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					        “Leptree fossil project”:
when Lepidopterists meet Paleontologists
Jae-Cheon SOHN1, Conrad LABANDEIRA2, and the LepTree Consorsium3
1) Department of Entomology, University of Maryland,
   College Park, MD 20742
2) Department of Paleontology, Smithsonian Institution          Paleontologist   Lepidopterist
   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.

 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
Difficulties of using fossils for
      evolutionary study

               • Limited taxon availability
                   - the record is sparse

               • Incomplete preservation or visibility of

               • Taxomomic misinterpretation/ uncertainty
                 of Phylogenetic placement
                    - synapomorphies not known, or not

               • Geological uncertainty
Types of Lepidopteran Fossils

* Preservation type
  1) Permineralization
    - amber/copal
    - silification              A moth inclusion in Baltic amber Prodryas persephone
                                (Eocene) www.harunyahya.com      (Florrisant, Late Eocene) From Meyer (2003)

  2) Compression/impression
  3) Sieved residue

                                    A fossil bat Palaeochiropteryx tupaiodon
                                    (Eocene) from Richter & Storch (1980)

* Forms of fossils
  1) Body fossil (egg/larva/pupa/adult)                                        A leaf-mine fossil (Eocene)
                                                                               From Grimaldi & Engel (2005)

  2) Trace fossil (feeding damage/coprolite/gut contents)
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)
 Problems in deciphering the fossil
       record of Lepidoptera

1) Limited taxon availability
- Why are compression fossils of Lepidoptera rare?

 <Taphonomy of Lepidoptera>

   4 weeks

 Fast decay in water          Ant consumption -> few remains
 (Duncan, 1997)               (Smith, 1998)
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

                      fauna trapped in the pine resin
                                                        After Skalski (1976)
Problems in deciphering the fossil
      record of Lepidoptera
2) taxonomic misinterpretation
- misidentification by external analogy
  ex) Palaeontinidae (Homoptera)


    Palaeontina oolitica (after Butler, 1873)
Problems in deciphering the fossil
      record of Lepidoptera
2) taxonomic misinterpretation
 - wrongly characterized by the distorted images of fossils
                                      Borkhausenites bachofeni Rebel (Oecophoridae)
                                      Baltic amber (Eocene) from Skalski (1977)

                              Original drawing by Rebel (1934)

                              A photograph of holotype

                              Wing venation redrawn by Skalski (1977)
 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)

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                                               www.amberworkshop.com

- reworked fossil-bearing bed (ex. Burmese amber)

                    Authors                          Age

                Noetling (1892)                    Miocene

                Cockerell (1917)               Late Cretaceous

         Stuart (1923), Cockerell (1922)            Eocene

             Grimaldi et al. (1997)        Paleocene or Early Eocene

               Zherikhin & Ross (2000) – the fossil beds reworked
                                         (Late Cretaceous)
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)
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)
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
So, how to overcome these problems?


                 Paleontologists   Lepidopterists

   Geological                                        Incomplete
   Uncertainty                                      Preservation
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
AToL (Alcohol Tubes of Lepidoptera)
Build a frozen tissue collection for all to use
        Molecular “backbone”          Twig Leaders
        phylogeny and dating
         26 nuclear genes
         127 families

                                           • Compile current knowledge
                                           relevant to phylogeny: character
                                           descriptions & images, data
                                           matrices, trees, literature, primers,
Lepidoptera                                availability of material…
   fossil            Collaborative
                                           • Compile morphological
 database              Website             homologies glossary
                                           with reconciled terminologies,
                                           providing characters for future
        AToLep                             higher phylogeny studies
        Collection             Training
How does “Fossil Project” progress?
STEP I                                   STEP III
                    STEP II              Critical reviews of
A preliminary
catalogue and       Constructing         the records in the
data-base of the    fossil image data-   fossil data-base
known fossil        base                 by the invited
Lepidoptera                              experts and

 STEP VI            STEP V               STEP IV
 Improving our      Providing dates      A comprehensive
 knowledge of the   for molecular        data-base &
 evolutionary       dating in the        Present/absent
 history of         Lepidoptera          chart along
 Lepidoptera        phylogeny            geologic time
Launched Website

                                  Family Name

                               Species Name

      Synopsis of Leptree fossil data base entries
  Composition by identification category                       Superfamily composition (439 specimens)
                       Categories                     No.
Assigned to Lepidoptera, with supporting evidence     594                                                                  6%
Tentatively assigned to Lepidoptera                   16                4%
Assigned to Lepidoptera, no supporting evidence       28               Tortricoidea
                                                                            5%                                                Micropterigoidea
Assigned to but subsequently excluded from            27                              Tineoidea                                     5%
Lepidoptera                                                                              11%
                          Total                       665
                                                                                Papilionoidea            Noctuoidea
                                                                                     15%                    9%

 Composition by preservation type                               Composition by geological age
 (526 specimens)
                                            Sieved    350                                               316
             2%                                       300


                compression/         44%              150                                                               128
                    53%                               100
                                                               1              10
                                                            Triassic       Jurassic     Cretaceous   Paleogene       Neogene
Species information sheet
1) Body fossil
 - Taxonomic information
 - 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
Interaction in “Fossil Project”
* “comments” section in each sheet
  - To find new evidence of the known fossil
    ex) Axel Hausmann’s 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)
LepTree Twig leaders        Smithsonian Paleontology
                             Finnegan Marsh
LepTree Collaborators
 Axel Hausmann               Wenying Wu
 Joaquin Baixeras

LepTree Informatics Staff
 John Park
 Giho Kim

  Erika Koenig

Helper for Translation
 Kim Mitter
 Ada Szczepaniec