Rhythms of Insect Evolution

DONG REN, CHUNGKUN SHIH, TAIPING GAO, YONGJIE WANG and YUNZHI YAO are all professors in the College of Life Sciences, Capital Normal University, Beijing, China. Dr. Shih is also a volunteer Research Associate in the National Museum of Natural History, Smithsonian Institution, Washington, D.C., USA.

Preface xi

List of Contributors xiii

Acknowledgements xvii

1 Jurassic-Cretaceous Non-Marine Stratigraphy and Entomofaunas in Northern China 1
Dong Ren

1.1 Introduction 1

1.2 Non-marine Jurassic and Cretaceous Insect Fossil-Bearing Lithostratigraphic Division and Correlation in Northern China 1

1.2.1 Yumen-Jiuquan Basin in Gansu Province 1

1.2.2 Intermountain Volcanic Basins in Beijing-Northern Hebei-Western Liaoning-Southeastern Inner Mongolia 3

1.3 Non-marine Jurassic and Cretaceous Entomofaunas in Northern China 4

1.3.1 Yanliao Entomofauna 5

1.3.2 Jehol Entomofauna in the Yanliao Area 8

1.3.3 Fuxin Entomofauna 10

1.4 Geological Ages of Non-marine Jurassic and Cretaceous Strata and Entomofaunas in Northern China 10

References 12

2 Coexisting Animals and Plants in the Ecosystems 17
Chungkun Shih, Taiping Gao, and Dong Ren

2.1 Introduction 17

2.2 Representative Fossils of Coexisting Animals 17

2.3 Representative Fossils of Coexisting Plants 24

References 28

3 Insects – In the Spotlight 31
Taiping Gao, Chungkun Shih, and Dong Ren

3.1 Introduction to Insects 31

3.2 How to Identify an Insect 31

3.3 Origin and Evolution of Insects 35

References 38

4 A History of Paleoentomology in China 41
Dong Ren, Chungkun Shih, and Taiping Gao

4.1 Introduction 41

4.2 Early Foundational Studies (1923–1935) 41

4.3 Early Taxonomic Studies (1965–1985) 42

4.4 Major Taxonomic Studies (1985–Present) 43

4.5 Phylogenetic and Paleobiological Studies (1991–Present) 44

4.6 International Cooperative Studies (2000–Present) 45

References 48

5 Ephemeroptera – Mayflies 51
MeiWang, Qingqing Lin, Chungkun Shih, and Dong Ren

5.1 Introduction to Ephemeroptera 51

5.2 Progress in the Studies of Fossil Ephemeroptera 53

5.3 Representative Fossils of Ephemeroptera from Northern China 53

References 59

6 Odonata – Dragonflies and Damselflies 63
Qiang Yang, Dong Ren, Hong Pang, and Chungkun Shih

6.1 Introduction to Odonata 63

6.2 Progress in the Studies of Fossil Odonata 66

6.3 Representative Fossils of Odonata from Northern China 66

References 86

7 Blattaria – Cockroaches 91
Junhui Liang, Chungkun Shih, and Dong Ren

7.1 Introduction to Blattaria 91

7.2 Progress in the Studies of Fossil Blattodea 92

7.3 Representative Fossils of Blattaria from Northern China 93

References 109

8 Termitoidae – Termites 113
Zhipeng Zhao, Dong Ren, and Chungkun Shih

8.1 Introduction to Termitoidae 113

8.2 Progress in the Studies of Fossil Termites 115

8.3 Representative Fossils of Termites from Northern China 116

References 117

9 Orthoptera – Grasshoppers and Katydids 121
Jun-Jie Gu, Chungkun Shih, and Dong Ren

9.1 Introduction to Orthoptera 121

9.2 Progress in the Studies of Fossil Orthoptera 122

9.3 Representative Fossils of Orthoptera from Northern China 124

References 134

10 Notoptera – Rock Crawlers and Ice Crawlers 137
Yingying Cui, Chungkun Shih, and Dong Ren

10.1 Introduction to Notoptera (Mantophasmatodea and Grylloblattodea) 137

10.2 Progress in the Studies of Fossil Notoptera (Mantophasmatodea and Grylloblattodea) 138

10.3 Representative Fossils of Notoptera (Mantophasmatodea and Grylloblattodea) from Northern China 139

References 145

11 Dermaptera – Earwigs 149
Mingyue Ren, Chungkun Shih, Changyue Xing, and Dong Ren

11.1 Introduction to Dermaptera 149

11.2 Progress in the Studies of Fossil Dermaptera 150

11.3 Representative Fossils of Dermaptera from Northern China 150

References 155

12 Chresmodidae –Water-Walking Insects 157
Chaofan Shi, Chungkun Shih, and Dong Ren

12.1 Introduction to Chresmodidae 157

12.2 Progress in the Studies of Fossil Chresmodidae 158

12.3 Representative Fossils of Chresmodidae from Northern China 159

References 162

13 Phasmatodea – Stick Insects and Leaf Insects 165
Chaofan Shi, Chungkun Shih, Sha Chen, and Dong Ren

13.1 Introduction to Phasmatodea 165

13.2 Progress in the Studies of Fossil Phasmatodea 166

13.3 Representative Fossils of Phasmatodea from Northern China 168

References 172

14 Plecoptera – Stoneflies 175
Yingying Cui, Chungkun Shih, and Dong Ren

14.1 Introduction to Plecoptera 175

14.2 Progress in the Studies of Fossil Plecoptera 176

14.3 Representative Fossils of Plecoptera from Northern China 176

References 183

15 Psocoptera – Barklice and Booklice 185
RuiqianWang, Yunzhi Yao, Dong Ren, and Chungkun Shih

15.1 Introduction to Psocoptera 185

15.2 Progress in the Studies of Fossil Psocoptera 185

15.3 Representative Fossils of Psocoptera from Northern China 186

References 187

16 Homoptera – Cicadas and Hoppers 189
Ying Wang, Xiao Zhang, Tingying Zhang, Xue Liu, Chungkun Shih, Yunzhi Yao, and Dong Ren

