ScholarSphere

Archive of digital videos showing insect surface skimming behavior

Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half an eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?

Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.

These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.

Papers:

Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution. Science 266, 427-430.

Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.

Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.

Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.

Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.

Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.

Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.

Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.

Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.

Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.

Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.

Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.

Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.

Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.

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Metadata

Work Title Archive of digital videos showing insect surface skimming behavior
Access
Open Access
Creators
  1. James Marden
Keyword
  1. Evolution
  2. Flight
  3. Insects
  4. Protowings
  5. Wings
  6. Aquatic
  7. Skimming
  8. Surface-skimming
  9. Complexity
  10. Pterygota
  11. Pterygotes
  12. Fossils
  13. Entomology
  14. Stoneflies
  15. Plecoptera
  16. Mayflies
  17. Ephemeroptera
  18. Gills
  19. Wings from gills
License CC BY 4.0 (Attribution)
Work Type Project
Acknowledgments
  1. NASA NGT-40002 (1994-1995; "Mechanics of insect flight: an interdisciplinary examination of evolution, development, and physiology”.) NASA NGT-40002 (1996-1997; “Evolution of insect flight”.) NSF IBN-9600840 (1997-1999; “Integrative biology of insect muscle: molecular through ecological investigations of locomotor performance”) NSF IBN-9722196 (1997-2002; Career Development Award: “Evolution of Insect Flight”)
Publication Date June 18, 2025
DOI doi:10.26207/2xvh-jj02
Related URLs
  1. https://doi.org/10.26207/2xvh-jj02
Deposited June 18, 2025

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Work History

Version 1
published

  • Created

    June 18, 2025 13:14 by jhm10

  • Updated

    June 18, 2025 13:14 by [unknown user]

  • Updated Description, Publication Date Show Changes

    June 18, 2025 14:21 by jhm10

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half and eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution. Science 266, 427-430.
    • - movie: Six-leg skimming
    • - Television coverage: Scientific American Frontiers, see 34:25 to 39:48 of this external link, https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1)
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.
    • - movie: Sailing
    • Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.
    • Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.
    • - movie: Hind-leg skimming
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.
    • - movies: Gill flapping, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.
    • movies: Gill flapping, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.
    • - movie: Rowing
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • -movies: Gill flapping, Rowing, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.
    • - movie: Skim onto mud, Gill flap homology
    • Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.
    Publication Date
    • 2025-06-18
  • Updated Acknowledgments Show Changes

    June 18, 2025 14:23 by jhm10

    Acknowledgments
    • NASA NGT-40002 (1994-1995; "Mechanics of insect flight: an interdisciplinary examination of evolution, development, and physiology”.) NASA NGT-40002 (1996-1997; “Evolution of insect flight”.) NSF IBN-9600840 (1997-1999; “Integrative biology of insect muscle: molecular through ecological investigations of locomotor performance”) NSF IBN-9722196 (1997-2002; Career Development Award: “Evolution of Insect Flight”)
  • Added Creator James Marden

    June 18, 2025 14:23 by jhm10

  • Added Six-leg skimming.mp4

    June 18, 2025 14:26 by jhm10

  • Added Sailing.mp4

    June 18, 2025 14:26 by jhm10

  • Added Hind-leg skimming.mp4

    June 18, 2025 14:26 by jhm10

  • Added Gill flapping.mp4

    June 18, 2025 14:26 by jhm10

  • Added Swim-skim.mp4

    June 18, 2025 14:26 by jhm10

  • Added Four-leg skimming.mp4

    June 18, 2025 14:26 by jhm10

  • Added Mayfly four-leg skimming.mp4

    June 18, 2025 14:26 by jhm10

  • Added Jump.mp4

    June 18, 2025 14:26 by jhm10

  • Added Rowing.mp4

    June 18, 2025 14:26 by jhm10

  • Added Gill flap homology.mp4

    June 18, 2025 14:26 by jhm10

  • Added Skim onto mud.mp4

    June 18, 2025 14:26 by jhm10

  • Updated Description, License Show Changes

    June 18, 2025 14:28 by jhm10

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half and eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution. Science 266, 427-430.
    • - movie: Six-leg skimming
    • - Television coverage: Scientific American Frontiers, see 34:25 to 39:48 of this external link, https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1)
    • - Television: Scientific American Frontiers, see 34:25 to 39:48 at this external link, https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1)
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.
    • - movie: Sailing
    • Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.
    • Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.
    • - movie: Hind-leg skimming
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.
    • - movies: Gill flapping, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.
    • movies: Gill flapping, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.
    • - movie: Rowing
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • -movies: Gill flapping, Rowing, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.
    • - movie: Skim onto mud, Gill flap homology
    • Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.
    License
    • https://creativecommons.org/licenses/by/4.0/
  • Published

