Surface-Skimming Stoneflies: A Possible Intermediate Stage in Insect Flight Evolution

@article{Marden1994SurfaceSkimmingSA,
  title={Surface-Skimming Stoneflies: A Possible Intermediate Stage in Insect Flight Evolution},
  author={James H. Marden and Melissa G. Kramer},
  journal={Science},
  year={1994},
  volume={266},
  pages={427 - 430}
}
Insect wings appear to have evolved from gills used by aquatic forms for ventilation and swimming, yet the nature of intermediate stages remains a mystery. Here a form of nonflying aerodynamic locomotion used by aquatic insects is described, called surface skimming, in which thrust is provided by wing flapping while continuous contact with the water removes the need for total aerodynamic weight support. Stoneflies surface skim with wing areas and muscle power output severely reduced, which… 
Surface‐Skimming Stoneflies and Mayflies: The Taxonomic and Mechanical Diversity of Two‐Dimensional Aerodynamic Locomotion*
TLDR
The phylogenetic distribution and mechanistic diversity of surface skimming in stoneflies and mayflies supports the hypothesis that the common ancestor of stoneflies was a surface skimmer, and a synthetic model for the evolution of flying insects from surface skimmers is formed.
Molecular phylogenetic analysis of evolutionary trends in stonefly wing structure and locomotor behavior.
TLDR
The results show that basal stoneflies are surface skimmers, and that various forms of surface skimming are distributed widely across the plecopteran phylogeny, which supports the hypothesis that the first stoneflies were surface skimmer and that wing structures important for aerial flight have become elaborated and more diverse during the radiation of modern stoneflies.
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TLDR
It is shown that Allocapnia vivipara stoneflies use a non-flying form of aerodynamic locomotion which may exemplify a precursor to flight, and this support the hypothesis that insect wings evolved from articulated gill plates of aquatic ancestors through an intermediate semi-aquatic stage.
Flightlessness in mayflies and its relevance to hypotheses on the origin of insect flight
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Comparison of wing area of living mayflies with fossil species indicates that brachyptery could have already occurred in early flying insects (in the Permian) and it is argued that flight loss in Cheirogenesia has been made possible by the lack of fish predation in its natural habitats.
Paddling mode of forward flight in insects.
TLDR
By analyzing high-speed video of the fruit fly, a swimminglike mode of forward flight characterized by paddling wing motions is discovered that is as effective in air as in water and represents a common strategy for propulsion through aquatic and aerial environments.
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TLDR
This work examines the locomotor behaviour and gill morphology of a stonefly, Diamphipnopsis samali (Plecoptera), which retains abdominal gills in the winged adult stage, and suggests an ability to contribute to gas-exchange in an amphibious setting during a transition from aquatic to aerial locomotion and gas exchange.
Evolution Of Flight In Animals
TLDR
The evolution of flight in animals has long been debated and different hypotheses have been suggested for their origins, but particular stress has been laid on whether a gliding stage was included or not.
Plecopteran Surface-Skimming and Insect Flight Evolution
TLDR
Phylogenetic and fossil evidence support multiple origins of flight within insects, showing improved skimming performance an origin of flight in a common ancestor of eral times within Plecoptera alone, and Marden and Kramer suggest that Plecopmining homologies are speculative.
Honeybees use their wings for water surface locomotion
  • C. Roh, M. Gharib
  • Engineering
    Proceedings of the National Academy of Sciences
  • 2019
TLDR
The ability to self-propel on a water surface may increase the water-foraging honeybee’s survival chances when they fall on the water, and the findings may have biological implications on the survival of water foragers and preflight locomotion mechanisms.
Waxing and Waning of Wings.
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References

SHOWING 1-10 OF 23 REFERENCES
AERODYNAMICS, THERMOREGULATION, AND THE EVOLUTION OF INSECT WINGS: DIFFERENTIAL SCALING AND EVOLUTIONARY CHANGE
TLDR
Using physical models of Paleozoic insects in a wind tunnel, the potential effects of wings for increasing gliding distance, increasing dispersal distance during parachuting, improving attitude control or stability, and elevating body temperatures during thermoregulation are explored.
Bodybuilding Dragonflies: Costs and Benefits of Maximizing Flight Muscle
  • J. Marden
  • Environmental Science
    Physiological Zoology
  • 1989
TLDR
It is suggested that a high FMR enhances ability to compete in aerial contests, which enables greater short-term mating success and a trade-off made between flight ability and nutritional state that may affect longevity and long- term mating success is probably widespread among flying animals.
Origin of the insect wing and wing articulation from the arthropodan leg
The most primitive known pterygote terga, wing articulation, wings, and upper leg segments with exites, occur in gigantic Upper Carboniferous Paleodictyoptera, Homoiopteridae. Fossil features are
Origin and evolution of insect wings and their relation to metamorphosis, as documented by the fossil record
TLDR
All primitive Paleozoic pterygote nymphs are now known to have had articulated, freely movable wings reinforced by tubular veins, which suggests that the wings of early Pterygota were engaged in flapping movements, and that the immobilized, fixed, veinless wing pads of Recentnymphs have resulted from a later adaptation affecting only juveniles.
Thermoregulatory significance of thoracic lobes in the evolution of insect wings.
The evolution of broadly attached thoracic lobes could have increased the body temperature excess of ancient wingless insects by 55 percent over that of lobeless forms. The subsequent expansion of
Maximum Lift Production During Takeoff in Flying Animals
TLDR
Interspecific differences in short-duration powered flight and takeoff ability are shown to be caused primarily by differences in flight muscle ratio, which ranges from 0.115 to 0.560 among species studied to date.
Flight of Winter Moths Near 0�C
TLDR
The geometrids that are able to fly with a thoracic temperature near 0�C do so largely because of unusually low wing-loading, which permits a low energetic cost of flight.
Selective Factors in the Evolution of Insect Wings
TLDR
Flight represents a major innovation that has evolved independently in several different animal groups and how such key innovations can evolve has been one of the major questions in evolutionary biology since the time of Darwin.
Aerial Predation and Butterfly Design: How Palatability, Mimicry, and the Need for Evasive Flight Constrain Mass Allocation
TLDR
The need for evasive flight imposes constraints on body design that have potentially far-reaching effects on butterfly biology, by comparing design of butterflies and diurnal moths that vary in their need to evade aerial predators.
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