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Did dinosaurs have lice? Vince Smith, Tom Ford, Kevin Johnson*, Paul Johnson & Rod Page University of Glasgow, Glasgow, UK *Illinois Natural History Survey, USA

2004.V Smith.Dino Lice

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Did dinosaurs have lice?

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Page 1: 2004.V Smith.Dino Lice

Did dinosaurs have lice?

Vince Smith, Tom Ford, Kevin Johnson*,Paul Johnson & Rod Page

University of Glasgow, Glasgow, UK*Illinois Natural History Survey, USA

Page 2: 2004.V Smith.Dino Lice

• Collected Eckfeld maar nearEifel, Germany

• Middle Eocene (Middle Lutetian,44.3±0.4 Ma)

• First fossil louse

• Completes ordinal representationof fossil insects

• Excellent preservation

• Phylogenetic affinities withmodern feather lice

Fossil Lice

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 3: 2004.V Smith.Dino Lice

Fossil Lice

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 4: 2004.V Smith.Dino Lice

Fossil Lice

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 5: 2004.V Smith.Dino Lice

Fossil Lice

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 6: 2004.V Smith.Dino Lice

Fossil Lice

Psocoptera

Menacanthus /Menopon - complex

Ricinidae

Laemobothriidae

Boopiidae

Austromenopon- complex

Dennyus- complex

Colpocephalum- complex

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 7: 2004.V Smith.Dino Lice

Fossil Lice

• Parasite of Anseriformes orCharadriiformes

• Crown group position• Suggests lice are very old• Points to a long coevolutionary

history with birds• Useful molecular calibration point

Psocoptera

Menacanthus /Menopon - complex

Ricinidae

Laemobothriidae

Boopiidae

Austromenopon- complex

Dennyus- complex

Colpocephalum- complex

Wappler, Smith & Dalgleish,Proc.R.Soc., 2004.

Page 8: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

Page 9: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

ii. Establish calibration points- Crown group fossils

i. Test for molecular clock

Page 10: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

ii. Establish calibration points- Crown group fossils

- R8s (Sanderson, 2003)iii. Apply rate smoothing

- Penalized likelihood- Cross validation100

75

50

25

0

i. Test for molecular clock

Page 11: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

ii. Establish calibration points- Crown group fossils

- R8s (Sanderson, 2003)iii. Apply rate smoothing

- Penalized likelihood- Cross validation100

75

50

25

0

100

75

50

25

0

iv. Multiple approaches!- Bottom up

i. Test for molecular clock

Page 12: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

ii. Establish calibration points- Crown group fossils

- R8s (Sanderson, 2003)iii. Apply rate smoothing

- Penalized likelihood- Cross validation100

75

50

25

0

100

75

50

25

0

iv. Multiple approaches!- Bottom up- Top down - cospeciation dates

i. Test for molecular clock

Page 13: 2004.V Smith.Dino Lice

Approaches To Dating

i. Test for molecular clock

ii. Establish calibration points- Crown group fossils

- R8s (Sanderson, 2003)iii. Apply rate smoothing

- Penalized likelihood- Cross validation100

75

50

25

0

100

75

50

25

0

iv. Multiple approaches!- Bottom up- Top down - cospeciation dates

30

20

10

0

v. Compare ages30

20

10

0

100

75

50

25

0

Bot

tom

Up

Top Dow

n

35 ma

30 ma

i. Test for molecular clock

Page 14: 2004.V Smith.Dino Lice

Phthiraptera

Phthiraptera

18s rRNA COI (1st & 2nd pos.)

ML(GTR+I+G)

ML(TVM+I+G)

0.1 0.1

CrustaceaDictyoptera

Hemiptera

Psocoptera

Lepidoptera

Diptera

CrustaceaOdonata

DictyopteraPlecoptera

Hemiptera

Psocoptera

ColeopteraRaphidiodea

MegalopteraNeuroptera

Hymenoptera

TrichopteraLepidoptera

SiphonapteraMecoptera

Diptera / Strepsiptera

Thysanoptera

Page 15: 2004.V Smith.Dino Lice

EF1-α Wingless

ML(GTR+I+G)

ML(GTR+I+G)

0.1 0.1

Phthiraptera

Phthiraptera

Trichoptera

Lepidoptera

SiphonapteraMecoptera

Coleoptera

Hymenoptera

Diptera

Hemiptera

Thysanoptera

Odonata

Hymenoptera

Diptera

Lepidoptera

Coleoptera

Thysanoptera

Hemiptera

Crustacea

Page 16: 2004.V Smith.Dino Lice

0.1 0.1 0.1 0.1

18s rRNA COI EF1-α Wingless

Insect Phylogeny (unsmoothed)

Ages Fixed Age Constraints

Page 17: 2004.V Smith.Dino Lice

~185 ma~260 ma ~250 ma~295 ma(280-300 ma) (235-300 ma) (200-250 ma) (185-240 ma)

18s rRNA COI EF1-α Wingless

0100200300

Hym

enopteraD

ipteraLepidoptera

Coleoptera

ThysanopteraLIC

EH

emiptera

Hem

ipteraD

ipteraLepidoptera

0100200300400500

LICE

0100200300400

Lepidop.C

addisFleas

Mecop.

