Paleotaxonomy is more than describing and naming new species. When we discover new species, we discover new organisms with unique morphologies and behaviors that will add to our understanding of life and its evolution.
For example, the tiny flies of the genus Synneuron (family Canthyloscelidae) are easily recognized by the unique "X"-shape formed by the anterior veins of their wings. They are exceedingly rare; only two species exist today. Synneuron decipiens is found only in northwestern North America (Colorado, Washington, Alaska and Canada) while Synneuron annulipes has been found only in Finland and Russia. When the fossil Synneuron eomontana was found in the Kishenehn Formation, a review of the literature located another specimen that had been misidentified as a fungus gnat. It really belonged to the genus Synneuron. It was from the Cretaceous of Australia and was named S. jelli by Dalton de Souza Amorim, my collaborator at the University of São Paulo in Brazil. These two specimens are the only fossils of the genus.
For example, the tiny flies of the genus Synneuron (family Canthyloscelidae) are easily recognized by the unique "X"-shape formed by the anterior veins of their wings. They are exceedingly rare; only two species exist today. Synneuron decipiens is found only in northwestern North America (Colorado, Washington, Alaska and Canada) while Synneuron annulipes has been found only in Finland and Russia. When the fossil Synneuron eomontana was found in the Kishenehn Formation, a review of the literature located another specimen that had been misidentified as a fungus gnat. It really belonged to the genus Synneuron. It was from the Cretaceous of Australia and was named S. jelli by Dalton de Souza Amorim, my collaborator at the University of São Paulo in Brazil. These two specimens are the only fossils of the genus.
The ancient continent of Pangaea approximately 250 million years ago.
How can the occurrence of a species so far away from the rest of its close relatives in North America be explained? The answer lies in knowing that the Earth's geography has changed drastically over geologic time. The shape of the ancient continent of Pangaea, shown in the map at the right, helps to explain. When all of the land masses of the Earth were connected 250 million years ago, species of Synneuron must have been plentiful and very dispersed all over the world. In the map of Pangaea at the right, the extant species are represented by open and closed circles; the two fossil species are depicted by stars.
Documentation of the diversity of the insects that lived near the subtropical Kishenehn lakeshore 46 million years ago in what is now northwestern Montana is an important goal of my research. Unfortunately, we have barely begun; work to date on the Kishenehn Formation has described only a small fraction of its insect fauna.
My collaborators and I have recently concentrated on the diversity of the order Diptera (flies - insects with only two wings [from the Greek words "di" = two and "pteron" = wing]). We have identified a total of 34 different families of flies in the Kishenehn Formation fossil insect collection.
My collaborators and I have recently concentrated on the diversity of the order Diptera (flies - insects with only two wings [from the Greek words "di" = two and "pteron" = wing]). We have identified a total of 34 different families of flies in the Kishenehn Formation fossil insect collection.
A list of the 33 families (in red) of flies (Diptera) collected at the KIshenehn Formation.
A highly diverse and relatively basal group of flies, including the crane flies, mosquitos, gnats, robber flies, bee flies, snipe flies, dance flies and flower flies, are listed in the table above. Of the 70 families listed, 33 (in red) have been recorded in the Kishenehn Formation - eight of them (underlined) have not been found at other North American Eocene fossil sites. An additional eight families of this group, though not recorded from the Kishenehn Formation, have been found in one or more of the Green River, Florissant and Okanagan localities.
One can divide the approximately 150 families of Diptera into the group described above and the Schizophora, about 80 families of more advanced flies that use a ballon-like structure to apply pressure to their pupal case; with enough pressure, the case breaks and the adult escapes. Perhaps the most interesting observation is the scarcity of identified Schizophora in the Kishenehn Collection; only one family, Lauxaniidae, has been definitively identified. This contrasts with the 15 families of Schizophora that have been recorded at the Green River, Florissant and Okanagan localities. The basis for these numbers appears to relate to the way the insects were preserved. Families found only in the Kishenehn Formation are often very tiny - and the Kishenehn fossils display a strong bias towards the preservation of very tiny insects. A corollary of this is that a number of the Schizophora are large and robust (e.g. house flies, flesh flies, blue-bottle flies and dung flies) and are not well-preserved in the thin shales of the Kishenehn Formation; although several families of these larger flies are present, the morphological features required for definitive identification are absent. Fossils of the Schizophora will be a major focus of our future research.
The following is a list of new insect species that we have named from the Kishenehn Formation.
Eoeustochus Huber, gen. n.
