1. Introduction to head morphology of Ciidae 22 2. Material and methods 22 2.1. Specimens 22 2.2. Pictorial documentation 23 2.3. Terminology 23 2.4. Morphological directions 23 2.5. Abbreviations 24 3. Description of head of Cis chinensis 24 3.1. Sexes and male morphs 24 3.2. Head capsule 24 3.3. Clypeus, labrum and epipharynx 26 3.4. Mandibles 27 3.5. Maxillae 28 3.6. Labium and hypopharynx 29 3.7. Antennae 30 4. Discussion of head morphology 32 4.1. Notes on morphological terms and interpretations 32 4.2. Comparison with some other Ciidae 35 4.3. Comparison with Tenebrio molitor (Tenebrionidae) 37

Part II: Ciidae in Caspian-Hyrcanian Forest in Northern Iran 50

1. Introduction to Caspian-Hyrcanian forest 50 1.1. Geography of the Caspian Forest 50 1.2. History and significance of the Caspian Forest 51 1.3. Climate of the Caspian Forest 51 1.4. Geographic relationships of the Caspian Forest 51 1.5. Trees of the Caspian Forest 52 1.6. Fungi of the Caspian Forest 52 1.7. Ciidae of the Caspian Forest 53 2. Material and methods 53 2.1. Area covered and collecting localities 53 2.2. Collecting of Ciidae 54 2.3. Morphological preparation and pictorial documentation 54 2.4. Processing of images 54 2.5. Identification of fungus species 55 2.6. Identification of Ciidae species 55 2.7. Morphological data and terminologies 56 2.8. Abbreviations 56 3. Results on Ciidae in Caspian Forest 56 3.1. Survey of observed Ciidae species and their systematics 56 3.2. Survey of observed host fungus species and their systematics 59 3.3. Morphology and distinguishing characters of Ciidae 60 3.4. Results on Cis chinensis 69 3.5. Results on Cis submicans 71 3.6. Results on Cis comptus 73 3.7. Results on Cis striatulus 75 3.8. Results on Cis tomentosus 77 3.9. Results on Cis reitteri 79 3.10. Results on Cis castaneus 81 3.11. Results on Cis lugowoji 83 3.12. Results on Cis fissicollis 85 3.13. Results on Cis festivus /87 3.14. Results on Ennearthron cornutum 88 3.15. Results on Orthocis reflexicollis 90 3.16. Results on Strigocis bicornis 92 3.17. Results on Sulcacis fronticomis 94 3.18. Results on Sulcacis nitidus 96 3.19. Results on Ropalodontus baudueri 97 3.20. Results on Ropalodontusperrini 99 3.21. Results on Octotemnus rugosopunctatus 101 3.22. Results on Xylographus bostrichoides 104 3.23. Identification key to Ciidae of Caspian Forest 106 4. Discussion of Ciidae in Caspian Forest 109 4.1. Taxonomic distinctions 109 4.1.1. Cis multidentatus species group: Cis chinensis 109 4.1.2. Cis boleti species group: Cis submicans 110 4.1.3. Cis comptus species group: Cis comptus and Cis striatulus 110 4.1.4. Cis punctulatus species group: Cis tomentosus and Cis reitteri 111 4.1.5. Cis castaneus species group: Cis castaneus and Cis lugowoji 112 4.1.6. Cis fissicollis (not assigned to a species group) 113 4.1.7. Cis festivus species group: Cis festivus 113 4.1.8. Genus Ennearthron: Ennearthron cornutum 114 4.1.9. Genus Orthocis: Orthocis reflexicollis 114 4.1.10. Genus Strigocis: Strigocis bicornis 114 4.1.11. Genus Sulcacis: Sulcacis fronticomis and S. nitidus 115 4.1.12. Genus Ropalodontus: Ropalodontus baudueri and R. perrini 115 4.1.13. Genus Octotemnus: Octotemnus rugosopunctatus 115 4.1.14. Genus Xylographus: Xylographus bostrichoides 116 4.2. Morphological characters and systematic implications 116 4.2.1. Sensillifers of antennal club 116 4.2.2. Fovea on 1st abdominal ventrite 116

