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UNIVERSITI PUTRA MALAYSIA
TAXONOMIC REVISION OF STINOCHIINAE, TENEBRIONINAE AND DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) FROM MALAYSIA
DARYA KAREEM HWAYYIZ
FP 2017 6
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TAXONOMIC REVISION OF STINOCHIINAE, TENEBRIONINAE AND DIAPERINAE (COLEOPTERA:TENEBRIONIDAE) FROM MALAYSIA
By
DARYA KAREEM HWAYYIZ
Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia, in Fulfillment of the Requirements for the Degree of Doctor of Philosophy
March 2017
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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. Copyright © Universiti Putra Malaysia
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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.
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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
By
DARYA KAREEM HWAYYIZ
March 2017
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 Uloma 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.
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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
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 spesies terdiri daripada 64 genera. Jumlah spesies terbanyak dijumpai 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.
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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 their specimen.
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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)
Izfa Riza Hazmi, PhD Senior LecturerSchool 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
Date:
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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;
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
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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
Signature:
Name of Member of Supervisory Committee:
Dr. Lau Wei Hong
Signature:
Name of Member of Supervisory Committee:
Dr. Izfa Riza Hazmi
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TABLE OF CONTENTS
Page
ABSTRACT iABSTRAK iiACKNOWLEDGEMENTS iiiAPPROVAL ivDECLARATION viLIST OF TABLES xLIST OF FIGURES xiLIST OF ABBREVIATIONS xvii
CHAPTER
1 INTRODUCTION 11.1 Background of study 11.2 Significance of the study 21.3 Research Objective(s) 21.4 Outline of thesis 2
2 LITERATURE REVIEW 32.1 Taxonomic Status of family Tenebrionidae 32.2 Evolution of the Phylogenetic study 42.3 Family division 52.4 The economic importance of the family 52.5 Biological studies for the family 82.6 Distribution study 82.7 Distribution of the family 92.8 Molecular studies of the family 12
3 MATERIAL AND METHODS 133.1 Insect collection 133.2 Insect preservation 143.3 Morphological and Anatomical study 143.4 Image analysis 153.5 Identification 163.6 Identificationkey 163.7 Illustration 163.8 Checklistof Malaysian Tenebrionidae 17
4 RESULTS AND DISCUSSION 184.1 Identification and Key construction of subfamily
Stinochiinae, Tenebrioninae and Diaperinae (Tenebrionidae) 18
4.2 Distribution of the species 194.3 Identification Key 23
4.3.1 Key to subfamily Stinochiinae 234.3.2 Key to subfamily Tenebrioninae 244.3.3 Subfamily Diaperinae 25
4.4 Description and illustration 26
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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. 99 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
FOR FUTURE RESEARCH 167
5.1 Summary 167 5.2 Conclusion and recommendation for future research 167 REFERENCES 169APPENDICES 179BIODATA OF STUDENT 185LIST OF PUBLICATIONS 186
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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,
Tenebrioninae and Diaperinae (Tenebrionidae). 18
4.2 Less common and most common species location in the study states
with their GPS coordinates (Appendice). 22
4.3 Number of the species belonging to each genus 162 4.4 List of Malaysian Tenebrionidae (Coleoptera) and occurrence of
species in each Malaysian state. 163
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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
Dinolite 15
3.4 Promethis semisulcata showing a. Dorsal view b. Ventral view 17 4.1 Distribution of each species of family Tenebrionidae in Malaysia. 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.
(b) Labrum. (c) Labium. (d) Maxilla. (e). Mandible. 29
4.4 Thorax of Promethis semisulcata (a) Pronotum. (b)Fore leg.
(c)Midleg. (d)Hindleg 30
4.5 Elytra and Abdomen of Promethis semisulcata (a) Elytra. (b)
Abdomen. 31
4.6 Male and Female genitalia of Promethis semisulcata (a) Male
genitalia (b) Female genitalia 32
4.7 Antenna and mouth part of Promethis coracina (a) Antennae. (b)
Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 35
4.8 Thorax of Promethis coracina (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 36
4.9 Elytra and Aedeagus of Promethis coracina (a) Elytra. (b)
Aedeagus. 37
4.10 Antenna and mouth part of Promethis punctulator (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 39
4.11 Thorax of Promethis punctulator (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 40
4.12 Elytra and Aedeagus of Promethis punctulator (a) Elytra. (b)
Aedeagus. 41
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4.13 Antenna and mouth part of Promethis opaca (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
