Scott Hartman
← Research

Origin of avian flight

Moving past the ground-up versus trees-down debate by testing flight-origin models against a time-calibrated paravian record.

Time-calibrated phylogeny of paravian theropods, with avialans highlighted in blue
Time-calibrated phylogeny showing multiple origins of theropod aerial behavior, and a relatively late origin of avian flight.

The origin of birds from theropod dinosaurs is one of the great locomotory transitions in the history of life. Unlike the situation with basal members of the other flying vertebrate clades, bats and pterosaurs, the fossil record of early flying dinosaurs is remarkably complete, including excellent preservation of taxa with incipient flight anatomy. That means the ecological context and selective pressures that drove the transition to avian flight can be documented in a detail rarely available to us.

I have been working on the origin of birds for over two decades. My first platform presentation, at the 2000 Society of Vertebrate Paleontology meeting, was on the topic (Primary and caudal feathers as locomotory adaptations in maniraptoran theropods). Ongoing goals include disentangling the “ground up versus trees down” debate, building a framework that lets competing models of bird origins be tested more explicitly against the fossil record, and documenting how the history of dinosaur research and typological thinking has shaped modern ideas about bird origins (spoiler: not for the better).

Papers and presentations from this work, and the colleagues who have collaborated on it, are listed below.

Papers

  1. 2025

    Forelimb feathering, soft tissues, and skeleton of the flying dromaeosaurid Microraptor

    Grosmougin M, Wang X, Zheng X, Kaye TG, Chotard M, Barlow LA, Dececchi TA, Habib MB, Zariwala J, Hartman SA, Xu X, Pittman M

    BMC Ecology and Evolution 25(1): 6510.1186/s12862-025-02397-5

  2. 2025

    New information on the hind limb feathering, soft tissues and skeleton of Microraptor (Theropoda: Dromaeosauridae)

    Chotard M, Wang X, Zheng X, Kaye TG, Grosmougin M, Barlow L, Kundrát M, Dececchi TA, Habib MB, Zariwala J, Hartman SA, Xu X, Pittman M

    BMC Ecology and Evolution 25(1): 3710.1186/s12862-025-02372-0

  3. 2022

    Preserved soft anatomy confirms shoulder-powered upstroke of early theropod flyers, reveals enhanced early pygostylian upstroke, and explains early sternum loss

    Pittman M, Kaye TG, Wang X, Zheng X, Dececchi TA, Hartman SA

    Proceedings of the National Academy of Sciences 119(47): e220547611910.1073/pnas.2205476119

  4. 2019

    A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight

    Hartman S, Mortimer M, Wahl WR, Lomax DR, Lippincott J, Lovelace DM

    PeerJ 7: e724710.7717/peerj.7247

  5. 2017

    Basal paravian functional anatomy illuminated by high-detail body outline

    Wang X, Pittman M, Zheng X, Kaye TG, Falk AR, Hartman SA, Xu X

    Nature Communications 8(1): 1457610.1038/ncomms14576

  6. 2007

    The tenth skeletal specimen of Archaeopteryx

    MAYR G, POHL B, HARTMAN S, PETERS DS

    Zoological Journal of the Linnean Society 149(1): 97-11610.1111/j.1096-3642.2006.00245.x

Conference abstracts & talks

  1. 2025

    New Microraptor specimens bring insight into the forewing anatomy and its impact on flight behavior and ecology

    Grosmougin M, Wang X, Zheng X, Kaye TG, Chotard M, Barlow LA, Dececchi TA, Habib MB, Zariwala J, Hartman SA, Xu X, Pittman M

    International Congress of Vertebrate Morphology

  2. 2025

    Transition from occasional to habitual theropod flyers: origin of regular flight use in early birds and their close relatives

    Dececchi TA, Habib MB, Hartman SA, Larsson HCE, Pittman M

    International Congress of Vertebrate Morphology

  3. 2021

    Operationalizing a late origin of avian flight

    Hartman SA, Fitch A

    Northeast Natural History Conference (online)

  4. 2019

    Internal and external flight-related anatomy of early theropod fliers revealed by laser-stimulated fluorescence fills knowledge gaps in functional morphology and flight capability

    Pittman M, Kaye TG, Wang X, Zheng X, Hartman SA, Xu X

    Society of Vertebrate Paleontology 79th Annual Meeting

  5. 2019

    Origin of Pterosauria and stepwise evolution of the pterosaurian flight apparatus

    Fitch A, Pritchard A, Hartman SA, Bevitt J, Lovelace DM, Bhullar BA

    Geological Society of America Annual Meeting, Phoenix10.1130/abs/2019AM-340860

  6. 2018

    A testable macroevolutionary framework for character acquisition in the origin of avian flight

    Hartman SA, Mortimer M, Lovelace DM

    Society of Vertebrate Paleontology 78th Annual Meeting

  7. 2018

    Adaptive diversity in the early evolution of theropod flight

    Pittman M, Wang X, Kaye TG, Zheng X, Hartman SA, Navalón G, Xu X

    International Pennaraptoran Dinosaur Symposium, Hong Kong

  8. 2018

    Laser-stimulated fluorescence reveals preserved soft tissues which unravel the complexity of avian flight evolution

    Pittman M, Kaye TG, Wang X, Zheng X, Hartman SA, Navalón G, Xu X

    5th International Palaeontological Congress (IPC5), Paris

  9. 2017

    New information on a paravian theropod from the Morrison Formation

    Hartman SA, Mortimer M, Wahl WR, Lomax DR, Lovelace DM

    Society of Vertebrate Paleontology 77th Annual Meeting

  10. 2016

    Nothing but skin and bones: Anchiornis body outline revealed by laser fluorescence

    Kaye TG, Pittman M, Wang X, Zheng X, Xu X, Hartman SA, Falk AR

    Society of Vertebrate Paleontology 76th Annual Meeting

  11. 2007

    Re-evaluating wing shape in Archaeopteryx: information from the Thermopolis specimen

    Hartman SA

    Society of Vertebrate Paleontology Annual Meeting

  12. 2005

    Phylogenetic assessment of a maniraptoran from the Morrison Formation

    Hartman SA, Lovelace DM, Wahl WR

    Society of Vertebrate Paleontology Annual Meeting

  13. 2000

    Primary and caudal feathers as locomotory adaptations in maniraptoran theropods

    Hartman SA

    Society of Vertebrate Paleontology Annual Meeting