    June 18, 2025 14:28 by jhm10

  • Updated

    June 18, 2025 22:06 by [unknown user]

  • Updated Related URLs Show Changes

    June 19, 2025 11:03 by avs5190

    Related URLs
    • https://doi.org/10.26207/2xvh-jj02
  • Updated Description Show Changes

    June 19, 2025 11:04 by avs5190

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half and eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half an eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution. Science 266, 427-430.
    • - movie: Six-leg skimming
    • - Television: Scientific American Frontiers, see 34:25 to 39:48 at this external link, https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1)
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.
    • - movie: Sailing
    • Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.
    • Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.
    • - movie: Hind-leg skimming
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.
    • - movies: Gill flapping, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.
    • movies: Gill flapping, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.
    • - movie: Rowing
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • -movies: Gill flapping, Rowing, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.
    • - movie: Skim onto mud, Gill flap homology
    • Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.
  • Updated Description Show Changes

    June 24, 2025 10:27 by avs5190

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half an eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution. Science 266, 427-430.
    • - movie: Six-leg skimming
    • - Television: Scientific American Frontiers, see 34:25 to 39:48 at this external link, https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1)
    • Marden, J.H. and M.G. Kramer. 1994. [Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution.](https://doi.org/10.1126/science.266.5184.427) Science 266, 427-430.
    • - movie: [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true)
    • - Television: [Scientific American Frontiers](https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1), see 34:25 to 39:48
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.
    • - movie: Sailing
    • Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.
    • Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.
    • - movie: Hind-leg skimming
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.
    • - movies: Gill flapping, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.
    • movies: Gill flapping, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.
    • - movie: Rowing
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • -movies: Gill flapping, Rowing, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.
    • - movie: Skim onto mud, Gill flap homology
    • Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.
  • Updated Description Show Changes