43
4.14 Thorax of Promethis opaca (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 44
4.15 Elytra and Aedeagus of Promethis opaca (a) Elytra. (b)
Aedeagus. 45
4.16 Antenna and mouth part of Rophobas asperatus (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 47
4.17 Thorax of Rophobas asperatus (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 48
4.18 Elytra and Aedeagus of Rophobas asperatus (a) Elytra. (b)
Aedeagus. 49
4.19 Antenna and mouth part of Eucyrtus pretiosus (a) Antennae. (b)
Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 52
4.20 Thorax of Eucyrtus pretiosus (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 53
4.21 Antenna and mouth part of Eucyrtus pretiosus (a) Pronotum. (b)
Forleg. (c) Midleg. (d) Hindleg. 54
4.22 Antenna and mouth part of Eucyrtus anthracinus (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 57
4.23 Thorax of Eucyrtus anthracinus (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 58
4.24 Elytra and Aedeagus of Eucyrtus anthracinus (a) Elytra. (b)
Aedeagus. 59
4.25 Antenna and mouth part of Asbolodes humerosus (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 62
4.26 Thorax of Asbolodes humerosus (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 63
4.27 Elytra and Aedeagus of Asbolodes humerosus (a) Elytra. (b)
Aedeagus 64
4.28 Antenna and mouth part of Pseudonautes vagevittatus (a)
Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 66
4.29 Thorax of Pseudonautes vagevittatus (a) Pronotum. (b) Fore leg.
(c) Midleg. (d) Hindleg. 67
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4.30 Elytra and Aedeagus of Pseudonautes vagevittatus (a) Elytra. (b) Aedeagus
68
4.31 Antenna and mouth part of Gauromaia annulipes (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 70
4.32 Thorax of Gauromaia annulipes (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 71
4.33 Elytra and Aedeagus of Gauromaia annulipes (a) Elytra. (b)
Aedeagus 72
4.34 Antenna and mouth part of Gauromaia haagi (a) Antennae. (b)
Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 74
4.35 Thorax of Gauromaia haagi (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 75
4.36 Elytra and Aedeagus of Gauromaia haagi (a) Elytra.
(b)Aedeagus 76
4.37 Antenna and mouth part of Camptobrachys sp.n. (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 79
4.38 Thorax of Camptobrachys sp.n. (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 80
4.39 Elytra and Aedeagus of Camptobrachys sp.n. (a) Elytra. (b)
Aedeagus 81
4.40 Antenna and mouth part of n.gen. (a) Antennae. (b) Labrum. (c)
Labium. (d) Maxilla. (e) Mandible. 84
4.41 Thorax of n.gen. (a) Pronotum. (b) Fore leg. (c) Midleg. (d)
Hindleg. 85
4.42 Elytra and Aedeagus of n.gen.(a) Elytra. (b) Aedeagus 86 4.43 Antenna and mouth part of Strongylium varians (a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 89
4.44 Thorax of Strongylium varians (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 90
4.45 Elytra and Aedeagus of Strongylium varians (a) Elytra. (b)
Aedeagus 91
4.46 Antenna and mouth part of Strongylium erythrocephalum (a)
Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 93
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4.47 Thorax of Strongylium erythrocephalum (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
94
4.48 Elytra and Aedeagus of Strongylium erythrocephalum (a) Elytra.
(b) Aedeagus 95
4.49 Antenna and mouth part of Strongylium orientale(a) Antennae.
(b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 97
4.50 Thorax of Strongylium orientale (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 98
4.51 Elytra and Aedeagus of Strongylium orientale (a) Elytra. (b)
Aedeagus 99
4.52 Antenna and mouth part of Strongylium n.sp.(a) Antennae. (b)
Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 101
4.53 Thorax of Strongylium n.sp. (a) Pronotum. (b) Fore leg. (c)
Midleg. (d) Hindleg. 102
4.54 Elytra and Aedeagus of Strongylium n.sp.(a) Elytra. (b)
Aedeagus 103
4.55 Antenna and mouth part of Strongylium forcipicolle (a)
Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 105
4.56 Thorax of Strongylium forcipicolle (a) Pronotum. (b) Fore leg.
(c) Midleg. (d) Hindleg. 106
4.57 Elytra and Aedeagus of Strongylium forcipicolle (a) Elytra. (b)
Aedeagus 107
4.58 Antenna and mouth part of Leprocaulus rotundicollis (a)
Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 110
4.59 Thorax of Leprocaulus rotundicollis (a) Pronotum. (b) Fore leg.
(c) Midleg. (d) Hindleg. 111
4.60 Elytra and Aedeagus of Leprocaulus rotundicollis (a) Elytra. (b)
Aedeagus 112
4.61 Antenna and mouth part of Derosphaerus aeruginosus (a)
Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible. 114
4.62 Thorax of Derosphaerus aeruginosus (a) Pronotum. (b) Fore leg.
(c) Midleg. (d) Hindleg. 115
4.63 Elytra and Aedeagus of Derosphaerus aeruginosus (a) Elytra.
(b) Aedeagus 116
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4.64 Antenna and mouth part of Pseudoblaps javana (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
118
4.65 Thorax of Pseudoblaps javana (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
119
4.66 Elytra and Aedeagus of Pseudoblaps javana (a) Elytra. (b) Aedeagus
120
4.67 Antenna and mouth part of Amarygmus metallicus (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
122
4.68 Thorax of Amarygmus metallicus (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
123
4.69 Elytra and Aedeagus of Amarygmus metallicus (a) Elytra. (b) Aedeagus
124
4.70 Antenna and mouth part of Amarygmus aeneolus (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
126
4.71 Thorax of Amarygmus aeneolus (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
127
4.72 Elytra and Aedeagus of Amarygmus aeneolus (a) Elytra. (b) Aedeagus
128
4.73 Antenna and mouth part of Gonocephalum hispidocostatum (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
130
4.74 Thorax of Gonocephalum hispidocostatum (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
131
4.75 Elytra and Aedeagus of Gonocephalum hispidocostatum (a) Elytra. (b) Aedeagus
132
4.76 Antenna and mouth part of Uloma sextuberosa (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
135
4.77 Thorax of Uloma sextuberosa (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
136
4.78 Elytra and Aedeagus of Uloma sextuberosa (a) Elytra. (b) Aedeagus
137
4.79 Antenna and mouth part of Uloma excise (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
139
4.80 Thorax of Uloma excise (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
140
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4.81 Elytra and Aedeagus of Uloma excise (a) Elytra. (b) Aedeagus 141
4.82 Antenna and mouth part of Alphitobius diaperinus (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
144
4.83 Thorax of Alphitobius diaperinus (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
145
4.84 Elytra and Aedeagus of Alphitobius diaperinus (a) Elytra. (b) Aedeagus
146
4.85 Antenna and mouth part of Tribolium confusum (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
149
4.86 Thorax of Tribolium confusum (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
150
4.87 Elytra and Aedeagus of Tribolium confusum (a) Elytra. (b) Aedeagus
151
4.88 Antenna and mouth part of Tribolium castanum (a) Antennae. (b) Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
153
4.89 Thorax of Tribolium castanum (a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
154
4.90 Elytra and Aedeagus of Tribolium castanum (a) Elytra. (b) Aedeagus
155
4.91 Antenna and mouth part of Hemicera n.sp.(a) Antennae. (b)Labrum. (c) Labium. (d) Maxilla. (e) Mandible.
158
4.92 Thorax of Hemicera n.sp.(a) Pronotum. (b) Fore leg. (c) Midleg. (d) Hindleg.
159
4.93 Elytra and Aedeagus of Hemicera n.sp.(a) Elytra. (b) Aedeagus 160
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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
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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 materials 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 family 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)
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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 conduct 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 presented while Chapter 4 postulated the construction of the keys 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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REFERENCES Aalbu, R. L. (2005, b). The pimeliine tribe Cryptoglossini: classification, biology and
inferred phylogeny (Coleoptera: Tenebrionidae). InAnnales Zoologici (Vol. 55, No. 4, pp. 677-756). Museum and Institute of Zoology, Polish Academy of Sciences.
Aalbu, R. L. (2006). where are we at: Assessing the currents state of tenebrionidae
systematics on a global scale (Coleoptera: Tenebrionidae).Cahiers scientifiques-Muséum d'histoire naturelle de Lyon, (10), 55-70.
Aalbu, R. L., Triplehorn, C. A., Campbell, J. M., Brown, K. W., Somerby, R. E., &
Thomas, D. B. (2002). Tenebrionidae Latreille 1802 [pp. 463–509].American beetles, 2.
Aalbu., R. L., Triplehorn, C. A., Campbell, J. M., Brown, K. W., Somerby, R. E. &
Thomas, D. B. (2002 b). 106. Tenebrionidae Latreille 1802. Pp. 463–509 in Arnett, R. H., Jr., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Gainesville, Florida.
Abu-Asab, M. S., Peterson, P. M., Shetler, S. G., & Orli, S. S. (2001). Earlier plant
flowering in spring as a response to global warming in the Washington, DC, area. Biodiversity & Conservation, 10(4), 597-612.
Allsopp, P. G. (1979). Identification of false wireworms (Coleoptera: Tenebrionidae)
from southern Queensland and northern New South Wales. – Journal of the Australian Entomological Society 18: 277–286.
Allsopp, P. G. (1980 a). The biology of false wireworms and their adults (soil-
inhabiting Tenebrionidae) (Coleoptera): a review. – Bulletin of Entomological Research 70 (3): 343–379.
Allsopp, P. G. (1981). Development, longevity and fecundity of the false wireworms
Pterohelaeus darlingensis and P. alternatus (Coleoptera: Tenebrionidae). 1. Effect of constant temperature. – Australian Journal of Zoology 29 (4): 605–619.
