| Scansoriopteryx | |
|---|---|
 | |
| Skeletal restoration of the type specimen | |
Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Chordata |
| Clade: | Dinosauria |
| Clade: | Saurischia |
| Clade: | Theropoda |
| Family: | †Scansoriopterygidae |
| Genus: | †Scansoriopteryx Czerkas & Yuan,2002 |
| Species: | †S. heilmanni |
| Binomial name | |
| †Scansoriopteryx heilmanni Czerkas & Yuan, 2002 | |
| Synonyms | |
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Scansoriopteryx ("climbing wing") is agenus ofmaniraptorandinosaur. Described from only a single juvenilefossil specimen found inLiaoning,China,Scansoriopteryx is asparrow-sized animal that shows adaptations in the foot indicating an arboreal (tree-dwelling) lifestyle. It possessed an unusual, elongated third finger which may have supported a membranous wing, much like the relatedYi qi. The type specimen ofScansoriopteryx also contains the fossilized impression of feathers.[1]
Most researchers regard this genus as a synonym ofEpidendrosaurus, with some preferring to treatScansoriopteryx as the junior synonym,[2][3] though it was the first name to be validly published.[4]
The provenance of theScansoriopteryx type specimen is uncertain, as it was obtained from private fossil dealers who did not record exact geologic data. Czerkas and Yuan initially reported that it had likely come from theYixian Formation, though Wanget al. (2006), in their study of the age of theDaohugou Beds, suggested that it probably hails from the same beds, and thus is likely a synonym ofEpidendrosaurus. The Daohugou Beds supposedly date to the mid-lateJurassic Period,[5] but this is hotly contested. See theDaohugou Beds article for details.
Thetype specimen ofScansoriopteryx (type genus of the Scansoriopterygidae) and its arboreal adaptations were first presented in 2000 during the Florida Symposium on Dinosaur/Bird Evolution, at the Graves Museum of Archaeology & Natural History, though the specimen would not be formally described and named until 2002.[1]
Thetype specimen ofScansoriopteryx heilmanni (specimen number CAGS02-IG-gausa-1/DM 607) represents thefossilized remains of a hatchlingmaniraptoran dinosaur, similar in some ways toArchaeopteryx. A second specimen, theholotype ofEpidendrosaurus 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.[6] Because the only known specimens are juvenile, the size of a full-grownScansoriopteryx is unknown; the type specimen is a tiny,sparrow-sized creature.[6]
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 theoviraptorosaurs.
One distinctive feature ofScansoriopteryx is its elongated third finger, which is the longest on the hand, nearly twice as long as the second finger. This is unlike the configuration seen in most othertheropods, 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.[7] A relative ofScansoriopteryx,Yi, suggests that this elongated third finger supported a membranous wing of some kind alongside feathers.[8]
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.[7] 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" ofMicroraptor and other basal paravians.[7] It also had an unusually large first toe, orhallux, 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 thefemur, and ended in a fan of feathers.[6]
In describingScansoriopteryx, Czerkas & Yuan cited evidence for an arboreal (tree-dwelling) lifestyle. They noted that, unlike all modern bird hatchlings, the forelimbs ofScansoriopteryx are longer than the hind limbs. The authors argued that this anomaly indicates the forelimbs played an important role in locomotion even at an extremely early developmental stage.Scansoriopteryx has a well-preserved foot, and the authors interpreted the hallux as reversed, the condition of a backward-pointing toe being widespread among modern tree-dwelling birds. Furthermore, the authors pointed to the short, stiffened tail of theScansoriopteryx specimen as a tree-climbing adaptation. The tail may have been used as a prop, much like the tails of modernwoodpeckers. Comparison with the hands of modern climbing species with elongated third digits, likeiguanid lizards, also supports the tree-climbing hypothesis. Indeed, the hands ofScansoriopteryx are much better adapted to climbing than the modern tree-climbing hatchling of thehoatzin.[1]
TheEpidendrosaurus was also interpreted as arboreal based on the elongated hand and specializations in the foot.[6] The describing authors stated that the long hand and strongly curved claws are adaptations for climbing and moving around among tree branches. They viewed this as an early stage in the evolution of the bird wing, stating that the forelimbs became well-developed for climbing, and that this development later lead to the evolution of a wing capable of flight. They stated that long, grasping hands are more suited to climbing than to flight, since most flying birds have relatively short hands.
