Scansoriopteryx

Scansoriopteryx is an excellent example of a transitional on the lineage from pterosaur to primitive bird. It is a member of basal Paraves.

Feduccia and Czerkas article:

https://www.researchgate.net/publication/271658034_Jurassic_archosaur_is_a_non-dinosaurian_bird

ORIGINAL ARTICLE

Jurassic archosaur is a non-dinosaurian bird

Stephen A. Czerkas •Alan Feduccia

Received: 17 April 2014 / Revised: 4 June 2014 / Accepted: 13 June 2014 / Published online: 9 July 2014

ÓDt. Ornithologen-Gesellschaft e.V. 2014

Abstract Re-examination utilizing Keyence 3D digital

microscopy and low angled illumination of the fossil

Scansoriopteryx, a problematic sparrow-size pre-Archae-

opteryx specimen from the Jurassic Daohugou Biotas, pro-

vides new evidence which challenges the widely accepted

hypothesis that birds are derived from dinosaurs in which

avian flight originated from cursorial forms. Contrary to

previous interpretations in which S

https://www.sciencedaily.com/releases/2014/07/140709140203.htm

The re-examination of a sparrow-sized fossil [Scansoriopteryx] from China challenges the commonly held belief that birds evolved from ground-dwelling theropod dinosaurs that gained the ability to fly. The birdlike fossil is actually not a dinosaur, as previously thought, but much rather the remains of a tiny tree-climbing animal that could glide, say American researchers Stephen Czerkas of the Dinosaur Museum in Blanding, Utah, and Alan Feduccia of the University of North Carolina. The study appears in Springer’s Journal of Ornithology.

Feduccia added, “Instead of regarding birds as deriving from dinosaurs, Scansoriopteryx reinstates the validity of regarding them as a separate class uniquely avian and non-dinosaurian.”

ELONGATE OUTER FINGER

http://link.springer.com/article/10.1007/s10336-014-1098-9/fulltext.html (2014)

Jurassic archosaur is a non-dinosaurian bird

Stephen A. Czerkas, Alan Feduccia

Re-examination utilizing Keyence 3D digital microscopy and low angled illumination of the fossil Scansoriopteryx, a problematic sparrow-size pre-Archaeopteryx specimen from the Jurassic Daohugou Biotas, provides new evidence which challenges the widely accepted hypothesis that birds are derived from dinosaurs in which avian flight originated from cursorial forms. Contrary to previous interpretations in which Scansoriopteryx was considered to be a coelurosaurian theropod dinosaur, the absence of fundamental dinosaurian characteristics demonstrates that it was not derived from a dinosaurian ancestry and should not be considered as a theropod dinosaur. Furthermore, the combination in which highly plesiomorphic non-dinosaurian traits are retained along with highly derived features, yet only the beginnings of salient birdlike characteristics, indicates that the basal origins of Aves stemmed from outside the Dinosauria and further back to basal archosaurs. Impressions of primitive elongate feathers on the forelimbs and hindlimbs suggest that Scansoriopteryx represents a basal form of “tetrapteryx” in which incipient aerodynamics involving parachuting or gliding was possible. Along with unique adaptations for an arboreal lifestyle, Scansoriopteryx fulfills predictions from the early twentieth century that the ancestors of birds did not evolve from dinosaurs, and instead were derived from earlier arboreal archosaurs which originated flight according to the traditional trees-down scenario.

The most unusual feature is the extremely elongate outer finger, considered here to be digit IV as in Aves (Capek et al. 2013). It is the longest manual digit whereas the middle digit in theropods is the longest.

http://dinosaur-museum.org/featheredinosaurs/arboreal_maniraptoran.pdf

Scansoriopteryx heilmanni is the only known
saurischian, or theropod, which has the third digit of the manus elongated to nearly twice that of the second digit. 

http://www.labnews.co.uk/news/new-finding-challenges-belief-dinosaurs-evolved-from-birds

The investigations – published in Journal of Ornithology – found a combination of plesiomorphic or ancestral non-dinosaurian traits along with highly derived unambiguous birdlike features. The researchers specifically note the primitive elongated feathers on the fore- and hind limbs, suggesting Scansoriopteryx is an ancestral form of early birds that had mastered basic aerodynamic manoeuvres of parachuting or gliding from trees.
These findings fulfil a prediction first made in the 1900s that the ancestors of birds didn’t evolve from dinosaurs, but instead from earlier arboreal archosaurs which originated flight according to the tree-down scenario. These small tree-dwelling archosaurs had improved ability to fly, with feathers that enabled them to at least glide. This ‘tree-down’ view is in contrast with the ‘ground-up’ view many palaeontologists side with.
“Instead of regarding birds as deriving from dinosaurs, Scansoriopteryx reinstates the validity of regarding them as a separate class uniquely avian and non-dinosaurian,” said Alan Feduccia.