Allsopp, P. G. (1982). Development, longevity and fecundity of the false wireworms
Pterohelaeus darlingensis and P. alternatus (Coleoptera: Tenebrionidae) 2. Effect of food type. – Australian Journal of Zoology 30 (2): 233–242.
Allsopp, P. G. (1986 a). Development, longevity and fecundity of the false wireworms
Pterohelaeus darlingensis and P. alternatus (Coleoptera: Tenebrionidae) 4. Models for larval and pupal development under fluctuating temperatures. – Australian Journal of Zoology 34 (6): 815–825.
© COPYRIG
HT UPM
170
Allsopp, P. G. (1986 b). Identification of an additional false wirewormCestrinus trivialis Erichson (Coleoptera: Tenebrionidae) from south Queensland. – Journal of the Australian Entomological Society 25: 137–140.
Allsopp, P. G., Adams, G. D. & Butler, D. G. (1980). Response of larvae of
Pterohelaeus darlingensis Carter (Coleoptera: Tenebrionidae) to moisture, temperature and gravity. – Journal of the Australian Entomological Society 19 (4): 281–285.
Ardoin, P. (1961). Contribution à l’étude des ténébrionides malgaches. Deux
nouveaux genres d’Adeliini malgaches. – Bulletin de l’Académie Malgache (N.S.) 37: 31–33.
Arentt, R. H. (1963). The beetles of the United States. Catholic Univ. of America
Press, Washington, D.C. (No. 595.76 A75). Bates, F. (1868). Descriptions of New Genera and Species of
Heteromera. Transactions of the Royal Entomological Society of London,16(4), 309-326.
Beutel, R. G., & Leschen, R. A. B. (2005). Handbook of Zoology: A Natural History
of the Phyla of the Animal Kingdom. Volume IV Arthropoda: Insecta, Part 38. Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim).
Bouchard, P., & Löbl, I. (2008). Case 3401 Delognatha Lacordaire, 1859 (Insecta,
Coleoptera): proposed conservation. Bulletin of Zoological Nomenclature, 65, 3.
Borchmann, F. (1910 a): Coleopterorum Catalogus. Pars 2. Nilionidae, Othniidae,
Aegialitidae, Petriidae, Lagriidae. 32 pp. W. Junk, Berlin. Borchmann, F. (1910 b): Coleopterorum Catalogus. Pars 3. Alleculidae. 80 pp. W.
Junk, Berlin. Bouchard, P., Lawrence, J. F., Davies, A. E., & Newton, A. F. (2005, b). Synoptic
classification of the world Tenebrionidae (Insecta: Coleoptera) with a review of family-group names. In Annales Zoologici (Vol. 55, No. 4, pp. 499-530). Museum and Institute of Zoology, Polish Academy of Sciences.
Bouchard, P., Löbl, I., & Merkl, O. (2007, September). Nomenclatural notes on
tenebrionid beetles of the Palaearctic Region (Insecta: Coleoptera). InAnnales zoologici (Vol. 57, No. 3, pp. 385-394). Museum and Institute of Zoology, Polish Academy of Sciences.
Boving, A. G., & Craighead, F. C. (1931). Illustrated synopsis of the principal larval
forms of the order Coleoptera.
© COPYRIG
HT UPM
171
Brendell, M. J. D. 1975. Hand books for the identification of British insects. London published by the society and sold at its rooms 41, queens gate, S.W. Vol. V. Part 10 .
Brown, J. W. (2001). Species turnover in the leafrollers (Lepidoptera: Tortricidae) of Plummers Island, Maryland: Assessing a century of inventory data. Proceedings-Entomological Society of Washington,103(3), 673-673.
Calkins, C. O., & Kirk, V. M. (1973). Distribution and movement of adult false wireworms in a wheat field. Annals of the Entomological Society of America, 66(3), 527-532.
Carl, M. (1992). A revision of the genus Micriotelus Solier , (Coleoptera –Tenebrionidae). Zoology in the middle East 69-87.
Chernei, L. S. & Fedorenko, V. P. (2006). Identification keys to the darkling beetles (Coleoptera, Tenebrionidae) of the Ukrainian fauna. 248 pp.
Crowson, R. A. (1955). The Natural Classification of the Families of Coleoptera. 187 pp. Nathaniel Lloyd, London.
Curran, L. M., Trigg, S.N., McDonald, A.K. et al. (2004) “Lowland Forest Loss in Protected Areas of Indonesian Borneo.” Science, 303 (5660), 1000–1003. doi:10.1126/science.1091714.
Dobie, P. (1991). Host-plant resistance to insects in stored cereals and legumes. Ecology and Management of Food-Industry Pests, 373-383.
Doyen, J. T. & Tschinkel, W. R. (1982). Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera). Syst Entomol. 7: 127–183.
