Since Darwin's time, complex features
such as the vertebrate eye and the ability to fly have posed a
particular challenge for evolutionary biology. How can complex traits
evolve in a stepwise fashion when it is unclear what function or
selective advantage can be derived from "half an eye" or a "nub of a
wing"? In the case of insect flight, the challenge is to explain how a
flight-capable creature, with its complex interrelated nerves, muscles,
articulations, and wings, evolved from nonflying ancestors. We have
discovered that certain stoneflies use their wings to skim across water
surfaces (i.e. nonflying two-dimensional aerodynamic locomotion). These
surface-skimming stoneflies do not need to generate total weight
support, and they demonstrate a pathway for gradual evolution beginning
with rudimentary stages of wings and muscles, all the way to fully
weight-supported "true" flight. My students and I have traveled the
world (Australia, Chile, Europe) to survey the behavioral and
mechanical diversity of stonefly locomotion, and we have constructed a
molecular phylogeny to examine the evolutionary history of
these traits across the entire order of Plecoptera. As a result of
studies, we now have a reasonably well supported synthetic theory for
flight originated in insects.
Marden, J.H. and M.G. Kramer. 1994.
Surface-skimming stoneflies: a possible intermediate stage in insect
flight evolution. Science 266, 427-430.
Marden, J.H. 1995. Flying lessons from a flightless insect. Natural
History 104 (2), 4-8
Marden, J.H. and M.G. Kramer. 1995. Locomotor performance of insects
with rudimentary wings: sailing on water versus gliding in air. Nature
Marden, J.H. 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 Science 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:
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.
Hagner-Holler, S., A. Schoen, W. Erker, J.H. Marden, R. Rupprecht, H. Decker, T. Burmester. 2004. A respiratory hemocyanin from an insect. Proceedings of the National Academy of Sciences 101: 871-874.
Hagner-Holler, S., Pick, C., Girgenrath,S., Marden, J.H., and Burmester, T. 2007. Diversity of stonefly hexamerins and implication for the evolution of insect storage proteins. Insect Biochem. Mol. Biol. 37:1064-74
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. 2012. Reanalysis and experimental evidence indicates
that the earliest trace fossil of a winged insect was a
surface-skimming neopteran. Evolution, in press. DOI: