Scansoriopteryx

Scansoriopteryx ("climbing wing") is a genus of avialan dinosaur. Described from only a single juvenile fossil specimen found in Liaoning, China, Scansoriopteryx is a sparrow-sized animal that shows adaptations in the foot indicating an arboreal (tree-dwelling) lifestyle. It possessed an unusual, elongated third finger. The type specimen of Scansoriopteryx also contains the fossilized impression of feathers.[1]

Most researchers regard this genus as a synonym of Epidendrosaurus, with some preferring to treat Scansoriopteryx as the junior synonym,[2][3] though it was the first name to be validly published.

Description
The type specimen of Scansoriopteryx heilmanni (specimen number CAGS02-IG-gausa-1/DM 607) represents the fossilized remains of a hatchling maniraptoran dinosaur, similar in some ways to Archaeopteryx. A second specimen, the holotype of Epidendrosaurus ninchengensis (IVPP V12653), also shows features indicating it was a juvenile. The specimen is partially disarticulated, and most bones are preserved as impressions in the rock slab, rather than three-dimensional structures.[5] Because the only known specimens are juvenile, the size of a full-grown Scansoriopteryx is unknown–the type specimen is a tiny, sparrow-sized creature.[5]

Scansoriopteryx is also notable for its wide, rounded jaws. The lower jaw contained at least twelve teeth, larger in the front of the jaws than in the back. The lower jaw bones may have been fused together, a feature otherwise known only in the oviraptorosaurs.

One distinctive feature of Scansoriopteryx is its elongated third finger, which is the longest on the hand, nearly twice as long as the second finger (in most theropod dinosaurs, the second finger is the longest). This is unlike the configuration seen in most other theropods, where the second finger is longest. The long wing feathers, or remiges, appear to attach to this long digit instead of the middle digit as in birds and other maniraptorans. Shorter feathers are preserved attached to the second finger.

Scansoriopteryx had a non-perforated hip socket, which is more open in most, but not all, other dinosaurs. It also had a pubis (hip bone) which pointed forward, a primitive trait among theropods, and unlike some maniraptorans more closely related to birds, where the pubis points downward or backward.[6] The legs were short, and preserve small pebbly scales along the upper foot (metatarsus), as well as possible impressions of long feathers in the same area, possibly similar to the "hind wings" of Microraptor and other basal paravians.[6] It also had an unusually large first toe, or hallux, which was low on the foot and may have been reversed, allowing some grasping ability.[1]

The tail was long, six or seven times the length of the femur, and ended in a fan of feathers.

Classification
Scansoriopteryx lent its name to the family Scansoriopterygidae. Studies of dinosaur relationships have found Scansoriopteryx to be a close relative of true birds and a member of the clade Avialae.[7]

The status of the name Scansoriopteryx has been controversial. The type specimen was described only a few months after a very similar specimen, Epidendrosaurus ninchengensis, was described online, though the name Epidendrosaurus was not published in print until after Scansoriopteryx.[1] These two specimens are so similar that they may be the same genus, in which case Article 21 of the International Code of Zoological Nomenclature (ICZN) would give priority to Scansoriopteryx. The journal in which Scansoriopteryx appeared has a very small circulation, but was distributed on roughly 2002-09-02, before the print appearance of Epidendrosaurus, but well after the later's appearance on the Internet, enough time for the name Epidendrosaurus to have come into wide use by experts. This situation was used as an example in a proposed amendment to the ICZN by Jerry Harris that would consider electronic articles with Digital Object Identifiers (DOIs) that are subsequently available in print to qualify as "publication" for naming purposes. Harris noted that while the name Epidendrosarus appeared first, Scansoriopteryx was the first to be published in print and is therefore the valid name, but the fact that the ICZN does not recognize online names as valid has led to confusion over which has priority.[4] In scientific literature, the genus Scansoriopteryx has been treated as a senior synonym of Epidendrosaurus by some scientists, such as Thomas R. Holtz, Jr.[8] and Alan Feduccia,[3] and as a junior synonym by other such as Kevin Padian.

Alternate interpretation
Czerkas and Yuan used the suite of primitive and birdlike characters in Scansoriopteryx to argue for an unorthodox interpretation of dinosaur evolution. They stated that Scansoriopteryx was "clearly more primitive than Archaeopteryx", based on its primitive, "saurischian-style" pubis and robust ischia. Scansoriopteryx also lacks a fully perforated acetabulum, the hole in the hip socket which is a key characteristic of Dinosauria and has traditionally been used to define the group. While the authors allowed that the hole may have closed secondarily, having evolved from a more traditional dinosaurian hip socket, they cited the other primitive features to argue that it is a true primitive trait, which would make Scansoriopteryx among the most birdlike and the most primitive known dinosaurs. Czerkas and Yuan called it a "proto-maniraptoran", supporting the hypothesis of Gregory S. Paul that the lager, ground-dwelling maniraptorans like Velociraptor evolved from small, flying or gliding forms that lived in trees. The authors took this idea further than Paul, however, and lent support to George Olshevsky's 1992 "birds came first" hypothesis, that all true theropods are secondarily flightless or at least secondarily arboreal, having evolved from small, tree-dwelling, Scansoriopteryx-like ancestors. Czerkas and Yuan also argued that, contrary to most phylogenetic trees, maniraptorans form a separate lineage from other theropods, and that this split occurred very early in theropod evolution.[1]

In 2014, Czerkas, along with Alan Feduccia, published a paper further describing Scansoriopteryx and stating their opinion that certain archaic features of the skeleton and the hypothesis that it was arboreal ruled out the possibility that it was a theropod or even a dinosaur, but that Scansoriopteryx and all birds evolved from non-dinosaurian avemetatarsalian archosaurs like Scleromochlus.[6] Phylogenetic analyses have never supported this alternative interpretation, and the arguments of Czerkas and Fedducia have been criticized as unscientific.