Doyen, J. T. (1972). Familial and subfamilial classification of the Tenebrionoidea (Coleoptera) and a revised generic classification of the Coniontini (Tentyriidae). Quaestiones entomol. 8: 357–376.
Doyen, J. T., & Lawrence, J. F. (1979). Relationships and higher classification of some Tenebrionidae and Zopheridae (Coleoptera).Systematic entomology, 4(4), 333-377.
Doyen, J. T., & TSCHINKEL, W. R. (1982). Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera). Systematic entomology, 7(2), 127-183.
Doyen, J. T., Matthews, E. G. & Lawrence, J. F. (1990). Classification and annotated checklist of the Australian genera of Tenebrionidae (Coleoptera).– Invertebrate Taxonomy 3 (1989): 229–260.
Drinkwater, T. W. (1999). Regional distribution of Gonocephalum species (Coleoptera: Tenebrionidae) in maize fields in South Africa. African entomology, 7(2), 183-187.
© COPYRIG
HT UPM
172
Drinkwater, T. W., Giliomee, J. H., & Pringle, K. L. (1990). The geographical distribution of false wireworms, Somaticus species (Coleoptera: Tenebrionidae) associated with maize in South Africa. Journal of the Entomological Society of Southern Africa, 53(2), 127-136.
Drinkwater, T. W., Giliomee, J. H., & Pringle, K. L. (1996). Development rates of the
false wireworm, Somaticus angulatus (Faehraeus)(Coleoptera: Tenebrionidae), at different temperatures. African entomology, 4, 21-24.
Drinkwater, T.W.(1989). Occurrence of soil-inhabiting Tenebrionidae (Coleoptera) in
maize fields in South Africa. – Phytophylactica 21 (4): 397–398. Endrödy-Younga, S. & Schawaller, W. (2002). First record of Laenini (Coleoptera:
Tenebrionidae) from South Africa, with descriptions of two genera and several new species. – Annals of the Transvaal Museum 39: 9–21
Erwin, T. L. (1981). The ground beetles of a temperate forest site (Coleoptera:
Carabidae): An analysis of fauna in relation to size, habitat selection, vagility, seasonality, and extinction. Bulletin of the Biological Society of Washington, 5, 104-224.
Fairmaire (1899). et description d’une nouvelle espèce de Borneo (Coleoptera,
Tenebrionidae). Nouv. Rev. Ent. (N. S.) 17(3): 233–244 Fernald, H. T. and Sheparo, H. H. (1955). Applied Entomology An Introduce text
book of insect in there relations to man. Mc Grawltill book company, Inc. New xork. 385 p.
Ferrer, J. and Degard, O. (2005). New species of darkling beetles from central
America with systematic notes (Coleoptera:Tenebrionidae ). Annales Zoologici, (Warszawa), 55(4): 633-661
Ferrer, J. (1995). Contribution to the knowledge of the Tenebrionidae of Somalia
(Coleoptera). Frustula Entomologica, Nuova, Serie, 18, 1-76. Ferrer, J. (2006). Constitution du groupe indo-africain des Falsocossyphini, tribus
nova, et description d'un nouveau genre hypogee du Vietnam. Coleoptera, Tenebrionidae. Cahiers scientifiques-Muséum d'histoire naturelle de Lyon, (10), 75-83.
Ferrer, J., & Moragues, G. (2000). Révision des espèces appartenant au genre
Pheugonius Fairmaire (1899). et description d’une nouvelle espèce de Borneo (Coleoptera, Tenebrionidae). Nouv. Rev. Ent.(NS), 17(3), 233-244.
Freeman, I. (1980). Common insect pest of stored food productos . 6th ed. Br. T. M.
(Nat. Hist.) Econ. Ser. No. 15. 100 Gall, A. (1980) Are lesser mealworms worth the trouble they cause. Poultry Digest
Feb: 76/77.
© COPYRIG
HT UPM
173
Gebien, H. (1937). Katalog der Tenebrioniden. Teil 3, Unterfamilie Asidinae.Publicazioni del Museo entomologico “Pietro Rossi, 663-791.
Gebien, H. (1944). Katalog der Tenebrioniden (Coleoptera heteromera). Verlag J. Pfeiffer.
Grimm, R. (2004 a). Die Arten der Gattung Ulomina Baudi di Selve, mit Beschreibungen von drei neuen Arten aus Laos und Thailand (Coleoptera: Tenebrionidae: Palorinae). – Stuttgarter Beiträge zur Naturkunde. Ser.A664: 1–10.
Grimm, R. (2004 b). Neve Arten der Gallung Palorinus Blair, 1930 und Paloropsis Masumoto et Grimm, 2004 aus Indien und Thailand (Coleoptera, Tenebrionidae, Palorinae). – Entomologica Basiliensia 26: 293–298.
