Proavis
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Proavis refers to a hypothetical extinct species or hypothetical extinct
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
and was coined in the early 20th century in an attempt to support and explain the hypothetical evolutionary steps and anatomical adaptations leading from non-avian
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is the ...
to
birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
. The term has also been used by defenders of the thecodontian origin of birds.


History

The term "Proavis" was first coined, although under the form "Pro-Aves", by
English English usually refers to: * English language * English people English may also refer to: Peoples, culture, and language * ''English'', an adjective for something of, from, or related to England ** English national ...
osteologist Osteology () is the scientific study of bones, practised by osteologists. A subdiscipline of anatomy, anthropology, and paleontology, osteology is the detailed study of the structure of bones, skeletal elements, teeth, microbone morphology, funct ...
and
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
William Plane Pycraft in "The Origin of Birds", a 1906 article published in the magazine ''Knowledge and Scientific News''. Pycraft added to his article his own drawn depiction of the hypothetical animal, a restoration entitled "One of the Pro-Aves". Pycraft's "Pro-Avis" (singular of "Pro-Aves") was arboreal, as suggested Professor Osborn six years before, in 1900, in an article dealing with the hypothetical common ancestors of dinosaurs and birds. Pycraft assumed that birds had developed as tree-dwelling dinosaurs, gliding on membranes between the limbs and the trunk. These membranes would gradually have been covered by increasingly more elongated scales, which eventually would have evolved into feathers. A year after Pycraft's article, the Hungarian
aristocrat The aristocracy is historically associated with "hereditary" or "ruling" social class. In many states, the aristocracy included the upper class of people (aristocrats) with hereditary rank and titles. In some, such as ancient Greece, ancient R ...
and
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Franz Nopcsa Franz may refer to: People * Franz (given name) * Franz (surname) Places * Franz (crater), a lunar crater * Franz, Ontario, a railway junction and unorganized town in Canada * Franz Lake, in the state of Washington, United States – see Fran ...
, who, while residing at London, had seen Pycraft's restoration, drew his own vision of the animal, a picture and article published in a 1907 issue of the ''
Proceedings of the Zoological Society of London The ''Journal of Zoology'' is a scientific journal concerning zoology, the study of animals. It was founded in 1830 by the Zoological Society of London and is published by Wiley-Blackwell. It carries original research papers, which are targeted t ...
'' (an English science publication nowadays known as ''Journal of Zoology''). But this time, for his own "Pro-Avis" restoration, Nopcsa suggested a
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often u ...
origin, not arboreal. The "Pro-Avis" would thus have been a running animal, accelerating and prolonging its jumps by flapping with feathered forelimbs. There is a model of Nopcsa's restoration, still preserved, and recently restored, in the
Grant Museum of Zoology The Grant Museum of Zoology and Comparative Anatomy is a natural history museum that is part of University College London in London, England. It was established by Robert Edmond Grant in 1828 as a teaching collection of zoological specimens and ...
, the only remaining university zoological museum in London. When Pycraft published his book ''A History of Birds'', in 1910, he reinserted his 1906 restoration ''One of the Pro-Aves'' (in page 39). In the following years, as of 1913, Danish artist and amateur
zoologist Zoology ()The pronunciation of zoology as is usually regarded as nonstandard, though it is not uncommon. is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and d ...
Gerhard Heilmann Gerhard Heilmann (later sometimes spelt "Heilman") (25 June 1859 – 26 March 1946) was a Danish artist and paleontologist who created artistic depictions of ''Archaeopteryx'', '' Proavis'' and other early bird relatives apart from writing the 19 ...
also used and popularised the term ''Proavis'', this time presenting the public with more accurate and anatomically probable drawings. Heilmann favoured a scientific model in which the assumed ''Proavis'' was arboreal and of thecodontian descent, thus not a dinosaur. In 1961,
Czech Czech may refer to: * Anything from or related to the Czech Republic, a country in Europe ** Czech language ** Czechs, the people of the area ** Czech culture ** Czech cuisine * One of three mythical brothers, Lech, Czech, and Rus' Places * Czech, ...
paleontologist Josef Augusta included "Proavis" in his book ''Prehistoric Birds and Reptiles''. The animal illustration, as were all illustrations in the book, was painted by
Zdeněk Burian Zdeněk Michael František Burian (11 February 1905 in Kopřivnice, Moravia, Austria-Hungary – 1 July 1981 in Prague, Czechoslovakia) was a Czech painter, book illustrator and palaeoartist whose work played a central role in the development o ...
. During the 1970s the work of John Ostrom, breathing new life into the hypothesis that birds were dinosaurs, revived the interest in the direct ancestors of birds. Ostrom assumed these were cursorial. An illustration of a running dinosaur explicitly indicated as a "Proavis" was in 1976 published by
Dieter Stefan Peters {{Short pages monitor