16.1 Introduction to Homoptera 189

16.2 Progress in the Studies of Fossil Homoptera 190

16.3 Representative Fossils of Homoptera from Northern China 192

References 218

17 Heteroptera – True Bugs 225
Sile Du, Shan Lin, Chungkun Shih, Dong Ren, and Yunzhi Yao

17.1 Introduction to Heteroptera 225

17.2 Progress in the Studies of Fossil Heteroptera and Fossil Coleorrhyncha 227

17.3 Representative Fossils of Heteroptera from Northern China 228

References 262

18 Megaloptera – Dobsonflies, Fishflies, and Alderflies 269
Yongjie Wang, Chungkun Shih, and Dong Ren

18.1 Introduction to Megaloptera 269

18.2 Progress in the Studies of Fossil Megaloptera 269

18.3 Representative Fossils of Megaloptera from Northern China 271

References 273

19 Raphidioptera – Snakeflies 275
Hui Fang, Yongjie Wang, Dong Ren, and Chungkun Shih

19.1 Introduction to Raphidioptera 275

19.2 Progress in the Studies of Fossil Raphidioptera 275

19.3 Representative Fossils of Raphidioptera from Northern China 276

References 282

20 Neuroptera – Lacewings 285
Zhenzhen Chen, Shuo Huang, Yu Chang, Yongjie Wang, Chungkun Shih, and Dong Ren

20.1 Introduction to Neuroptera 285

20.2 Progress in the Studies of Fossil Neuroptera 287

20.3 Representative Fossils of Neuroptera from Northern China 290

References 32920.1 Introduction to Neuroptera 285

20.2 Progress in the Studies of Fossil Neuroptera 287

20.3 Representative Fossils of Neuroptera from Northern China 290

References 329

21 Coleoptera – Beetles 337
Yali Yu, Zhenhua Liu, Chungkun Shih, and Dong Ren

21.1 Introduction to Coleoptera 337

21.2 Progress in the Studies of Fossil Coleoptera 342

21.3 Representative Fossils of Coleoptera from Northern China 344

References 414

22 Hymenoptera – Sawflies and Wasps 429
Mei Wang, Longfeng Li, Chungkun Shih, Taiping Gao, and Dong Ren

22.1 Introduction to Hymenoptera 429

22.2 Progress in the Studies of Fossil Hymenoptera 435

22.3 Representative Fossils of Hymenoptera from Northern China 436

References 490

23 Diptera – True Flies with Two Wings 497
Ye Han, Xiuna Ye, Cuiping Feng, Kuiyan Zhang, Chungkun Shih, and Dong Ren

23.1 Introduction to Diptera 497

23.2 Progress in the Studies of Fossil Diptera 500

23.3 Representative Fossils of Diptera from Northern China 501

References 546

24 Mecoptera – Scorpionflies and Hangingflies 555
Xiaodan Lin, Chungkun Shih, Sheng Li, and Dong Ren

24.1 Introduction to Mecoptera 555

24.2 Progress in the Studies of Fossil Mecoptera 558

24.3 Representative Fossils of Mecoptera from Northern China 559

References 589

25 Siphonaptera – Fleas 597
Taiping Gao, Chungkun Shih, and Dong Ren

25.1 Introduction to Siphonaptera 597

25.2 Progress in the Studies of Fossil Fleas 598