    June 24, 2025 11:00 by avs5190

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half an eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. [Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution.](https://doi.org/10.1126/science.266.5184.427) Science 266, 427-430.
    • Marden, J.H. and M.G. Kramer. 1994. [Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution](https://doi.org/10.1126/science.266.5184.427). Science 266, 427-430.
    • - See also: Kaiser, J. 1994. [A new theory of insect wing origins takes off](https://doi.org/10.1126/science.266.5184.363). Science 266, 363.
    • - movie: [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true)
    • - Television: [Scientific American Frontiers](https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1), see 34:25 to 39:48
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects with rudimentary wings. Nature 377, 332-334.
    • - movie: Sailing
    • Marden, J.H. and M.G. Kramer. 1995. [Locomotor performance of insects with rudimentary wings](https://doi.org/10.1038/377332a0). Nature 377, 332-334.
    • - movie: [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true)
    • Marden, J.H. 1995. [How insects learned to fly](https://ezaccess.libraries.psu.edu/login?url=https://www.proquest.com/scholarly-journals/how-insects-learned-fly/docview/212702879/se-2?accountid=13158). The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. [Plecopteran surface-skimming and insect flight evolution - reply](https://doi.org/10.1126/science.270.5242.1685). Science 270, 1685.
    • Marden, J.H. 1995. How insects learned to fly. The Sciences 35, 26-30.
    • Kramer, M.G. and J.H. Marden. 1997. [Almost airborne](https://doi.org/10.1038/385403a0). Nature 385, 403-404.
    • Marden, J.H. and M.G. Kramer. 1995. Plecopteran surface-skimming and insect flight evolution - reply. Science 270, 1685.
    • - movie: [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true)
    • Kramer, M.G. and J.H. Marden. 1997. Almost airborne. Nature 385, 403-404.
    • - movie: Hind-leg skimming
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. [Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion](https://doi.org/10.1086/318109). Physiological and Biochemical Zoology 73, 751-764.
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion. Physiological and Biochemical Zoology 73, 751-764.
    • - movies: Gill flapping, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior. Proceedings of the National Academy of Sciences 97:13178-13183.
    • movies: Gill flapping, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. [Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior](https://doi.org/10.1073/pnas.230296997). Proceedings of the National Academy of Sciences 97:13178-13183.
    • Marden, J.H. and M.A. Thomas. 2003. Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills. Biological Journal of the Linnean Society 79: 341–349.
    • - movie: Rowing
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Marden, J.H. and M.A. Thomas. 2003. [Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills](https://doi.org/10.1046/j.1095-8312.2003.00192.x). Biological Journal of the Linnean Society 79: 341–349.
    • - movie: [Rowing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43075?download=true)
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • -movies: Gill flapping, Rowing, Sailing, Swim-skim, Six-leg skimming, Four-leg skimming, Mayfly four-leg skimming, Hind-leg skimming, Jump
    • Marden, J.H. 2008. Evolution and physiology of flight in aquatic insects. In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Rowing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43075?download=true), [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Marden, J.H. 2008. [Evolution and physiology of flight in aquatic insects](https://doi.org/10.1079/9781845933968.0230). In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. [Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran](https://doi.org/10.1111/j.1558-5646.2012.01743.x). Evolution 67, 274–280.
    • Marden, J.H. 2013. Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran. Evolution 67, 274–280.
    • - movie: Skim onto mud, Gill flap homology
    • - movie: [Skim onto mud](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43077?download=true), [Gill flap homology](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43076?download=true)
    • Marden, J.H. 2013. Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.” Evolution DOI: 10.1111/evo.12093.
    • Marden, J.H. 2013. [Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.”](https://doi.org/10.1111/evo.12093) Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. Origin and diversification of wings: insights from a neopteran insect. Proceedings of the National Academy of Science 112(52), 15946–15951.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. [Origin and diversification of wings: insights from a neopteran insect](https://doi.org/10.1073/pnas.1509517112). Proceedings of the National Academy of Science 112(52), 15946–15951.
  • Updated Description Show Changes

