The Diversification of Lice: Age

Vincent S. Smithwith Tom Ford, Kevin Johnson, Paul Johnson, Kazunori Yoshizawa, and Roderic D. M. Page

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.

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

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 !

Molecular dating

i. Build a tree

Cha

ract

er c

hang

e

Molecular dating

i. Build a tree

= ti

me

Molecular dating

i. Build a tree

ii. Test for molecular clockNow

Age of lice

Molecular dating

i. Build a tree

ii. Test for molecular clock

Rates of evolution vary

Now

Age of lice

Molecular dating

i. Build a tree

ii. Test for molecular clock

iii. Apply calibrations

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

0.1

Louse phylogeny

Oldest fossil Diomedeidae-

(Wappler et al 2004)

44 MyrFossil 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 MyrPelicaniform 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

0.1

Louse phylogeny

(Wappler et al 2004)

44 MyrFossil louse, Megamenopon

Fossil Louse

0.1

Louse phylogeny

(Benton 1993)

19.8 MyrBase of the Columbidae (fossil)

Dove-Louse Cospeciation

0.1

Louse phylogeny

(Tambussi 1995)

60.0 MyrOldest Fossil Rhea

Rhea/Ostrich-Louse Cospeciation

0.1

Louse phylogeny

(Reed et al 2004)

5.0 MyrFossil & Molecular data

Chimpanzee-human louse cospeciation

0.1

Louse phylogeny

(Reed et al 2004)

5.0 MyrFossil & Molecular data

Chimpanzee-human louse cospeciation

0.1

Louse phylogeny

Oldest fossil Diomedeidae-

(Wappler et al 2004)

44 MyrFossil 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 MyrPelicaniform 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

Amblycera96 Myr(76.4-124.9 Myr)

Ischnocera88.9 Myr(76.3-102.7 Myr)

Anoplura73.4 Myr(61.4-87.5 Myr)

Rhynchophthirina

0100 Myr

KT96141

116 Myr

• Lice and hosts must have

a long shared history

Louse phylogeny

Amblycera96 Myr(76.4-124.9 Myr)

Ischnocera88.9 Myr(76.3-102.7 Myr)

Anoplura73.4 Myr(61.4-87.5 Myr)

Rhynchophthirina

0100 Myr

KT96141

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

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

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?

Questions?