Grimm, R. (2008). Guanobius borneensis n. gen., n. sp. from Borneo (Coleoptera: Tenebrionidae: Alphitobiini). Stuttgarter Beiträge zur Naturkunde A, Neue Serie, 1, 375-379.
Halstead, D. G. H. (1967): A revision of the genus Palorus (sens. lat.) (Coleoptera: Tenebrionidae). – Bulletin of the British Museum (Natural History), Entomology 19: 61–148.
Hill, D. S. (2008). Pests of crops in warmer climates and their control. Springer Science & Business Media.
Hiremath, I. G., Lingappa, S., Viraktamath, C. A., & Ali, T. M. M. (1986). Occurrence of the ground beetle, Gonocephalum hofmannseggi (Coleoptera: Tenebrionidae) on finger millet (Eleusine coracana (L.) Gaertn). Journal of soil biology & ecology. 6 (1): 62–66.
Hoffman, R. L., & Steiner, W. E. (2005). Jadera haematoloma, another insect on its way north (Heteroptera: Rhopalidae). Banisteria, 26, 7-10.
Horion, A. (1956). Faunistik der mitteleuropäischen Käfer. Band V. Heteromera.. Museum G. Frey, Tutzing bei München. xv + 336 pp.
Imms, A. D. 1964. A General Textbook of Entomology. Ninth Edition. London: Methuen and Co. Ltd. 886 pp.
Junk, W., & Schenkling, S. (1910). Coleopterorum catalogus. W. Junk.
Junk,W.(1911).Coleopterorum Calalogus.Berin W. I5, Kurfuerstendamm 20I. 24. III. 1911.
Kaszab, Z. (1980). Neue Tenebrioniden (Coleoptera) aus Sri Lanka. I. II. – Acta Zoologica Academiae Scientiarum Hungaricae 26 (1–3, 4): 123–196, 285–375.
© COPYRIG
HT UPM
174
Kaszab, Z. (1985). Beiträge zur Kenntnis der Pazifischen Tenebrioniden (Coleoptera). Folia Entomol. Hung, 46(1), 23-63.
Kaszab, Z.(1979): Tenebrioniden aus Südindien (Coleoptera). – Acta Zoologica
Academiae Scientiarum Hungaricae 25 (3–4): 259–310. Kobayashi, T., & Aguero, G. D. (1988). Singular occurrence of soybean insect pests
and the control of them under the severe drought condition in Paraguay. JARQ, Japan Agricultural Research Quarterly, 22(2), 157-160.
Koch, C. (1955). Monograph of the Tenebrionidae of southern Africa (No. 7).
Transvaal Museum. Koh, L.P., Wilcove D.S. (2008) Is oil palm agriculture really destroying tropical
biodiversity? Conserv Lett 1, 60–64. Krombein, K. V. (1963). Annotated list of the wasps (Hymenoptera): Bethyloidea,
Scolioidea, Vespoidea, Pompiloidea, Sphecoidea). Natural history of Plummers Island, Maryland. Proceedings of the Biological Society of Washington 76:255–280
Lacordaire, T. (1859). Rhipiphorides. Histoire naturelle des insectes. Genera des
Coldopteres. Librairie Encyclop6dique de Roret, Paris, 5(pt 1), 616-633. Latreille, P. A. (1802). Histoire naturelle, générale et particulière des crustacés et des
insectes. Familles naturelles des genres. Vol. 3. 387 pp. 16–37 pls. F. Dufart, Paris.
Lawrence, J. F. (1994). The larva of Sirrhas variegatus, sp. nov., with notes on the
Perimylopidae, Ulodidae (stat. nov.), Zopheridae and Chalcodryidae (Coleoptera: Tenebrionoidea). Invertebrate Systematics, 8(2), 329-349.
Lawrence, J. F., & Slipinski, S. A. (1999). Phylogeny and classification of Zopheridae
sensu novo (Coleoptera: Tenebrionoidea) with a review of the genera of Zopheridae (excluding Monommatini). In Annales zoologici (Vol. 49, pp. 1-53).
Lawrence, J. F., Newton, A. F., Pakaluk, J., & Slipinski, S. A. (1995). Families and
Subfamilies of Coleoptera: With Selected Genera, Notes, References and Data on Family-group Names. Museum i Instytut Zoologii PAN.
LeConte, J. L., Horn, G. H., & Henshaw, S. (1883). Classification of the coleoptera
of North America (Vol. 26, No. 4). Smithsonian institution. Lillig, M., & Pavlíček, T. (2003). The Darkling Beetles of the Sinai Peninsula:
Coleoptera: Tenebrionidae (excl. Lagriinae Et Alleculinae) (Vol. 1). Kasparek Verlag.
© COPYRIG
HT UPM
175
Lillig, M., Barthet, H. B., & Mifsud, D. (2012). An identification and informative guide to the Tenebrionidae of Malta (Coleoptera). Bulletin of the entomological Society of malta, 5, 121-160.