25.3 Representative Fossils of Siphonaptera from Northern China 600

References 605

26 Trichoptera – Caddisflies 607
Mei Wang,Weiting Zhang, Chungkun Shih, and Dong Ren

26.1 Introduction to Trichoptera 607

26.2 Progress in the Studies of Fossil Trichoptera 607

26.3 Representative Fossils of Trichoptera from Northern China 608

References 616

27 Lepidoptera – Butterflies and Moths 619
Weiting Zhang, Chungkun Shih, and Dong Ren

27.1 Introduction to Lepidoptera 619

27.2 Progress in the Studies of Fossil Lepidoptera 620

27.3 Representative Fossils of Lepidoptera from Northern China 622

References 629

28 Insect Feeding 631
Chungkun Shih, Taiping Gao, Yunzi Yao, and Dong Ren

28.1 Introduction 631

28.2 PollinationMutualism – Feeding on Pollination Drops Before Angiosperms 631

28.2.1 Scorpionflies with Elongate Siphonate Mouthparts 631

28.2.2 Kalligrammatid Lacewings with Elongate Siphonate Mouthparts 632

28.3 Predation – Preying on Other Insects 635

28.3.1 Mantispid Lacewings with Raptorial Forelegs 635

28.3.2 Dipteromantispidae with Raptorial Forelegs 636

28.3.3 Hangingflies – Bittacidae and Cimbrophlebiidae 636

28.4 Blood Feeding by Ectoparasite Fleas 636

28.5 Blood Feeding by True Bugs 639

28.6 Herbivores: Feeding on Plant Matter as Evidenced by Insect Damage Types 643

References 646

29 Camouflage, Mimicry or Eyespot Warning 651
Chungkun Shih, YongjieWang, and Dong Ren

29.1 Introduction 651

29.2 Camouflage by Fossil Insects 651

29.2.1 Irregular Light and Dark Patches Covering the EntireWing 651

29.2.2 Irregular Dark or Light Markings on Part of the Margin and/or Center ofWing 653

29.2.3 Dispersed Dark Spots Large or Small 654

29.2.4 Dispersed Small Light Spots or Large Light Spots Surrounded by Small Dark Spots 654

29.2.5 Regular Transverse (from Anterior to Posterior Margin) Stripes of Light and Dark Bands 654

29.2.6 Regular Longitudinal (from Base to the Apex) Stripes of Light and Dark Bands 655

29.3 Mimicry by Fossil Insects 655

29.3.1 Ancient Pinnate Leaf Mimicry among Lacewings 656

29.3.2 Mimicry and Mutualism among Hangingflies and Ginkgo Plants 657

29.4 EyespotWarning for Fossil Insects 659

29.4.1 Eyespots and Spots on the Forewings of Kalligrammatids 659

29.5 Summary and Prospects 660

References 662

30 Gene Propagation – Courtship, Mating, and Next Generation 667
Chungkun Shih, Taiping Gao, and Dong Ren

30.1 Introduction 667

30.2 Extreme Sexual Display 667

30.3 Serenade with Love Songs 668

30.4 Sensing and Locating PotentialMates with Ramified Antennae 670

30.5 Forever Love –The Hitherto Earliest Record of Copulating Insects 670

30.6 Long Ovipositors Used for Laying Eggs into Hosts 673

30.7 Breeding – Oviposition, Gall and Leaf Mining 675

References 679

Index 683