promethis semisulcata (a) Male 32 genitalia (b) Female genitalia

4.7 Antenna and mouth part of Promethis coracina (a) Antennae. (b) 35 Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.8 Thorax of Promethis coracina (a) Pronotum. (b) Fore leg. (c) 36 Midleg. (d) Hindleg.

4.9 Elytra and Aedeagus of Promethis coracina (a) Elytra. (b) 37 COPYRIGHTAedeagus.

4.10 Antenna and mouth part of Promethis punctulator (a) Antennae. 39 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. © 4.11 Thorax of Promethis punctulator (a) Pronotum. (b) Fore leg. (c) 40 Midleg. (d) Hindleg.

4.12 Elytra and Aedeagus of Promethis punctulator (a) Elytra. (b) 41 Aedeagus.

4.13 Antenna and mouth part of Promethis opaca (a) Antennae. (b) 43 Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.14 Thorax of Promethis opaca (a) Pronotum. (b) Fore leg. (c) 44 Midleg. (d) Hindleg.

4.15 Elytra and Aedeagus of Promethis opaca (a) Elytra. (b) 45 Aedeagus.

4.16 Antenna and mouth part of Rophobas asperatus (a) Antennae. 47 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.17 Thorax of Rophobas asperatus (a) Pronotum. (b) Fore leg. (c) 48 Midleg. (d) Hindleg.

4.18 Elytra and Aedeagus of Rophobas asperatus (a) Elytra. (b) 49 Aedeagus. UPM

4.19 Antenna and mouth part of Eucyrtus pretiosus (a) Antennae. (b) 52 Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.20 Thorax of Eucyrtus pretiosus (a) Pronotum. (b) Fore leg. (c) 53 Midleg. (d) Hindleg.

4.21 Antenna and mouth part of Eucyrtus pretiosus (a) Pronotum. (b) 54 Forleg. (c) Midleg. (d) Hindleg.

4.22 Antenna and mouth part of Eucyrtus anthracinus (a) Antennae. 57 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.23 Thorax of Eucyrtus anthracinus (a) Pronotum. (b) Fore leg. (c) 58 Midleg. (d) Hindleg.

4.24 Elytra and Aedeagus of Eucyrtus anthracinus (a) Elytra. (b) 59 Aedeagus.

4.25 Antenna and mouth part of Asbolodes humerosus (a) Antennae. 62 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.26COPYRIGHT Thorax of Asbolodes humerosus (a) Pronotum. (b) Fore leg. (c) 63 Midleg. (d) Hindleg.

4.27 Elytra and Aedeagus of Asbolodes humerosus (a) Elytra. (b) 64 © Aedeagus

4.28 Antenna and mouth part of Pseudonautes vagevittatus (a) 66 Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.29 Thorax of Pseudonautes vagevittatus (a) Pronotum. (b) Fore leg. 67 (c) Midleg. (d) Hindleg. xii

4.30 Elytra and Aedeagus of Pseudonautes vagevittatus (a) Elytra. (b) 68 Aedeagus

4.31 Antenna and mouth part of Gauromaia annulipes (a) Antennae. 70 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.32 Thorax of Gauromaia annulipes (a) Pronotum. (b) Fore leg. (c) 71 Midleg. (d) Hindleg.

4.33 Elytra and Aedeagus of Gauromaia annulipes (a) Elytra. (b) 72 Aedeagus

4.34 Antenna and mouth part of Gauromaia haagi (a) Antennae. (b) 74 Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.35 Thorax of Gauromaia haagi (a) Pronotum. (b) Fore leg. (c) 75 Midleg. (d) Hindleg. UPM

4.36 Elytra and Aedeagus of Gauromaia haagi (a) Elytra. 76 (b)Aedeagus

4.37 Antenna and mouth part of Camptobrachys sp.n. (a) Antennae. 79 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.38 Thorax of Camptobrachys sp.n. (a) Pronotum. (b) Fore leg. (c) 80 Midleg. (d) Hindleg.

4.39 Elytra and Aedeagus of Camptobrachys sp.n. (a) Elytra. (b) 81 Aedeagus

4.40 Antenna and mouth part of n.gen. (a) Antennae. (b) Labrum. (c) 84 Labium. (d) Maxilla. (e) Mandible.

4.41 Thorax of n.gen. (a) Pronotum. (b) Fore leg. (c) Midleg. (d) 85 Hindleg.

4.42 Elytra and Aedeagus of n.gen.(a) Elytra. (b) Aedeagus 86

4.43 Antenna and mouth part of Strongylium varians (a) Antennae. 89 COPYRIGHT(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.44 Thorax of Strongylium varians (a) Pronotum. (b) Fore leg. (c) 90 Midleg. (d) Hindleg. © 4.45 Elytra and Aedeagus of Strongylium varians (a) Elytra. (b) 91 Aedeagus

4.46 Antenna and mouth part of Strongylium erythrocephalum (a) 93 Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

xiii

4.47 Thorax of Strongylium erythrocephalum (a) Pronotum. (b) Fore 94 leg. (c) Midleg. (d) Hindleg.

4.48 Elytra and Aedeagus of Strongylium erythrocephalum (a) Elytra. 95 (b) Aedeagus

4.49 Antenna and mouth part of Strongylium orientale(a) Antennae. 97 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.50 Thorax of Strongylium orientale (a) Pronotum. (b) Fore leg. (c) 98 Midleg. (d) Hindleg.