    June 24, 2025 11:02 by avs5190

    Description
    • Understanding the evolution of insect wings and flight is a long-standing challenge for biology. Darwin's concept of evolution by natural selection was immediately challenged by critics asking "What good is half an eye or a nub of a wing?". In other words, how do complex traits evolve in a gradualistic fashion if there are no apparent scenarios for incremental improvement?
    • Starting in 1994, the Marden lab in the Dept. of Biology at Penn State University discovered and began publishing what would become a series of papers on flight-like behaviors of adult forms of aquatic insects that use their wings to skim across the surface of water. A key feature of these behaviors is that they use wings to accomplish 2-dimensional movement across water surfaces without requiring aerodynamic forces sufficient to support body weight. The first paper in 1994 showed that clipping the wings to mere nubs, and/or reducing body temperature to near freezing (a proxy for reducing the force and power output of flight muscles) still allowed surface skimming, thereby providing for the first time an empirical demonstration of a setting in which protowings and flight muscles could have been used for a flight-like behavior prior to the evolution of powered 3-dimensional flight.
    • These papers were accompanied by URLs where readers could find videos of these behaviors, but over time those URLs became no longer functional. This archive serves to correct that loss of video material for this collection of work. Below are the papers describing behaviors and the accompanying videos, all of which show various species of stoneflies except where otherwise indicated by the video title or text in the video itself.
    • Papers:
    • Marden, J.H. and M.G. Kramer. 1994. [Surface-skimming stoneflies: a possible intermediate stage for insect flight evolution](https://doi.org/10.1126/science.266.5184.427). Science 266, 427-430.
    • - See also: Kaiser, J. 1994. [A new theory of insect wing origins takes off](https://doi.org/10.1126/science.266.5184.363). Science 266, 363.
    • - movie: [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true)
    • - Television: [Scientific American Frontiers](https://www.youtube.com/watch?v=2hQ0MK-cTSU&list=PLeTVzyt4IP4vx8iu7toZWVbTLWWRBNWKu&index=1), see 34:25 to 39:48
    • Marden, J.H. 1995. Flying lessons from a flightless insect. Natural History 104, 4-8.
    • Marden, J.H. and M.G. Kramer. 1995. [Locomotor performance of insects with rudimentary wings](https://doi.org/10.1038/377332a0). Nature 377, 332-334.
    • - movie: [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true)
    • Marden, J.H. 1995. [How insects learned to fly](https://ezaccess.libraries.psu.edu/login?url=https://www.proquest.com/scholarly-journals/how-insects-learned-fly/docview/212702879/se-2?accountid=13158). The Sciences 35, 26-30.
    • Marden, J.H. and M.G. Kramer. 1995. [Plecopteran surface-skimming and insect flight evolution - reply](https://doi.org/10.1126/science.270.5242.1685). Science 270, 1685.
    • Kramer, M.G. and J.H. Marden. 1997. [Almost airborne](https://doi.org/10.1038/385403a0). Nature 385, 403-404.
    • - movie: [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true)
    • Marden, J.H., B.C. O’Donnell, M.A. Thomas, and J.Y. Bye. 2000. [Surface-skimming stoneflies and mayflies: the taxonomic and mechanical diversity of two-dimensional aerodynamic locomotion](https://doi.org/10.1086/318109). Physiological and Biochemical Zoology 73, 751-764.
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Thomas, M.A., K.A. Walsh, M.R. Wolf, B.A. McPheron, and J.H. Marden. 2000. [Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior](https://doi.org/10.1073/pnas.230296997). Proceedings of the National Academy of Sciences 97:13178-13183.
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Marden, J.H. and M.A. Thomas. 2003. [Rowing locomotion by a stonefly that possesses the ancestral pterygote condition of co-occurring wings and abdominal gills](https://doi.org/10.1046/j.1095-8312.2003.00192.x). Biological Journal of the Linnean Society 79: 341–349.
    • - movie: [Rowing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43075?download=true)
    • Marden, J.H. 2003. The surface-skimming hypothesis for the evolution of insect flight. Proceedings of the 2nd International Congress of Paleoentomology. Acta Zoologica Cracoviensia. 46: 73-84.
    • - movies: [Gill flapping](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43070?download=true), [Rowing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43075?download=true), [Sailing](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43068?download=true), [Swim-skim](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43071?download=true), [Six-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43067?download=true), [Four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43072?download=true), [Mayfly four-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43073?download=true), [Hind-leg skimming](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43069?download=true), [Jump](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43074?download=true)
    • Marden, J.H. 2008. [Evolution and physiology of flight in aquatic insects](https://doi.org/10.1079/9781845933968.0230). In: Aquatic Insects: Challenges to Populations, ed. J. Lancaster. CABI Press.
    • Marden, J.H. 2013. [Reanalysis and experimental evidence indicate that the earliest trace fossil of a winged insect was a surface-skimming neopteran](https://doi.org/10.1111/j.1558-5646.2012.01743.x). Evolution 67, 274–280.
    • - movie: [Skim onto mud](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43077?download=true), [Gill flap homology](https://scholarsphere.psu.edu/resources/d37c8365-019c-4914-bfca-fefce05aa7e3/downloads/43076?download=true)
    • Marden, J.H. 2013. [Reply to “Comment on Marden (2013) regarding the interpretation of the earliest trace fossil of a winged insect.”](https://doi.org/10.1111/evo.12093) Evolution DOI: 10.1111/evo.12093.
    • Medved V, Marden JH, Fescemyer HW, Der J, Mahfooz N, Popadić, A. 2015. [Origin and diversification of wings: insights from a neopteran insect](https://doi.org/10.1073/pnas.1509517112). Proceedings of the National Academy of Science 112(52), 15946–15951.
  • Updated Keyword Show Changes

    June 24, 2025 13:31 by avs5190

    Keyword
    • Evolution, Flight, Insects, Protowings, Wings, Aquatic, Skimming, Surface-skimming, Complexity, Pterygota, Pterygotes, Fossils, Entomology, Stoneflies, Plecoptera, Mayflies, Ephemeroptera, Gills, Wings from gills