Löbl, I., Ando, K., Bouchard, P., Egorov, L. V., Iwan, D., Lillig, M., ... & Petterson, R. (2008). Family Tenebrionidae Latreille, 1802. Catalogue of Palaearctic Coleoptera, 5, 105-352.
Martin, O. Y., Gubler, A., Wimmer, C., Germann, C., & Bernasconi, M. V. (2012). Infections with Wolbachia and Spiroplasma in the Scathophagidae and other Muscoidea. Infection, Genetics and Evolution, 12(2), 315-323.
Masumoto, K (1996). Fourteen new Laena (Coleoptera, Tenebrionidae) from China, Vietnam and Thailand.– Bulletin of the National Science Museum, Series A, 22 (3): 165–178.
Masumoto, K (2001). Anew genus and species of the Tribe Adeliini (Coleoptera, Tenebrionidae) from the Upper Hypogean Zone of Southwest China. – Journal of the Speleological Society of Japan 26: 44–49.
Masumoto, K. & Grimm, R. (2004). A new genus and species of the Palorinae (Coleoptera: Tenebrionidae) from Japan. – Entomological Review of Japan 59 (1): 127 – 130.
Masumoto, K.(1990): New Himalayan species of Laena (Coleoptera, Tenebrionidae) preserved in the collection of the National Science Museum, Tokyo. – Bulletin
Matthews, E. G. & Bouchard, P. (2007). Australian Faunal Directory. Checklist for Tenebrionidae Coleoptera. (http://www.environment.gov.au/cgibin/ abrs/fauna/details.plpstrVol=TENEBRIONIDAE;pstrTaxa=131;pstrChecklistMode=1). (Accessed 30 October 2008.)
Matthews, E. G. (1986). A revision of the troglophilic genus Brises Pascoe, with a discussion of the Cyphaleini (Coleoptera, Tenebrionidae). Records of the South Australian Museum, 19(6), 77-90.
Matthews, E. G. (1998). Classification, phylogeny and biogeography of the genera of Adeliini (Coleoptera: Tenebrionidae). Invertebrate Taxonomy, 12, 685-824.
Matthews, E. G. (2003 b). The Palorus group – a new subfamily of Tenebrionidae. – Spixiana 26: 49–55.
Matthews, E. G., & Bouchard, P. (2008). Tenebrionid beetles of Australia: description of tribes, keys to genera, catalogue of species. Australian Biological Resources Study. viii + 398 pp.
Matthews, E. G., & Bouchard, P. (2008). Tenebrionid beetles of Australia: description of tribes, keys to genera, catalogue of species. Australian Biological Resources Study. . viii + 398 pp.
© COPYRIG
HT UPM
176
Mccolloch, J. W. (1918). Notes on false wireworms with especial reference to Eleodes tricostata Say. Journal of Economic Entomology, 11, 212-224.
Mccolloch, J. W. (1919). Eleodes opaca Say, an important enemy of wheat in the Great
Plains area. Journ. Econ. Ent, 12, 183-194. Medvedev, G. S. (1968). Coleoptera. Darkling beetles (Tenebrionidae). Subfamily
Opatrinae. Tribes Platynotini, Dendarini, Pedinini, Dissonomini, Pachypterini, Opatrini (part) and Heterotarsini, 19(Part 2), 1-285.
Medvedev, G. S. (1973). position of the genera Leichenum Dej. and Idisia
Pask.(Coleoptera, Tenebrionidae) in the system, and the description of new genus from the northern Karakums. Entomological review.
Medvedev, G. S. (2006). To the systematics and nomenclature of tenebrionid beetles
of the tribes Phaleriini, Lachnogyini, Klewariini, and Blaptini (Coleoptera, Tenebrionidae). Entomological Review, 86(7), 820-839.
Nabozhenko, M. V. (2001). On the classification of the tenebrionid tribe Helopini,
with a review of the genera Nalassus Mulsant and Odocnemis Allard (Coleoptera, Tenebrionidae) of the European part of CIS and the Caucasus. Entomological Review, 81(8), 909-942.
Pascoe, F. P. (1869). Descriptions of New Genera and species of Tenebrionidae from
Australia and Tasmania.– Annals and Magazine of Natural History (4) 3:29–45.
Peck, S. B. (2005). A checklist of the beetles of Cuba with data on distributions and
bionomics. Insecta, Coleoptera. Arthrropods of Florida and neighboring land areas, 18, 1-214.
Peck, S. B.(2006). The Beetles of the Galápagos Islands, Ecuador: Evolution, Ecology,
and Diversity (Insecta: Coleoptera). 314 pp. NRC Research Press, Ottawa, Ontario, Canada.
Reddy, M. V., Kumar, V. K., & Reddy, B. K. (1992). Occurrence of Gonocephalum
sp.(Tenebrionidae: Coleoptera) as a pest of groundnut in India. Tropical Pest Management 38 (1): 108.