Eoeustochus kishenehn Huber sp. n.
Eoeustochus borchersi Huber, sp. n.
Eoanaphes Huber, gen. n.
Eoanaphes stethynioides Huber, sp. n.
Gonatocerus greenwalti Huber, sp. n.
Gonatocerus kootenai Huber, sp. n.
Gonatocerus rasnitsyni Huber, sp. n.
Ptenidium kishenehnicum Shockley and Greenwalt, sp. n.
Culiseta kishenehn, Harbach, sp. n.
Culiseta lemniscata Harbach, sp. n.
Pseudotettigonia leona Greenwalt and Rust, sp.n.
Tettigonia bricei Greenwalt and Rust, sp.n.
Dolichoderus dlusskyi LaPolla, sp. n.
Ktunaxia LaPolla, gen. n.
Ktunaxia jucunda LaPolla, sp. n
Proiridomyrmex rotundatus LaPolla, sp. n.
Protazteca eocenica LaPolla, sp. n.
Formica annosa LaPolla, sp. n.
Lasius glom LaPolla, sp. n.
Crematogaster aurora LaPolla, sp. n.
Solenopsites abdita LaPolla, sp. n.
Ponerites kishenehne LaPolla, sp. n.
Pseudomyrmex saxulum LaPolla, sp. n.
Eoformica brevipetiola LaPolla, sp. n.
Eoformica latimedia LaPolla, sp. n.
Eogyropsylla paveloctogenarius Ouvrard and Burckhardt, sp. n.
Phasmatopelecinus Greenwalt and Engel, gen. n.
Phasmatopelecinus leonae Greenwalt and Engel, sp. n.
Latiblattella avita Vidlička and Greenwalt, sp. n.
Anthrax succini Greenwalt and Evenhuis, sp. n.
Eoanomala Greenwalt and Evenhuis, gen. n.
Eoanomala melas Greenwalt and Evenhuis, sp. n.
Lutetialestes uniformis Bechly and Greenwalt, sp. n.
Eolestidae Bechly and Greenwalt, fam. n.
Eolestes syntheticus Bechly and Greenwalt, sp. n.
Eolestes ramosus Bechly and Greenwalt, sp. n.
Dixella intacta Greenwalt and Moulton, sp.n.
Dixella eomarginata Greenwalt and Moulton, sp. n.
Dixella spinilobata Greenwalt and Moulton, sp. n.
Dixella curvistyla Greenwalt and Moulton, sp. n.
Bolitophila warreni Greenwalt and Blagoderov, sp. n.
Tipula (Trichotipula) fji De Jong, sp. n.
Ellipteroides (Ellipteroides) kishenehn De Jong, sp. n.
Cyttaromyia lynnae De Jong, sp. n.
Cyttaromyia fuscula De Jong, comb. n.
Sylvicola silibrarius Greenwalt, sp. n.
Efcookella nigra Greenwalt, sp. n.
Bibiodes kishenehnensis Fitzgerald, sp. n.
Eosciarites hermaea Greenwalt, sp. n.
Rymosia hypnolithica Kerr, sp. n.
Litoleptis araeostylus Greenwalt, sp. n.
Kishenehnoasilus bhl Dikow gen. n.
Kishenehnoasilus bhl Dikow sp. n.
Salishomyia eocenica Bickel, gen. n.
Salishomyia eocenica Bickel, sp. n.
Drapetis adelomedos Greenwalt, sp. n.
Agathomyia eocenica Tkoč sp. n.
Aenigmatias kishenehnensis Brown sp. n.
Lonchoptera eocenica Amorim and Brown, sp. nov.
Synneuron eomontana Amorim and Greenwalt sp. nov.
Synneuron jelli Amorim and Greenwalt sp. nov.
The following is a list of publications on paleotaxonomy from our laboratory with , where possible, links to pdfs of the manuscripts.
Huber, J.T. and D. Greenwalt. 2011. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales, with description of two new genera and review of Tertiary amber genera. ZooKeys, 130:473‒494. Link to pdf
Harbach, R. E. and D. Greenwalt. 2012. Two Eocene species of Culiseta (Diptera: Culicidae) from the Kishenehn Formation in Montana. Zootaxa, 3530:25-34.
Shockley, F. W. and D. Greenwalt. 2013. Ptenidium kishenehnsis, a new fossil described from the Kishenehn oil shales (Coleoptera: Ptiliidae), with a checklist of previously known fossil ptiliids. Proceedings of the Entomological Society of Washington, 115 (2):173-181.
