a timescale for the evolution of lice vincent s. smith with tom ford, kevin johnson, paul johnson,...
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A timescale for the evolution of lice
Vincent S. Smith
with Tom Ford, Kevin Johnson, Paul Johnson, Kazunori Yoshizawa, and Jessica E. Light
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Louse phylogeny (eight years on…)
• 60+ phylogenies! (circa 25 in 2003)
• Now cover 90% of louse genera
• Mix of molecules & morphology
• Beginning to reconcile differences
• Very unusual mit. & nuclear genome
• New questions emerging via next gen.
• One full genome (more coming)
AmblyceraAmblycera
IschnoceraIschnocera
RhynchophthirinaRhynchophthirina
AnopluraAnoplura
[ “[ “Psocoptera”Psocoptera”]]
68 spp. (15%)18 Genera1427 seq.+29 seq.*
1 spp. (33%)1 Genus12 seq.+3 seq.*
509 spp. (17%)95 Genera2264 seq.+285 seq.*
134 spp. (10%)37 Genera326 seq.+29 seq.*
286 spp.113 Genera1218 seq.+478 seq.*
GENBANK(June 2011)
*Change since Feb. 2010http://phylota.net/
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Why deep branch louse phylogeny is important?
• Permanent obligate parasites
• Has ecological replicates
• Opportunity to identify ancestral host
• Tied to origin of feathers and hair
• Model for cospeciation studies at different scales
• Need to date the origin of lice
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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.
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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
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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 !
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Molecular dating
i. Build a tree
Cha
ract
er c
hang
e
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Molecular dating
i. Build a tree
= ti
me
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Molecular dating
i. Build a tree
ii. Test for molecular clockNow
Age of lice
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Molecular dating
i. Build a tree
ii. Test for molecular clock
Rates of evolution vary
Now
Age of lice
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Molecular dating
i. Build a tree
ii. Test for molecular clock
iii. Apply calibrations
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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
- BEAST (Drummond et al, 2006)
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0.1
Louse phylogeny
Amblycera
Ischnocera
Anoplura
Rhynchophthirina
• 3 Genes (COI, EF1, 18s)
• No molecular clock
• Bayesian Analysis (MrBayes)- 4 chains, 10 million generations
• Recovers classical
louse phlogeny
• Eight calibration points
(Grimaldi & Engel 2006)
100 Ma min.Fossil Liposcelid booklouse
19.8 Ma min. Base of the Columbidae (fossil) (Benton 1993)
Oldest fossil Diomedeidae-37 - 53.25 Ma
Oldest fossil Procellariform(Benton 1993)
49 Ma min.Oldest Pelicaniform fossil(Mayr 2002)
20 - 25 Ma Old World Monkeys & apes(Reed et al 2004)
5-7 Ma Chimpanzees & humans(Reed et al 2004)
4.75 - 30 Ma Last extant gopher fossil -
(Russel 1968)gophers split from relatives
44 Ma min. (Wrappler et al 2004)Fossil Megamenopon louse
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0.1
Louse phylogeny
(Wappler et al 2004)
44 MyrFossil louse, Megamenopon
Fossil Louse
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0.1
Louse phylogeny
(Benton 1993)
19.8 MyrBase of the Columbidae (fossil)
Dove-Louse Cospeciation
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0.1
Louse phylogeny
(Benton ed. 1993)
37 - 53 MaRange for Diomedeidae based on fossils
Albatross-Louse Cospeciation
Hosts:Albatrosses
Parasites:Feather Lice
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0.1
Louse phylogeny
(Reed et al 2004)
5-7 MaFossil & Molecular data
Chimpanzee-human louse cospeciation
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0.1
Louse phylogeny
(Reed et al 2004)
20-25 MaFossil & Molecular data
Old World Monkeys - Apes cospeciation
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Louse phylogeny
Amblycera79 Ma(57-102 Ma)
Ischnocera98 Ma(74-125 Ma)
Anoplura74 Ma(53-98 Ma)
Rhynchophthirina
0100 Myr
KT105161
130 Myr
0.1
(Grimaldi & Engel 2006)
100 Ma min.Fossil Liposcelid booklouse
19.8 Ma min. Base of the Columbidae (fossil) (Benton 1993)
Oldest fossil Diomedeidae-37 - 53.25 Ma
Oldest fossil Procellariform(Benton 1993)
49 Ma min.Oldest Pelicaniform fossil(Mayr 2002)
20 - 25 Ma Old World Monkeys & apes(Reed et al 2004)
5-7 Ma Chimpanzees & humans(Reed et al 2004)
4.75 - 30 Ma Last extant gopher fossil -
(Russel 1968)gophers split from relatives
44 Ma min. (Wrappler et al 2004)Fossil Megamenopon louse
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• Lice and hosts must have
a long shared history
Louse phylogeny
• Three main clades of lice
parasitizing mammals
• Lice are approx. 130 Ma- 161 - 105 Ma, 95% conf. intervals
• Major louse radiation
before the KT boundary
Amblycera79 Ma(57-102 Ma)
Ischnocera98 Ma(74-125 Ma)
Anoplura74 Ma(53-98 Ma)
Rhynchophthirina
0100 Myr
KT105161
130 Myr
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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
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Summary
• Just one fossil louse, but..
• Many other calibration points
• Lice are approx. 130 Ma 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?
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PressSmith et al 2011. Multiple lineages of lice pass through the K–Pg boundary. Biology Letters doi:10.1098/rsbl.2011.0105
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Questions?
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