4.51 Elytra and Aedeagus of Strongylium orientale (a) Elytra. (b) 99 Aedeagus

4.52 Antenna and mouth part of Strongylium n.sp.(a) Antennae. (b) 101 Labrum. (c) Labium. (d) Maxilla. (e) Mandible. UPM

4.53 Thorax of Strongylium n.sp. (a) Pronotum. (b) Fore leg. (c) 102 Midleg. (d) Hindleg.

4.54 Elytra and Aedeagus of Strongylium n.sp.(a) Elytra. (b) 103 Aedeagus

4.55 Antenna and mouth part of Strongylium forcipicolle (a) 105 Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.56 Thorax of Strongylium forcipicolle (a) Pronotum. (b) Fore leg. 106 (c) Midleg. (d) Hindleg.

4.57 Elytra and Aedeagus of Strongylium forcipicolle (a) Elytra. (b) 107 Aedeagus

4.58 Antenna and mouth part of Leprocaulus rotundicollis (a) 110 Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.59 Thorax of Leprocaulus rotundicollis (a) Pronotum. (b) Fore leg. 111 (c) Midleg. (d) Hindleg.

4.60COPYRIGHT Elytra and Aedeagus of Leprocaulus rotundicollis (a) Elytra. (b) 112 Aedeagus

4.61 Antenna and mouth part of Derosphaerus aeruginosus (a) 114 © Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.62 Thorax of Derosphaerus aeruginosus (a) Pronotum. (b) Fore leg. 115 (c) Midleg. (d) Hindleg.

4.63 Elytra and Aedeagus of Derosphaerus aeruginosus (a) Elytra. 116 (b) Aedeagus xiv

4.64 Antenna and mouth part of Pseudoblaps javana (a) Antennae. 118 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.65 Thorax of Pseudoblaps javana (a) Pronotum. (b) Fore leg. (c) 119 Midleg. (d) Hindleg.

4.66 Elytra and Aedeagus of Pseudoblaps javana (a) Elytra. (b) 120 Aedeagus

4.67 Antenna and mouth part of Amarygmus metallicus (a) Antennae. 122 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.68 Thorax of Amarygmus metallicus (a) Pronotum. (b) Fore leg. (c) 123 Midleg. (d) Hindleg.

4.69 Elytra and Aedeagus of Amarygmus metallicus (a) Elytra. (b) 124 Aedeagus UPM 4.70 Antenna and mouth part of Amarygmus aeneolus (a) Antennae. 126 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.71 Thorax of Amarygmus aeneolus (a) Pronotum. (b) Fore leg. (c) 127 Midleg. (d) Hindleg.

4.72 Elytra and Aedeagus of Amarygmus aeneolus (a) Elytra. (b) 128 Aedeagus

4.73 Antenna and mouth part of Gonocephalum hispidocostatum (a) 130 Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.74 Thorax of Gonocephalum hispidocostatum (a) Pronotum. (b) 131 Fore leg. (c) Midleg. (d) Hindleg.

4.75 Elytra and Aedeagus of Gonocephalum hispidocostatum (a) 132 Elytra. (b) Aedeagus

4.76 Antenna and mouth part of Uloma sextuberosa (a) Antennae. (b) 135 Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 4.77COPYRIGHT Thorax of Uloma sextuberosa (a) Pronotum. (b) Fore leg. (c) 136 Midleg. (d) Hindleg.

4.78 Elytra and Aedeagus of Uloma sextuberosa (a) Elytra. (b) 137 © Aedeagus 4.79 Antenna and mouth part of Uloma excise (a) Antennae. (b) 139 Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.80 Thorax of Uloma excise (a) Pronotum. (b) Fore leg. (c) Midleg. 140 (d) Hindleg. xv 4.81 Elytra and Aedeagus of Uloma excise (a) Elytra. (b) Aedeagus 141

4.82 Antenna and mouth part of Alphitobius diaperinus (a) Antennae. 144 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.83 Thorax of Alphitobius diaperinus (a) Pronotum. (b) Fore leg. (c) 145 Midleg. (d) Hindleg.

4.84 Elytra and Aedeagus of Alphitobius diaperinus (a) Elytra. (b) 146 Aedeagus

4.85 Antenna and mouth part of Tribolium confusum (a) Antennae. 149 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.86 Thorax of Tribolium confusum (a) Pronotum. (b) Fore leg. (c) 150 Midleg. (d) Hindleg. UPM 4.87 Elytra and Aedeagus of Tribolium confusum (a) Elytra. (b) 151 Aedeagus

4.88 Antenna and mouth part of Tribolium castanum (a) Antennae. 153 (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.89 Thorax of Tribolium castanum (a) Pronotum. (b) Fore leg. (c) 154 Midleg. (d) Hindleg.

