Leptree fossil project: when Lepidopterists meet Paleontologists

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Leptree fossil project when Lepidopterists
meet Paleontologists

• Jae-Cheon SOHN1, Conrad LABANDEIRA2, and the
  LepTree Consorsium3

• Department of Entomology, University of Maryland,
  
• College Park, MD 20742
• 2) Department of Paleontology, Smithsonian
  Institution
• National Museum of Natural History,
  Washington, DC 20560
• 3) Charles Mitter, Don Davis, John Brown, Susan
  Weller,
• Cynthia Parr, Akito Kawahara, Andreas Zwick,
  Jerry Regier,
• Michael Cummings, Soowon Cho and external
  contributors.

Lepidopterist
Paleontologist
Lepidopterists
Paleontologists
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Fossils in Phylogeny inference- Why are they
important

• Morphological phylogenetics
• 1) bridging problematic morphological gaps
  subdividing long branches
• (Gauthier et al., 1988)
• 2) offering morphological snapshot close to
  the splitting of lineages (Novacek, 1992)
• 3) assessing the direction of character
  evolution (Hennig, 1966)

Molecular Phylogenetics (critical for LepTree
project) Calibration (relaxed) molecular clock,
allowing all major lineages to be
dated using molecular tree
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Difficulties of using fossils for evolutionary
study

• Limited taxon availability
• - the record is sparse
• Incomplete preservation or visibility of
  characters
• Taxomomic misinterpretation/ uncertainty of
  Phylogenetic placement
• - synapomorphies not known, or not
• observable
• Geological uncertainty

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Types of Lepidopteran Fossils

• Preservation type
• 1) Permineralization
• - amber/copal
• - silification
• 2) Compression/impression
• 3) Sieved residue
• Forms of fossils
• 1) Body fossil (egg/larva/pupa/adult)
• 2) Trace fossil (feeding damage/coprolite/gut
  contents)

Prodryas persephone (Florrisant, Late Eocene)
From Meyer (2003)
A moth inclusion in Baltic amber (Eocene)
www.harunyahya.com
A fossil bat Palaeochiropteryx tupaiodon (Eocene)
from Richter Storch (1980)
A leaf-mine fossil (Eocene) From Grimaldi
Engel (2005)
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Previous reviews of Fossil Lepidoptera

• Taxonomic reviews
• Scudder (1875) Fossil butterflies
• Kusnezov (1941), Skalski (1976), Poinar (1992)
  Lepidoptera inclusions in ambers
• Bachofen-Echt (1949) Lepidoptera in Baltic amber
• Leestmans (1983) Lepidoptera fossils found in
  France
• Kozlov (1988) all known Lepidoptera fossils
• Danilevsky Martynova (1991) Fossil records of
  the Lepidopterous families
• Carpenter (1992) Generic review of the known
  fossil Lepidoptera
• Petrulevicius Martins-Neto (2000) South
  American Cenozoic Lepidoptera
• Rasnitsyn Ross (2000) Lepidoptera in the
  Burmese amber collection at BMNH
• Meyer (2003) Lepidoptera from the Florissant
  fossil beds
• Grimaldi Engel (2005) Review of lepidopteran
  fossil record

Numerical estimation Kristensen Skalski
(1998) about 600-700 specimens known
(ca. 500
preserved in resin)
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Problems in deciphering the fossil record of
Lepidoptera

• 1) Limited taxon availability
• Why are compression fossils of Lepidoptera rare?
• fragility
• buoyancy

ltTaphonomy of Lepidopteragt Fast decay
in water Ant
consumption -gt few remains (Duncan, 1997)
(Smith, 1998)
4 weeks
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Problems in deciphering the fossil record of
Lepidoptera

• Biased sampling of Lepidoptera in Amber
• - taxonomic composition of Microlepidoptera
  trapped in mordern pine resin differs from
  actual composition of fauna (Skalski, 1976)

Actual microlepidoptera fauna in the forest
Microlepidoptera fauna trapped in the pine resin
After Skalski (1976)
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Problems in deciphering the fossil record of
Lepidoptera

• 2) taxonomic misinterpretation
• - misidentification by external analogy
• ex) Palaeontinidae (Homoptera)

www.fossilmuseum.net
Palaeontina oolitica (after Butler, 1873)
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Problems in deciphering the fossil record of
Lepidoptera

• 2) taxonomic misinterpretation
• - wrongly characterized by the distorted images
  of fossils