Ouvrard, D., Burckhardt, D. and Greenwalt, D. 2013. The oldest jumping plant-louse (Insecta: Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea. Acta Musei Moraviae, Scientiae biologicae (Brno), 98(2):21-33.
Greenwalt, D. and Rust, J. 2014. A new species of Pseudotettigonia Zeuner 1937 (Orthoptera:Tettigoniidae) with an intact stridulatory field and reexamination of the subfamily Pseudotettigoniinae. Systematic Entomology, 39 (2):256-263.
Greenwalt, D. and Michael S. Engel, M. S. 2014. A diminutive pelecinid wasp from the Eocene Kishenehn Formation of northwestern Montana (Hymenoptera: Pelecinidae). Novitates Paleoentomologicae, 8:1-9.
Link to pdf
Greenwalt, D. E. and Bechly, G. 2014. A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America. Zootaxa, 3887 (2):138–156.
LaPolla, J.S. and Greenwalt, D.E. 2015. Fossil Ants (Hymenoptera: Formicidae) of the Middle Eocene Kishenehn Formation. Sociobiology, 62(2):163-174. Link to pdf
Greenwalt, D.E. and Vidlička, L. 2015. Latiblattella avita sp.n. (Blattaria: Ectobiidae) from the Eocene Kishenehn Formation, Montana, U.S.A. Palaeontologia Electronica, 18(1):16A. Link to pdf
Greenwalt D.E., Wingerath J.G. and Evenhuis N.L. 2015. Two new and disparate fossil bee flies (Bombyliidae: Anthracinae) from the Americas and reassessment of Anthrax dentoni Lewis, 1969. Palaeontologia Electronica, 18(3):1-10 (51A). Link to pdf
Greenwalt, D.E. and Moulton, J.K. 2016. The first fossil New World Dixidae with a critical discussion of generic definitions. Palaeontologia Electronica, 19(3):1-32. Link to pdf
Greenwalt, D.E and Blagoderov, V.A. 2019. Review of the fossil record of Bolitophilidae (Diptera: Sciaroidea). Zootaxa, 4567(3): 546-560.
Greenwalt, D.E., and Bickel, D.J., Kerr, P.H., Curler, G.R., Brown, B.V., de Jong, H., Scott J. Fitzgerald, S.J., Dikow, T., Tkoč, M., Kehlmaier, C. and Amorim, D.S. 2019. Diptera of the Kishenehn Formation. I. Documentation of diversity at the family level. Palaeontologia Electronica, 22.2.50. Link to pdf
Amorim, D. De S. and Greenwalt, D. E. 2020. Cretaceous and Eocene fossils of the rare extant genus Synneuron (Diptera: Canthyloscledidae): evidence of a true Pangean clade. Cladistics. 36(4): 413-423.
Additional manuscripts from other laboratories are listed below. The 1989 paper by Kurt Constenius et al., was the first to describe fossil insects from the Kishenehn Formation.
Constenius, K.N., Dawson, M.R., Pierce, H.G., Walter, R.C., and Wilson, M.V.H. 1989. Reconnaissance paleontologic study of the Kishenehn Formation, northwestern Montana and southeastern British Columbia. In: D.E. French and R.F. Grabb (eds.), 1989 Field Conference Guidebook: Montana Centennial Edition, Vol. 1, 189–203. Geological Resources of Montana, Billings.
Krzemiński, W. 1991. A first fossil Helius (Diptera, Limoniidae) from North America. Acta Zoologica Cracoviensia, 34(1):311-313.
Talamas EJ, Buffington ML (2015) Fossil Platygastroidea in the National Museum of Natural History, Smithsonian Institution. Journal of Hymenoptera Research 47: 1–52.
Brunke, A.J., Schillhammer, H. and Chatzimanolis, S., 2017. The first fossil rove beetle from the middle Eocene Kishenehn Formation (North America) provides evidence for ancient Eocene relicts within the hyperdiverse Staphylinini (Coleoptera: Staphylinidae: Staphylininae). Journal of Systematic Palaeontology, 15(12), pp.1015-1025.
Makarkin, V., Archibald, S., & Jepson, J. (2019). The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America. The Canadian Entomologist, 1-10.
Eoeustochus Huber, gen. n.
Eoeustochus kishenehn Huber sp. n.
Eoeustochus borchersi Huber, sp. n.
Eoanaphes Huber, gen. n.
Eoanaphes stethynioides Huber, sp. n.
