page_title: Origin of birds
section_title: Cursorial ("from the ground up") theory
hierarchical_section_title: Origin of birds / Origin of bird flight / Cursorial ("from the ground up") theory
caption_attribution_description: STAGES IN THE DEVELOPMENT OF FLIGHT Tetrapteryx, Archaeopteryx, Hypothetical Stage, Modern Bird. After Beebe.
context_page_description: The scientific question of within which larger group of animals birds evolved has traditionally been called the 'origin of birds'. The present scientific consensus is that birds are a group of maniraptoran theropod dinosaurs that originated during the Mesozoic Era.
  A close relationship between birds and dinosaurs was first proposed in the nineteenth century after the discovery of the primitive bird Archaeopteryx in Germany. Birds and extinct non-avian dinosaurs share many unique skeletal traits. Moreover, fossils of more than thirty species of non-avian dinosaur with preserved feathers have been collected. There are even very small dinosaurs, such as Microraptor and Anchiornis, which have long, vaned arm and leg feathers forming wings. The Jurassic basal avialan Pedopenna also shows these long foot feathers. Paleontologist Lawrence Witmer concluded in 2009 that this evidence is sufficient to demonstrate that avian evolution went through a four-winged stage. Fossil evidence also demonstrates that birds and dinosaurs shared features such as hollow, pneumatized bones, gastroliths in the digestive system, nest-building and brooding behaviors.
context_section_description: The cursorial theory of the origin of flight was first proposed by Samuel Wendell Williston, and elaborated upon by Baron Nopcsa. This hypothesis proposes that some fast-running animals with long tails used their arms to keep their balance while running. Modern versions of this theory differ in many details from the Williston-Nopcsa version, mainly as a result of discoveries since Nopcsa's time.
  Nopcsa theorized that increasing the surface area of the outstretched arms could have helped small cursorial predators keep their balance, and that the scales of the forearms elongated, evolving into feathers. The feathers could also have been used to trap insects or other prey. Progressively, the animals leapt for longer distances, helped by their evolving wings. Nopcsa also proposed three stages in the evolution of flight. First, animals developed passive flight, in which developing wing structures served as a sort of parachute. Second, they achieved active flight by flapping the wings. He used Archaeopteryx as an example of this second stage. Finally, birds gained the ability to soar.
  Current thought is that feathers did not evolve from scales, as feathers are made of different proteins. More seriously, Nopcsa's theory assumes that feathers evolved as part of the evolution of flight, and recent discoveries prove that assumption is false.
  Feathers are very common in coelurosaurian dinosaurs (including the early tyrannosauroid Dilong). Modern birds are classified as coelurosaurs by nearly all palaeontologists, though not by a few ornithologists. The modern version of the "from the ground up" hypothesis argues that birds' ancestors were small, feathered, ground-running predatory dinosaurs (rather like roadrunners in their hunting style) that used their forelimbs for balance while pursuing prey, and that the forelimbs and feathers later evolved in ways that provided gliding and then powered flight. The most widely suggested original functions of feathers include thermal insulation and competitive displays, as in modern birds.
  All of the Archaeopteryx fossils come from marine sediments, and it has been suggested that wings may have helped the birds run over water in the manner of the Jesus Christ Lizard (common basilisk).
  Most recent refutations of the "from the ground up" hypothesis attempt to refute the modern version's assumption that birds are modified coelurosaurian dinosaurs. The strongest attacks are based on embryological analyses that conclude that birds' wings are formed from digits 2, 3, and 4, (corresponding to the index, middle, and ring fingers in humans. The first of a bird's three digits forms the alula, which they use to avoid stalling in low-speed flight—for example, when landing). The hands of coelurosaurs, however, are formed by digits 1, 2, and 3 (thumb and first two fingers in humans). However, these embryological analyses were immediately challenged on the embryological grounds that the "hand" often develops differently in clades that have lost some digits in the course of their evolution, and that birds' "hands" do develop from digits 1, 2, and 3. This debate is complex and not yet resolved - see "Digit homology".
caption: Proposed development of flight in a book from 1922: Tetrapteryx, Archaeopteryx, Hypothetical Stage, Modern Bird
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