4.90 Elytra and Aedeagus of Tribolium castanum (a) Elytra. (b) 155 Aedeagus

4.91 Antenna and mouth part of Hemicera n.sp.(a) Antennae. 158 (b)Labrum. (c) Labium. (d) Maxilla. (e) Mandible.

4.92 Thorax of Hemicera n.sp.(a) Pronotum. (b) Fore leg. (c) Midleg. 159 (d) Hindleg.

xvi LIST OF ABBREVIATIONS

IMUMS Insect museum of Universti Malaysia Sabah

IMUPM Insect Museum of Universti Putra Malaysia.

CIS Center of Insect Systematic Universiti Kebangsaan Malaysia.

KCCS Kinabalu Commercial College Sabah.

FRC Forest research Center in Sabah, Sandakan NRMS Swedish of Natural history museum UPM

xvii CHAPTER ONE

INTRODUCTION

1.1 Background of the study

Tenebrionidae is the fifth largest and one of the most diverse families within order Coleoptera that contains more than 18,000 insect species (15,000 described) world wide (Matthews & Bouchard, 2008) and exhibits an extraordinary wide range of superficial dissimilarity. It makes up the large group of nocturnal beetles and usually referred as “Darkling Beetles” that is a common name for this large family.

Family Tenebrionidae is of a great economic importance as it contains insect pests that are cosmopolitan in nature and most imperatively are associated with stored products (Martin et al., 2012). Generally, these insect pests feed on plant materialsUPM including decaying matter, wood, leaf litters, pollen, fungal and algal matters. Some are also scavengers whereas few of them are predatory especially the wood boring beetles. In Tenebrionidae, insect species are mostly large in size and flightless beetles however other insect species within similar family living in rotten wood and stored products are also small in size (Soldati & Soldati, 2003).

The greater numbers of these species occur in warmer climates particularly in the more arid regions such as in the Namib and Gobi deserts. The varied adaptations of the species enable them to survive in extreme high temperature and the periods of prolonged drought. These facts have caused the Tenebrionidae to be regarded as the most highly evolved family of the Cucujoidea (Crowson, 1955). These darkling beetles that inhabit in the most torrid deserts can survive in a temperature of 50°C, they normally burrow under the stones, bark and leaf litters and they have long legs that keep their bodies at a safe distance from the burning sand and enable them to move speedily, many of these are excellent burrowers and can bury themselves in the sand immediately to escape from the heat (Schawaller, 1996; Bouchard et al., 2005).

The great importance of family-group names for the classification and nomenclatural stability of Coleoptera was clearly demonstrated by Lawrence and Newton (1995). But, due to close resemblances to the members of other families, some of the adults in familyCOPYRIGHT Tenebrionidae have been often wrongly identified in a preliminary sorting stage especially in most Carabidae (Chrysomelidae) identification (Watt, 1974). Previously, the most systematic work has been carried out at the specific and generic © levels and few have attempted to improve the higher classification within the family.

The morphological diversity of the members of this family is very difficult to diagnose the entire fauna (Aalbu et al., 2002), however a synthetic treatment of the taxonomy, biology and distribution of Tenebrionidae has been previously presented. But it still largely comprises a synthesis of the classifications given by Leconte & Horn (1883)

and Lacordaire (1859) which were mostly based on adults because of the great similarities between the species of this family so it was important to show the different characters of the species, larvae in this family tend to be more uniform and easily recognisable superficially.

Therefore, this study has been attempted to provide a stable classification scheme for the family Tenebrionidae especially for three Subfamilies such as Stinochiinae, Tenebrioninae and Diaperinae with a synoptic classification of the world fauna that summarizes recent knowledge about the relationships among suprageneric groups as well as a catalogue of family-group names especially in Malaysian agro-ecosystem.

1.2 Significance of the study

Due to the economic damages caused by some of the species of the subfamilies Stinochiinae, Tenebrioninae and Diaperinae, it is essential to conductUPM a study for their proper identification. There is still a gap in their taxonomical study of the insect species from these subfamilies especially in Malaysia therefore it is an important to identify these species in order to control their damages.

1.3 Research Objective(s)

The objectives of this study are:

1- to identify the selected species of family Tenebrionidae and study their distribution in Malaysia. 2- to construct the keys of selected species of family Tenebrionidae in Malaysia. 3- to describe and illustrate the selected morphology characters of each species and to describe the genitalia. 4- to develop a checklist of Malaysian Tenebrionidae.

1.4 Outline of thesis

This thesis is divided into five chapters. After an introductory Chapter 1, Chapter 2 presented some background of the research with a literature addressing two important topics of taxonomy and key identification. In Chapter 3, material and methods are presentedCOPYRIGHT while Chapter 4 postulated the construction of the ke ys of each species, identification, description, illustration and a checklist are also provided. Finally, in Chapter 5, summary of this study and recommendations for future research have been © presented.

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