Zhanget al. also noted that the foot ofEpidendrosaurus is unique among non-avian theropods. While theEpidendrosaurus specimen does not preserve a reversed hallux, the backward-facing toe seen in modern perching birds, its foot was very similar in construction to more primitive perching birds likeCathayornis andLongipteryx. These adaptations for grasping ability in all four limbs makes it likely thatEpidendrosaurus spent a significant amount of time living in trees.
Scansoriopteryx fossils preserve impressions of wispy, down-likefeathers around select parts of the body, forming V-shaped patterns similar to those seen in modern down feathers. The most prominent feather impressions trail from the left forearm and hand. The longer feathers in this region led Czerkas and Yuan to speculate that adult scansoriopterygids may have had reasonably well-developed wing feathers which could have aided in leaping or rudimentary gliding, though they ruled out the possibility thatScansoriopteryx could have achieved powered flight. Like other maniraptorans,Scansoriopteryx had a semilunate (half-moon shaped) bone in the wrist that allowed for bird-like folding motion in the hand. Even if powered flight was not possible, this motion could have aided maneuverability in leaping from branch to branch.[1] Reticulatescales were preserved in the proximal portion of the second metatarsal,[7] and theEpidendrosaurus specimen also preserved faintfeather impressions at the end of the tail, similar to the pattern found inMicroraptor.[6]
The holotype skeleton ofEpidendrosaurus was recovered from the Daohugou fossil beds of northeastern China. In the past, there has been some uncertainty regarding the age of these beds. Various papers have placed the fossils here anywhere from theMiddle Jurassic period (169 million years ago) to theEarly Cretaceous period (122 ma).[9] The age of this formation has implications for the relationship betweenEpidendrosaurus and similar dinosaurs, as well as for the origin of birds in general. A Middle Jurassic age would mean that the bird-like dinosaurs in the Daohugou beds are older than the "first bird",Archaeopteryx, which wasLate Jurassic in age. The provenance ofScansoriopteryx is uncertain, though Wanget al. (2006), in their study of the age of the Daohugou, suggest that it probably hails from the same beds, and thus is likely a synonym ofEpidendrosaurus.
Scansoriopteryx lent its name to the familyScansoriopterygidae. Studies of dinosaur relationships have foundScansoriopteryx to be a close relative of truebirds and a member of thecladeAvialae.[10]
The status of the nameScansoriopteryx has been controversial. The type specimen was described only a few months after a very similar specimen,Epidendrosaurus ninchengensis, was described online, though the nameEpidendrosaurus was not published in print until afterScansoriopteryx.[1] These two specimens are so similar that they may be the same genus, in which case Article 21 of theInternational Code of Zoological Nomenclature (ICZN) would give priority toScansoriopteryx. The journal in whichScansoriopteryx appeared has a very small circulation, but was distributed on roughly 2002-09-02, before the print appearance ofEpidendrosaurus, but well after the later's appearance on the Internet, enough time for the nameEpidendrosaurus 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 nameEpidendrosarus 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 genusScansoriopteryx has been treated as a senior synonym ofEpidendrosaurus by some scientists, such as Alan Feduccia,[3] and as a junior synonym by others such asThomas R. Holtz, Jr.[11] and Kevin Padian.[2]
Czerkas and Yuan used the suite of primitive and birdlike characters inScansoriopteryx to argue for an unorthodox interpretation of dinosaur evolution. They stated thatScansoriopteryx was "clearly more primitive thanArchaeopteryx", based on its primitive, "saurischian-style" pubis and robust ischia.Scansoriopteryx also lacks a fullyperforated 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 makeScansoriopteryx among the most birdlikeand the most primitive known dinosaurs. Czerkas and Yuan called it a "proto-maniraptoran", supporting the hypothesis ofGregory S. Paul that the larger, ground-dwelling maniraptorans likeVelociraptor 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, thatall 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 withAlan Feduccia, published a paper further describingScansoriopteryx 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 thatScansoriopteryx and all birds evolved from non-dinosaurianavemetatarsalian archosaurs likeScleromochlus.[7] This conclusion has been overwhelmingly rejected by other paleontologists due to the clear morphological similarities scansoriopterygids have to other maniraptoran dinosaurs.[12]
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