1. Traumatic Insemination in the plant bug genus Coridromius (Heteroptera: Miridae) Nik Tatarnic University of Sydney & Australian Museum

2. Q. What is Traumatic Insemination? A. Male punctures female with hypodermic genitalia, injects sperm into body cavity

3. Q. Why Practice Traumatic Insemination? A. Thought to be a means for males to circumvent female mating resistance

4. Q. Why Resist Mating? A. Males and Females have differing reproductive optima = Sexual Conflict

5. Coercive mating in waterstriders Westlake Rowe & Currie (2000)

6. Crudgington, H.S. and Siva-Jothy, M.T. (2000) Genital Damage in Bean Weevils

7. Sexual Conflict Coevolutionary Arms Race Males evolve structures and behaviours which increase their own fitness, but reduce female fitness Females evolve structures and behaviours to mitigate costs

8. e.g. Traumatic Insemination & Paragenitalia Sexual Conflict Coevolutionary Arms Race Males evolve structures and behaviours which increase their own fitness, but reduce female fitness Females evolve structures and behaviours to mitigate costs

9. Who are the Traumatic Inseminators? Non-insect invertebrates: rotifers, acanthocephalans, pinworms, gastropod snails, free-living flatworms Insects: Strepsiptera, Cimicomorpha (Heteroptera) e.g. Bedbugs

10. Who are the Traumatic Inseminators? Non-insect invertebrates: rotifers, acanthocephalans, pinworms, gastropod snails, free-living flatworms Insects: Strepsiptera, Cimicomorpha (Heteroptera) e.g. Bedbugs

11. Heteroptera (True Bugs)

12. Cimicimorpha Heteroptera (True Bugs)

13. Pachynomidae Thaumastocoridae Miridae (Coridromius) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae Schuh & Stys (1991)

14. Pachynomidae Thaumastocoridae Miridae (Coridromius) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae 1 1 Genitalic TI Schuh & Stys (1991)

15. Genitalic Traumatic Insemination: Prostemmatinae vaginal copulation genital tract wall pierced with phallus sperm travel through hemocoel to ovarioles C. Weisenböhler Deutschland / Baden-Württemberg, Vellberg III.2005

16. Pachynomidae Thaumastocoridae Miridae ( Coridromius ) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae 1 2 1 Genitalic TI 2 Extragenitalic TI Schuh & Stys (1991)

17. Extragenitalic Traumatic Insemination (I) Male punctures the female body wall with his genitalia –“Acus” in Plokiophilidae & Lyctocoridae –Aedeagus + left paramere in Anthocoridae*, Polyctenidae and Cimicidae *Not all Anthocoridae practice TI

18. Carpentiro & Dellapé (2005) Carayon (1972) Plokiophilidae Lyctocoridae

19. Pachynomidae Thaumastocoridae Miridae (Coridromius) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae 1 2 3 1 Genitalic TI 2 Extragenitalic TI 3 Aedeagus & Paramere coupled Schuh & Stys (1991)

20. Extragenitalic Traumatic Insemination (II) Male punctures the female body wall with his genitalia –“Acus” in Plokiophilidae & Lyctocoridae –Aedeagus + left paramere in Anthocoridae*, Polyctenidae and Cimicidae *Not all Anthocoridae

21. Anthocoridae, Polyctenidae & Cimicidae Stutt and Siva-Jothy (2001)

22. Female Response Females have evolved paragenitalia to mitigate costs Spermalege = External “Ectospermalege” & Internal “Mesospermalege”

23. Plokiophilidae Cimicidae Stutt and Siva-Jothy (2001) Carayon (1974)

24. A New Case of Traumatic Insemination Family Miridae Genus Coridromius +32 species (11 described) Pacific Islands, Australia, SE Asia, Africa Extragenitalic TI with paramere & aedeagus coupled *Tatarnic, N.J., Cassis G. & Hochuli D.F. 2006. Biology Letters 2: 58-61

25. 1. Genitalic TI Pachynomidae Thaumastocoridae Miridae (Coridromius) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae 1 2,3 2 3 1 Genitalic TI 2 Extragenitalic TI 3 Aedeagus & Paramere coupled Schuh & Stys (1991)

26. The males

27. And the females?

31. 1. Genitalic TI Pachynomidae Thaumastocoridae Miridae (Coridromius) Tingidae Plokiophilidae Lyctocoridae Anthocoridae Polyctenidae Cimicidae Lasiochilidae Nabidae (Prostemmatinae) Medocostidae Microphysidae Joppeidicae Reduviidae 1 2,3 2 3 1 Genitalic TI 2 Extragenitalic TI 3 Aedeagus & Paramere coupled Schuh & Stys (1991)

32. Conclusions I Traumatic Insemination has arisen multiple times Coupling of paramere and aedeagus has arisen twice

33. Part 2 Taxonomic descriptions Phylogeny

35. 86 75 55 54 98 63 92 59 85 68 96 50 86 56 72

36. Asia + AfricaPacific 86

40. Enough biogeography – back to the sex!

42. Conclusions II Female paragenitalic evolution does not follow a simple transition series Instead, multiple independent cases of elaborate paragenital development = Strong sexual selection

43. Summary Traumatic Insemination has evolved at least 3 times in the Cimicomorpha TI drives elaborate female morphology in Coridromius Question: What predisposes Cimicomorpha to such deviant sexual practices?

44. Acknowledgements Gerry Cassis, AM Dieter Hochuli, U Sydney Michael Wall, AM/AMNH Locke Rowe, U Toronto Mike Siva-Jothy, U Sheffield NSF: Planetary Biodiversity Inventory