FliesH

ymenop.

Coleop.

LICE

Bugs

ThripsO

donata

0100200300400

Strepsip.Flies

Fleas/Mecop.

Tric./Lep.H

ymenop.

Neurop.

Beetles

LICE

BugsB

ark Lice

Insect Phylogeny (rate smoothed)

Page 18: 2004.V Smith.Dino Lice

Cospeciation Events

Constraints

Min. Age 44 ma

Louse Fossil

Combined ML

0.1

(18s, COI, EF1-α)

Am

blyceraIschnocera

Anoplura

Rhyn.

Louse Phylogeny (unsmoothed)

Page 19: 2004.V Smith.Dino Lice

Cospeciation Events

Constraints

Combined ML

0.1

(18s, COI, EF1-α)

Am

blyceraIschnocera

Anoplura

Rhyn.

Louse Phylogeny (unsmoothed)

Dove-LouseCospeciation

Age Fixed 19.8 ma

Page 20: 2004.V Smith.Dino Lice

Cospeciation Events

Constraints

Combined ML

0.1

(18s, COI, EF1-α)

Rhea/Ostrich-Louse Cospeciation

Age Fixed 60 ma

Am

blyceraIschnocera

Anoplura

Rhyn.

Louse Phylogeny (unsmoothed)

Page 21: 2004.V Smith.Dino Lice

Cospeciation Events

Constraints

Combined ML

0.1

(18s, COI, EF1-α)

Am

blyceraIschnocera

Anoplura

Rhyn.

Louse Phylogeny (unsmoothed)

Primate-LouseCospeciation

Age Fixed 4.5 ma

Page 22: 2004.V Smith.Dino Lice

Ricinus spLaemobothrion atrumAustromenopon elliottiTrinoton querquedulaeHolomenopon spDennyus hirundinisMachaerilaemus spColimenopon urocoliusDocophoroides brevisFulicoffula longipilaBovicola spFelicola subrostratusNeotrichodectes spGeomydoecus spPectinopygus sulaePectinopygus brevicornisPectinopygus punctatusPectinopygus variusAnatoecus icterodesTrichophilopterus babakotophilusColumbicola columbaeColumbicola passerinaeStruthioliperus nanduStruthiolipeurus struthionisCraspedorrhynchus spDegeeriella fulvaAustrophilopterus subsimilusColilipeurus coliusCampanulotes comparGoniodes spNaubates spPerineus nigrolimbatusHaematomyzus elephantsPediculus humanus var capitisPediculus schaeffiEchinophthirius horridus bPolyplax spinulosaHaematopinus tuberculatusHaematopinus suisHaematopinus pacochoeriHoplopleura trispinosaHoplopleura spPterophthirus imitansLinognathoides marmotaeLinognathoides marmotae bNeohaematopinus sciuriNeohaematopinus sciuri b

0100200

Louse Phylogeny (rate smoothed)

Combined ML(18s, COI, EF1-α)

Am

blyceraIschnocera

Anoplura

Rhyn.

~190 ma(150-225 ma)

KT

Result very sensitiveto smoothing

parameter used

Page 23: 2004.V Smith.Dino Lice

Host Implications

Paleocene

Cretaceous

LateCretaceous

EarlyCretaceousLateJurassic

157ma ±

146ma ±

97ma ±

65ma ±

Modern Birds

‘Ratites’and

tinamousNeognathae

Neornithes

Archaeopteryx

Paleognathae

Tertiary ‘big bang’ model formodern bird evolutionsensu Feduccia, 1995

Figure adaptedfrom Feduccia,

1995

010

020

0

‘Big bang’ followed bya ‘big switch’?

Modern birds much older- Gondwanic origin?

Revised classic modelsensu Cracraft, 2001

KT

Modern Birds

Page 24: 2004.V Smith.Dino Lice

Host Implications

Figure adaptedfrom ???, Science,

????

2-300ma ?

Page 25: 2004.V Smith.Dino Lice

Summary

• Lice are old (150-300ma)• Major radiation of lice before the KT (65ma)• Modern birds & lice have a very long

coevolutionary history

• Origin of lice associated with origin of feathers?

• Recent threopod dinosaurs probably had lice?

• Molecular clocks run fast at times of rapidradiation?

Caveats

Page 26: 2004.V Smith.Dino Lice

Acknowledgements

• Wellcome Trust