Gonatocerus greenwalti Huber, sp. n.
Gonatocerus kootenai Huber, sp. n.
Gonatocerus rasnitsyni Huber, sp. n.
Ptenidium kishenehnicum Shockley and Greenwalt, sp. n.
Culiseta kishenehn, Harbach, sp. n.
Culiseta lemniscata Harbach, sp. n.
Pseudotettigonia leona Greenwalt and Rust, sp.n.
Tettigonia bricei Greenwalt and Rust, sp.n.
Dolichoderus dlusskyi LaPolla, sp. n.
Ktunaxia LaPolla, gen. n.
Ktunaxia jucunda LaPolla, sp. n
Proiridomyrmex rotundatus LaPolla, sp. n.
Protazteca eocenica LaPolla, sp. n.
Formica annosa LaPolla, sp. n.
Lasius glom LaPolla, sp. n.
Crematogaster aurora LaPolla, sp. n.
Solenopsites abdita LaPolla, sp. n.
Ponerites kishenehne LaPolla, sp. n.
Pseudomyrmex saxulum LaPolla, sp. n.
Eoformica brevipetiola LaPolla, sp. n.
Eoformica latimedia LaPolla, sp. n.
Eogyropsylla paveloctogenarius Ouvrard and Burckhardt, sp. n.
Phasmatopelecinus Greenwalt and Engel, gen. n.
Phasmatopelecinus leonae Greenwalt and Engel, sp. n.
Latiblattella avita Vidlička and Greenwalt, sp. n.
Anthrax succini Greenwalt and Evenhuis, sp. n.
Eoanomala Greenwalt and Evenhuis, gen. n.
Eoanomala melas Greenwalt and Evenhuis, sp. n.
Lutetialestes uniformis Bechly and Greenwalt, sp. n.
Eolestidae Bechly and Greenwalt, fam. n.
Eolestes syntheticus Bechly and Greenwalt, sp. n.
Eolestes ramosus Bechly and Greenwalt, sp. n.
Dixella intacta Greenwalt and Moulton, sp.n.
Dixella eomarginata Greenwalt and Moulton, sp. n.
Dixella spinilobata Greenwalt and Moulton, sp. n.
Dixella curvistyla Greenwalt and Moulton, sp. n.
Bolitophila warreni Greenwalt and Blagoderov, sp. n.
Tipula (Trichotipula) fji De Jong, sp. n.
Ellipteroides (Ellipteroides) kishenehn De Jong, sp. n.
Cyttaromyia lynnae De Jong, sp. n.
Cyttaromyia fuscula De Jong, comb. n.
Sylvicola silibrarius Greenwalt, sp. n.
Efcookella nigra Greenwalt, sp. n.
Bibiodes kishenehnensis Fitzgerald, sp. n.
Eosciarites hermaea Greenwalt, sp. n.
Rymosia hypnolithica Kerr, sp. n.
Litoleptis araeostylus Greenwalt, sp. n.
Kishenehnoasilus bhl Dikow gen. n.
Kishenehnoasilus bhl Dikow sp. n.
Salishomyia eocenica Bickel, gen. n.
Salishomyia eocenica Bickel, sp. n.
Drapetis adelomedos Greenwalt, sp. n.
Agathomyia eocenica Tkoč sp. n.
Aenigmatias kishenehnensis Brown sp. n.
Lonchoptera eocenica Amorim and Brown, sp. nov.
Synneuron eomontana Amorim and Greenwalt sp. nov.
Synneuron jelli Amorim and Greenwalt sp. nov.
The following is a list of publications on paleotaxonomy from our laboratory with , where possible, links to pdfs of the manuscripts.
Huber, J.T. and D. Greenwalt. 2011. Compression fossil Mymaridae (Hymenoptera) from Kishenehn oil shales, with description of two new genera and review of Tertiary amber genera. ZooKeys, 130:473‒494. Link to pdf
Harbach, R. E. and D. Greenwalt. 2012. Two Eocene species of Culiseta (Diptera: Culicidae) from the Kishenehn Formation in Montana. Zootaxa, 3530:25-34.
Shockley, F. W. and D. Greenwalt. 2013. Ptenidium kishenehnsis, a new fossil described from the Kishenehn oil shales (Coleoptera: Ptiliidae), with a checklist of previously known fossil ptiliids. Proceedings of the Entomological Society of Washington, 115 (2):173-181.