3 4.2.3. Cephalofoveae on forehead 117 4.3. Distribution and ecology of individual species 117 4.3.1. Distribution and ecology of Cis chinensis 118 4.3.2. Distribution and ecology of Cis submicans 122 4.3.3. Distribution and ecology of Cis comptus 122 4.3.4. Distribution and ecology of Cis striatulus 123 4.3.5. Distribution and ecology of Cis tomentosus 123 4.3.6. Distribution and ecology of Cis reitteri 124 4.3.7. Distribution and ecology of Cis castaneus 124 4.3.8. Distribution and ecology of Cis lugowoji 124 4.3.9. Distribution and ecology of Cis fissicollis 125 4.3.10. Distribution and ecology of Cis festivus 125 4.3.11. Distribution and ecology of Ennearthron comutum 125 4.3.12. Distribution and ecology of Orthocis reflexicollis 126 4.3.13. Distribution and ecology of Strigocis bicomis 126 4.3.14. Distribution and ecology of Sulcacis fronticomis 127 4.3.15. Distribution and ecology of Sulcacis nitidus 127 4.3.16. Distribution and ecology of Ropalodontus baudueri 128 4.3.17. Distribution and ecology of Ropalodontusperrini 128 4.3.18. Distribution and ecology of Octotemnus rugosopunctatus 129 4.3.19. Distribution and ecology of Xylographus bostrichoides 129 4.4. Overview of distribution ranges 130 4.5. Overview of elevation ranges 131 4.6. Overview of fungus host ranges 132 4.7. Ciidae fauna of the wider region 136 4.8. Faunal links and dispersal of Ciidae of Caspian Forest 137

Acknowledgements 250

References 251

Appendix: Collecting data, host fungi, co-occurrence 257

1. Cis chinensis 257 1.1. Samples of Cis chinensis collected 257 1.2. Host fungi of Cis chinensis 258 1.3. Ciidae species found associated with Cis chinensis 258 2. Cis submicans 260 2.1. Samples of Cis submicans collected 260 2.2. Host fungi of Cis submicans 262 2.3. Ciidae species found associated with Cis submicans 262 3. Cis comptus 267 3.1. Samples of Cis comptus collected 267 3.2. Host fungi of Cis comptus 268 3.3. Ciidae species found associated with Cis comptus 268 4. Cis striatulus 270 4.1. Samples of Cis striatulus collected 270 4.2. Host fimgi of Cis striatulus 271 4.3. Ciidae species found associated with Cis striatulus 272

4 5. Cis tomentosus 273 5.1. Samples of Cis tomentosus collected 273 5.2. Host fungi of Cis tomentosus 274 5.3. Ciidae species found associated with Cis tomentosus 275 6. Cis reitteri 277 6.1. Samples of Cis reitteri collected 277 6.2. Host fungi of Cis reitteri 277 6.3. Ciidae species found associated with Cis reitteri 277 7. Cis castaneus 278 7.1. Samples of Ciscastaneus collected 278 7.2. Host fungi of Cis castaneus 280 7.3. Ciidae species found associated with Cis castaneus 280 8. Cis lugowoji 282 8.1. Samples of Cis lugowoji collected 283 8.2. Host fungi of Cis lugowoji 283 8.3. Ciidae species found associated with Cis lugowoji 283 9. Cisfissicollis 284 9.1. Samples of Cis fissicollis collected 284 9.2. Host fungi of Cis fissicollis 285 9.3. Ciidae species found associated with Cis fissicollis 285 10. Cis fiestivus 287 10.1. Samples of Cis festivus collected 287 10.2. Host fungi of Cis festivus 287 10.3. Ciidae species found associated with Cis festivus 287 11. Ennearthron comutum 288 11.1. Samples of Ennearthron comutum collected 288 11.2. Host fungi of Ennearthron comutum 288 11.3. Ciidae species found associated with Ennearthron comutum 288 12. Orthocis reflexicollis 289 12.1. Samples of Orthocis reflexicollis collected 289 12.2. Host fungi of Orthocis reflexicollis 290 12.3. Ciidae species found associated with Orthocis reflexicollis 290 13. Strigocis bicornis 290 13.1. Samples of Strigocis bicornis collected 290 13.2. Host fungi of Strigocis bicornis 291 13.3. Ciidae species found associated with Strigocis bicornis 292 14. Sulcacis fronticornis 293 14.1. Samples of Sulcacis fronticornis collected 293 14.2. Host fungi of Sulcacis fronticornis 294 14.3. Ciidae species found associated with Sulcacisfronticornis 294 15. Sulcacis nitidus 295 15.1. Samples of Sulcacis nitidus collected 295 15.2. Host fungi of Sulcacis nitidus 297 15.3. Ciidae species found associated with Sulcacis nitidus 297 16. Ropalodontus baudueri 298 16.1. Samples of Ropalodontus baudueri collected 298 16.2. Host fungi of Ropalodontus baudueri 300 16.3. Ciidae species found associated with Ropalodontus baudueri 300 17. Ropalodontus perrini 302 17.1. Samples of Ropalodontus perrini collected 302

5 17.2. Host fungi of Ropalodontus perrini 303 17.3. Ciidae species found associated with Ropalodontusperrini 304 18. Octotemnus rugosopunctatus 305 18.1. Samples of Octotemnus rugosopunctatus collected 305 18.2. Host fungi of Octotemnus rugosopunctatus 307 18.3. Ciidae species found associated with Octotemnus rugosopunctatus 307 19. Xylographus bostrichoides 311 19.1. Samples of Xylographus bostrichoides collected 311 19.2. Host fungi of Xylographus bostrichoides 311 19.3. Ciidae species found associated with Xylographus bostrichoides 312