Reitter, E. (1917). Bestimmungs-Tabelle der palaearktischer Coleopteren. 82.Heft:
Tenebrionidae, Unterfamilie Asidini. Sonderabdruck aus dem, 55, 1-74. Robertson, L. N., & Simpson, G. B. (1989). Sampling and dispersion of Pterohelaeus
alternatus Pascoe and Gonocephalum macleayi (Blackburn)(Coleoptera: Tenebrionidae) larvae in soil. Queensland Journal of Agricultural and Animal Sciences, 45(2), 189-193.
Schawaller, W. (1992). Leochrini (Coleoptera: Tenebrionidae) aus dem Nepal-
Himalaya. Stuttgarter Beitr. zur Naturkunde. Ser. A, Biologie. 482: 1–15.
© COPYRIG
HT UPM
177
Schawaller, W. (1996). Tenebrionidae (Coleoptera) aus Nord-Griechenland: Habirare, Artengesellschaften und Verbreitung. Entomologische Blatter fur Biologie und Systematik der Kafer, 92(1), 3-17.
Schawaller, W. (1996). Tenebrionidae (Coleoptera) aus Nord-Griechenland: Habitate,
Artengeselschaften und Verbreitung. Entomologische Blätter, 92: 3-18. Schawaller, W. (2001a). Nepalolaena kira gen. nov., sp. nov. from the Nepal
Himalayas (Coleoptera, Tenebrionidae). Entomologica Basiliensia 23: 277–282.
Schawaller, W. (2002). The genus Laena Latreille in Nepal, with taxonomic and
faunistic notes on species from the adjacent Himalayas (Coleoptera: Tenebrionidae). Staatl. Museum für Naturkunde. Serie A (Biologie) 641: 1–69.
Schawaller, W. (2008). The genus Laena Latreille (Coleoptera: Tenebrionidae) in
China (part 2), with descriptions of 30 new species and a new identification key. Stuttgarter Beiträge zur Naturkunde Serie A (Biologie), 1, 387-411.
Schawaller, W.(1998 a). Borneolaena gen. n. riedeli sp. n. from Sarawak, the first
species of Laenini (Coleoptera: Tenebrionidae) from the Sunda Islands. – Stuttgarter Beiträge zur Naturkunde, Ser. A (Biologie) 575: 1–8.
Skopin, N. G. (1964). Die Larven der Tenebrioniden des Tribus Pycnocerini
(Coleoptera Heteromera). Koninklijk Museum voor Midden-Afrika. Sciences Zoologiques 127: 1–35, 16 pls.
Snodgrass, R. E. (1935). Principles of insect morphology. McGraw-HillBook
company , Inc. New York and London 664 pp. Soldati, F., & Soldati, L. (2003). Réactualisation de la liste systématique des
Coléopteres Tenebrionidae (Alleculinae exclus) de France continentale et de Corse. Bulletin mensuel de la Société linnéenne de Lyon,72(10), 331-349.
Sodhi, N.S., Koh L.P., Brook B.W., Ng P.K.L. (2004) Southeast Asian biodiversity:
an impending disaster. Trends Ecol Evol 19, 654–660. Steiner, W. E. (1999). Flightless beetles in Appalachian “deserts”: studies on the
distribution and localized habitats of some species of Tenebrionidae (Coleoptera). Virginia Museum of Natural History Special Publication, 7, 125-144.
Swenk, M. H. (1909). Eleodes as an enemy of planted grain. Journal of Economic
Entomology, 2(5), 332-336. Tschinkel, W. R., & Doyen, J. T. (1980). Comparative anatomy of the defensive
glands, ovipositors and female genital tubes of tenebrionid beetles (Coleoptera). International Journal of Insect Morphology and Embryology,9(5), 321-368.
© COPYRIG
HT UPM
178
Wade, J. S. (1921). Notes on ecology of injurious Tenebrionidae (Col.). Ent. News, 32, 1-6.
Wade, J. S., & ST George, R. A. (1923). Biology of the false wireworm.Eleodes
suturalis. Watt, J. C. (1967). A review of classifications of Tenebrionidae
(Coleoptera).Entomologist's Monthly Magazine, 102(1966), 80-86. Watt, J. C. (1974). A revised subfamily classification of Tenebrionidae
(Coleoptera). New Zealand journal of zoology, 1(4), 381-452. Watt, J. C. (1987). The family and subfamily classification and New Zealand genera
of Pythidae and Scraptiidae (Coleoptera). Systematic entomology, 12(1), 111-136.
Zhang, D. Z., YU, Y. Z., & Ren, J. Z. (2003). The biological characters and behavior
of Mantichorula semenowi [J]. Entomological Knowledge, 4, 011.