Ouvrard, D., Burckhardt, D. and Greenwalt, D. 2013. The oldest jumping plant-louse (Insecta: Hemiptera: Sternorrhyncha) with comments on the classification and nomenclature of the Palaeogene Psylloidea. Acta Musei Moraviae, Scientiae biologicae (Brno), 98(2):21-33.
Greenwalt, D. and Rust, J. 2014. A new species of Pseudotettigonia Zeuner 1937 (Orthoptera:Tettigoniidae) with an intact stridulatory field and reexamination of the subfamily Pseudotettigoniinae. Systematic Entomology, 39 (2):256-263.
Greenwalt, D. and Michael S. Engel, M. S. 2014. A diminutive pelecinid wasp from the Eocene Kishenehn Formation of northwestern Montana (Hymenoptera: Pelecinidae). Novitates Paleoentomologicae, 8:1-9.
Link to pdf
Greenwalt, D. E. and Bechly, G. 2014. A re-description of the fossil damselfly Eolestes syntheticus Cockerell, 1940 (Odonata: Zygoptera: Eolestidae n. fam.) with description of new taxa from the Eocene of North America. Zootaxa, 3887 (2):138–156.
LaPolla, J.S. and Greenwalt, D.E. 2015. Fossil Ants (Hymenoptera: Formicidae) of the Middle Eocene Kishenehn Formation. Sociobiology, 62(2):163-174. Link to pdf
Greenwalt, D.E. and Vidlička, L. 2015. Latiblattella avita sp.n. (Blattaria: Ectobiidae) from the Eocene Kishenehn Formation, Montana, U.S.A. Palaeontologia Electronica, 18(1):16A. Link to pdf
Greenwalt D.E., Wingerath J.G. and Evenhuis N.L. 2015. Two new and disparate fossil bee flies (Bombyliidae: Anthracinae) from the Americas and reassessment of Anthrax dentoni Lewis, 1969. Palaeontologia Electronica, 18(3):1-10 (51A). Link to pdf
Greenwalt, D.E. and Moulton, J.K. 2016. The first fossil New World Dixidae with a critical discussion of generic definitions. Palaeontologia Electronica, 19(3):1-32. Link to pdf
Greenwalt, D.E and Blagoderov, V.A. 2019. Review of the fossil record of Bolitophilidae (Diptera: Sciaroidea). Zootaxa, 4567(3): 546-560.
Greenwalt, D.E., and Bickel, D.J., Kerr, P.H., Curler, G.R., Brown, B.V., de Jong, H., Scott J. Fitzgerald, S.J., Dikow, T., Tkoč, M., Kehlmaier, C. and Amorim, D.S. 2019. Diptera of the Kishenehn Formation. I. Documentation of diversity at the family level. Palaeontologia Electronica, 22.2.50. Link to pdf
Amorim, D. De S. and Greenwalt, D. E. 2020. Cretaceous and Eocene fossils of the rare extant genus Synneuron (Diptera: Canthyloscledidae): evidence of a true Pangean clade. Cladistics. 36(4): 413-423.
Additional manuscripts from other laboratories are listed below. The 1989 paper by Kurt Constenius et al., was the first to describe fossil insects from the Kishenehn Formation.
Constenius, K.N., Dawson, M.R., Pierce, H.G., Walter, R.C., and Wilson, M.V.H. 1989. Reconnaissance paleontologic study of the Kishenehn Formation, northwestern Montana and southeastern British Columbia. In: D.E. French and R.F. Grabb (eds.), 1989 Field Conference Guidebook: Montana Centennial Edition, Vol. 1, 189–203. Geological Resources of Montana, Billings.
Krzemiński, W. 1991. A first fossil Helius (Diptera, Limoniidae) from North America. Acta Zoologica Cracoviensia, 34(1):311-313.
Talamas EJ, Buffington ML (2015) Fossil Platygastroidea in the National Museum of Natural History, Smithsonian Institution. Journal of Hymenoptera Research 47: 1–52.
Brunke, A.J., Schillhammer, H. and Chatzimanolis, S., 2017. The first fossil rove beetle from the middle Eocene Kishenehn Formation (North America) provides evidence for ancient Eocene relicts within the hyperdiverse Staphylinini (Coleoptera: Staphylinidae: Staphylininae). Journal of Systematic Palaeontology, 15(12), pp.1015-1025.
Makarkin, V., Archibald, S., & Jepson, J. (2019). The oldest Inocelliidae (Raphidioptera) from the Eocene of western North America. The Canadian Entomologist, 1-10.