1. The Diversification of Lice: Age Vincent S. Smith with Tom Ford, Kevin Johnson, Paul Johnson, Kazunori Yoshizawa, and Roderic D. M. Page

2. Psocoptera Menacanthus / Menopon -complex Ricinidae Laemobothriidae Boopiidae Austromenopon -complex Dennyus -complex Colpocephalum -complex Collected Eckfeld maar near Eifel, Germany Middle Eocene (Middle Lutetian, 44.3±0.4 Ma) First fossil louse Completes ordinal representation of fossil insects Excellent preservation Phylogenetic affinities with modern feather lice Parasite of Anseriformes or Charadriiformes Crown group position Suggests lice are very old Points to a long coevolutionary history with birds Useful molecular calibration point Wappler, Smith & Dalgleish First fossil louse Proc. R. Soc., 2004.

3. Not fossil lice! Amblyceropsis indica Kumar (2004) considered a mite by Dalgleish, Palma, Price, & Smith 2006 fused head and thorax (cephalothorax) Anopluropsis khatamaensis Kumar (2004) considered an orbatid mite by Dalgleish, Palma, Price, & Smith 2006

4. Not fossil lice! Saurodectes vrsanski Rasnitsyn & Zherikhin (1999) insect of uncertain ordinal affinities by Grimaldi and Engel 2005 & Dalgleish, Palma, Price, & Smith 2006 pterosaur louse !

5. Molecular dating i.Build a tree Character change

6. Molecular dating i.Build a tree = time

7. Molecular dating i.Build a tree ii.Test for molecular clock Now Age of lice

8. Molecular dating i.Build a tree ii.Test for molecular clock Rates of evolution vary Now Age of lice

9. Molecular dating i.Build a tree ii.Test for molecular clock iii.Apply calibrations

10. Molecular dating i.Build a tree ii.Test for molecular clock iii.Apply calibrations iv.Smooth rates across the tree -R8s with Penalized Likelihood (Sanderson, 2003) -Multidivtime (Thorne et al, 1998) Now Age of lice

11. Louse phylogeny Oldest fossil Diomedeidae- (Wappler et al 2004) 44 Myr Fossil louse, Megamenopon 4.75 - 30 Myr Last extant gopher fossil - (Russel 1968) 19.8 Myr Base of the Columbidae (fossil) (Benton 1993) 60 Myr Oldest fossil Rhea (Tambussi 1995) 37 - 53.25 Myr Oldest fossil Procellariform (Benton 1993) 49 Myr Pelicaniform fossil (Mayr 2002) 20 - 25 Myr Fossil & molecular data (Reed et al 2004) 5 Myr Fossil & molecular data (Reed et al 2004) gophers split from relatives Amblycera Ischnocera Anoplura Rhynchophthirina 3 Genes (COI, EF1, 18s) No molecular clock Bayesian Analysis - 4 chains, 5 million generations Recovers classical louse phlogeny Eight calibration points

12. Louse phylogeny (Wappler et al 2004) 44 Myr Fossil louse, Megamenopon Fossil Louse

13. Louse phylogeny (Benton 1993) 19.8 Myr Base of the Columbidae (fossil) Dove-Louse Cospeciation

14. Louse phylogeny (Tambussi 1995) 60.0 Myr Oldest Fossil Rhea Rhea/Ostrich-Louse Cospeciation

15. Louse phylogeny (Reed et al 2004) 5.0 Myr Fossil & Molecular data Chimpanzee-human louse cospeciation

16. Louse phylogeny (Reed et al 2004) 5.0 Myr Fossil & Molecular data Chimpanzee-human louse cospeciation

17. Louse phylogeny Oldest fossil Diomedeidae- (Wappler et al 2004) 44 Myr Fossil louse, Megamenopon 4.75 - 30 Myr Last extant gopher fossil - (Russel 1968) 19.8 Myr Base of the Columbidae (fossil) (Benton 1993) 60 Myr Oldest fossil Rhea (Tambussi 1995) 37 - 53.25 Myr Oldest fossil Procellariform (Benton 1993) 49 Myr Pelicaniform fossil (Mayr 2002) 20 - 25 Myr Fossil & molecular data (Reed et al 2004) 5 Myr Fossil & molecular data (Reed et al 2004) gophers split from relatives Amblycera 96 Myr (76.4-124.9 Myr) Ischnocera 88.9 Myr (76.3-102.7 Myr) Anoplura 73.4 Myr (61.4-87.5 Myr) Rhynchophthirina 0 100 Myr KT 96 141 116 Myr

18. Lice and hosts must have a long shared history Louse phylogeny Amblycera 96 Myr (76.4-124.9 Myr) Ischnocera 88.9 Myr (76.3-102.7 Myr) Anoplura 73.4 Myr (61.4-87.5 Myr) Rhynchophthirina 0 100 Myr KT 96 141 116 Myr Three main clades of lice parasitizing mammals Lice are approx. 116 Myr old - 141 - 96 Myr, 95% conf. intervals Major louse radiation before the KT boundary

19. Host implications Lice radiated on birds first Mammal lice are young These groups colonized mammals Placental Mammals Marsupials Birds What were the first louse hosts? Did lice radiate with feathers? feathers hair

20. Summary Just one fossil louse, but.. Many other calibration points Lice are approx. 120 Myr old Radiated before dinosaurs when extinct First radiated on the lineage leading to birds Either modern birds are older than we think, or… Perhaps the bird-like Theropod dinosaurs had lice? Radiated in response to the evolution of feathers?

21. Questions?