Borkhausenites bachofeni Rebel (Oecophoridae) Balt
ic amber (Eocene) from Skalski (1977)
Original drawing by Rebel (1934)
A photograph of holotype
Wing venation redrawn by Skalski (1977)
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Problems in deciphering the fossil record of
Lepidoptera

• 2) taxonomic misinterpretation
• - confusion between plant material and
  Lepidoptera

misidentified as a lepidopteran ex) Daudet
(1876) Satyrites incertus coniferous
male cone (Nel Nel, 1985)
misidentified as plants Reid (1915) Aralia
fruits Chandler (1961) Zelkova fruits
caterpillar coprolite (Lancucka-Srodoniowa,
1964)
www.mntca.org
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Problems in deciphering the fossil record of
Lepidoptera

• 3) geological uncertainty
• - often absolute age estimation unavailable
• - a fossil bed possible combination of
• variously aged layers
• - reworked fossil-bearing bed (ex. Burmese amber)

www.amberworkshop.com
Zherikhin Ross (2000) the fossil beds
reworked
(Late Cretaceous)
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Problems in deciphering the fossil record of
Lepidoptera

• 4) incomplete preservation
• - Fossil taxa rarely displaying diagnostic
  characters (Kristensen Skalski, 1998)

Protolepis cuprealata (Late Jurassic) After
Kozlov (1989)
Lepidopteran groundplan autapomorphies Modifed
from Kristensen Skalski (1998)
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Problems in deciphering the fossil record of
Lepidoptera

• 4) incomplete preservation
• - Fossils causing lots of missing data and
  negatively affecting phylogenetic study
• - decreasing the resolution of tree
• (Gauthier et al., 1988)
• - lowering support values over whole
  tree
• (Wilkinson Benton, 1996)
• - bad influence on otherwise
  well-supported clades
• (Wilkinson, 1995)

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Problems in deciphering the fossil record of
Lepidoptera

• 4) incomplete preservation
• - contamination
• new techniques
• ex) Micro-tomography (ESRF synchrotron
  Lak et al., 2008)

After http//www.esrf.eu/news/general/amber
X-ray image
Micro-tomographic image
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So, how to overcome these problems?
Incomplete Taxon Sampling
Taxonomic Misinterpretation
Lepidopterists
Paleontologists
New Technologies
Incomplete Preservation
Geological Uncertainty
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Molecular Phylogeny Resolve backbone of
lepidopteran relationships
Training Workshops Develop and share molecular
markers protocols
Lepidoptera Fossil Database Compilation and
review of fossils for dating and hypothesis
testing

• LepInformatics project
• Provide the community with an interactive website
  and tools to present, discuss and expand our
  collective knowledge

Morphology Initiative Develop a comprehensive
illustrated glossary of descriptive terms based
on homologies
AToL (Alcohol Tubes of Lepidoptera) Build a
frozen tissue collection for all to use
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Molecular backbone phylogeny and dating 26
nuclear genes 127 families
Twig Leaders
Morphology Projects

• Compile current knowledge relevant to phylogeny
  character descriptions images, data matrices,
  trees, literature, primers, availability of
  material
• Compile morphological homologies glossary
• with reconciled terminologies, providing
  characters for future higher phylogeny studies

Lepidoptera fossil database
AToLep Tissue Collection
Training Workshops
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How does Fossil Project progress?
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Launched Website
Family Name
Species Name
http//www.leptree.net/fossil
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Synopsis of Leptree fossil data base entries
Superfamily composition (439 specimens)
Composition by identification category
Composition by preservation type (526 specimens)
Composition by geological age
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Species information sheet

• 1) Body fossil
• - Taxonomic information
• rank
• - Specimen information
• preservation depository
• - Geologic information
• age locality
• - Diagnostic character by authors
• - References
• - Image Description
• 2) Trace fossil (leaf mines)
• - host plant information

To be in future
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Interaction in Fossil Project

• comments section in each sheet
• - To find new evidence of the known fossil
• ex) Axel Hausmanns reevaluation of
  Hydriomena(?) protrita
• Cockerell, 1922 (Florissant, Late
  Eocene)
• - To find erroneously placed taxa and new
  position for them
• - To place the fossil taxa remaining unsettled

Hydriomena (?) protrita (Florissant, Late
Eocene) After Cockerell (1922)
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Acknowledgements
LepTree Twig leaders LepTree Collaborators
Axel Hausmann Joaquin Baixeras LepTree
Informatics Staff John Park Giho
Kim Technician Erika Koenig Helper for
Translation Kim Mitter Ada Szczepaniec
Smithsonian Paleontology Finnegan Marsh
Wenying Wu