Traumatic Insemination in the plant bug genus Coridromius

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

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

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

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

Coercive mating in waterstriders Westlake Rowe & Currie (2000)

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

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

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 e. g. Traumatic Insemination & Paragenitalia

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

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

Heteroptera (True Bugs)

Heteroptera (True Bugs) Cimicimorpha

R M r ic Schuh & Stys (1991) d p Jo i pe e da a Th um M iri (C ae T id g in e M e c do a id N ab os Pr t ( d ili i ch o si La ae ae il d lo P ph o ki L to yc ae ae rid o c c A n o th ae id or e ct y ol P d ni ae hy p o os tid id id vi u ed t as ic ae i or ic si ae ae om r d im P yn h ac e da rid o oc m em C e da a iu ae i om s) e) a tin

) e na i at R M r ic d p Jo i pe e da a Th um M iri (C ae id g in T os c do M e tid e a id ab N ( d ili i ch o si La ae ae il d ph o ki P lo 1 1 Schuh & Stys (1991) Genitalic TI L to yc ae ae rid o c c A n o th ae id or e ct y ol P d ni ae hy p o i or os r P id id vi u ed t as ic o oc ae ic si ae ae om r d im P yn h ac e da a rid C i om iu e e da m m te s)

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

P u ed R M r ic hy p o pp o J d ei m au h T o to as d ae M iri (C oc id ng Ti id t os ae ed M t os r P ae i e da ab N ( hi c o si La e ae ae a d id h rid ori n p o e c io oc ho yct k t o c nt Pol Pl Ly A ae il d 1 2 Schuh & Stys (1991) 1 Genitalic TI 2 Extragenitalic TI ae id vi co e a ic rid e a li id id yn h ac ae d si ae m ro rid m em C im ic e da a iu ae i om s) e) a tin

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

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

R M um a Th o ae M d iri (C id t os ae oc d T gi in ed M ae ( N i ch io as d i ab ae il d L e ae lid i ph io k lo P ri o ct y L a rid e da o oc h nt A 1 3 2 Schuh & Stys (1991) 1 Genitalic TI 2 Extragenitalic TI 3 Aedeagus & Paramere coupled en t c ly Po e a id ae ei pp o J t as ri t os r P ae id ph ro ic c di om r d im ic P ae d i ys e da ii uv d e ae rid o oc m em C e da a iu ae i om n hy ac s) e) a tin

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

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

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

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

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

The males

e fe m al es ?

) s u e) a tin P ii R uv d e M ae d i ys e da ph ro ic pp o J ei c di um a Th t as M d iri 2, 3 ae i or (C id t os ae oc d T gi in ed M t os r P ae ae ( N i ch io as d i ab ae il d L lid i ph io k lo P e ae ri o ct y L a rid e da o oc h nt A 1 3 2 Schuh & Stys (1991) 1 Genitalic TI 1. Genitalic TI 2 Extragenitalic TI 3 Aedeagus & Paramere coupled en t c ly Po e a id ae i om n hy ac ae rid o oc im ic e da a m em C ae om r d id i

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

Part 2 • Taxonomic descriptions • Phylogeny

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

Asia + Africa Pacific 86

Enough biogeography – back to the sex!

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

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?

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