Pterosaur fingers: 
http://www.reptileevolution.com/pterosaur-wings.htm

CAUDAL RODS

 http://www.sci-news.com/paleontology/science-scansoriopteryx-hypothesis-birds-evolved-dinosaurs-02059.html

The techniques made it possible to interpret the natural contours of the bones. Many aspects of the [Scansoriopteryx] fossil’s pelvis, forelimbs, hind limbs, and tail were confirmed, while it was discovered that it had elongated tendons along its tail vertebrae similar to Velociraptor.

http://onlinelibrary.wiley.com/doi/10.1111/1755-6724.12009/abstract

In the tails of dromaeosaurids dinosaurs and rhamphorhynchid pterosaurs, elongate osteological rods extend anteriorly from the chevrons and the prezygapophyses. These caudal rods are positioned in parallel and are stacked dorsoventrally.

http://pterosaur-net.blogspot.ca/2013/01/guest-post-dragon-tails-what-pterosaurs.html

The tail of Deinonychus and its raptor relatives is bizarre, but it is not (as Professor Ostrom himself realized) unique. Among all known vertebrates, a similar tail anatomy has evolved in one other group [rhamphorhynchid pterosaurs].

Consider the below images of a tail of a Bambiraptor and of a Velociraptor. Both are dromaeosaurids with caudal-rod bearing tails and both are fully articulated.

ACETABULUM

http://www.aoucospubs.org/doi/full/10.1525/auk.2013.130.1.1

A partially closed acetabulum is seen in basal archosaurs and
is characteristic of the scansoriopterids and Jurassic feathered
forms such as Anchiornis initially described as near Aves by Xu et al. (2009).

http://en.wikipedia.org/wiki/Scansoriopteryx

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. 

http://dinosaur-museum.org/featheredinosaurs/arboreal_maniraptoran.pdf

Scansoriopteryx is clearly more primitive
than Archaeopteryx in many respects such as its
saurischian-style pelvis which has remarkably short
pubes; elongate and robust ischia; and
comparatively small pubic peduncles. These
primitive features further suggest that the nearly
closed acetabulum is not a reversal, but a true
plesiomorphic condition.

http://en.wikipedia.org/wiki/Pterosaur

Pterosaur's hip sockets are oriented facing slightly upwards, and the head of the femur (thigh bone) is only moderately inward facing, suggesting that pterosaurs had a semi-erect stance. It would have been possible to lift the thigh into a horizontal position during flight as gliding lizards do.

GENERAL

http://en.wikipedia.org/wiki/Scansoriopterygidae

Scansoriopteryx (and its likely synonym Epidendrosaurus) was the first non-avian dinosaur found that had clear adaptations to an arboreal or semi-arboreal lifestyle–it is likely that they spent much of their time in trees. Both specimens showed features indicating they were juveniles, which made it difficult to determine their exact relationship to other non-avian dinosaurs and birds. It was not until the description of Epidexipteryx in 2008 that an adult specimen was known.

The scansoriopterygids would have lived alongside synapsids such as the aquatic Castorocauda and arboreal gliding mammal Volaticotherium, the rhamphorhynchoid pterosaurs Jeholopterus and Pterorhynchus, as well as a diverse range of insect life (including mayflies and beetles) and several species of salamander.[14][15]

A monophyletic Scansoriopterygidae was recovered by Godefroit et al. (2013); the authors found scansoriopterygids to be basalmost members of Paraves and the sister group to the clade containing Avialae and Deinonychosauria.[9] Agnolín and Novas (2013) recovered scansoriopterygids as non-paravian maniraptorans and the sister group to Oviraptorosauria.[10]

http://dinosaur-museum.org/featheredinosaurs/arboreal_maniraptoran.pdf

Scansoriopteryx heilmanni is the only known
saurischian, or theropod, which has the third digit
of the manus elongated to nearly twice that of the
second digit. Scansoriopteryx closely resembles
Archaeopteryx, but differs in the following: a
definite contact between an elongate ventral process
of the postorbital and the ascending process of the
jugal; the lower jaw is equipped with a large
fenestra; the tail has a greater development in the
articulation of the zygapophyses. The pelvis is
similar to that of Archaeopteryx in having the same
number of sacrals and general shape of the ilia, but
differs in having a small, unexpanded pubic
peduncle; a significantly short pubis which is not
retroverted; longer ischia; and an acetabulum which
is not entirely perforated.

Scansoriopteryx is clearly more primitive
than Archaeopteryx in many respects such as its
saurischian-style pelvis which has remarkably short
pubes; elongate and robust ischia; and
comparatively small pubic peduncles. These
primitive features further suggest that the nearly
closed acetabulum is not a reversal, but a true
plesiomorphic condition.

http://www.researchgate.net/publication/259438884_Agnoln_and_Novas._2013._Avian_ancestors

It is important to mention that scansoriopterygids retained a caudoventrally oriented glenoid, a subrectangular coracoid with reduced biceps tubercle, and a distally fan-shaped scapular blade, all representing plesiomorphic character states in respect to paravians.

Agnolin and Novas provide no support for their assertion that scansoriopterygids had a caudoventrally oriented glenoid.
http://www.researchgate.net/publicat...vian_ancestors
In Figure 5.2 they diagram other taxa but not scansoriopteryids. 

In the published material by those who have analyzed the fossils, I see no mention of a caudoventrally oriented glenoid. (If anyone does see a mention, please give us the link and copy and paste please). 
https://bio.unc.edu/files/2011/04/Jo...logy-20141.pdf
http://www.dinosaur-museum.org/feath...niraptoran.pdf

http://en.wikipedia.org/wiki/Scansoriopteryx

Zhang et al. also noted that the foot of Epidendrosaurus is unique among non-avian theropods. While the Epidendrosaurus 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 like Cathayornis and Longipteryx. These adaptations for grasping ability in all four limbs makes it likely that Epidendrosaurus spent a significant amount of time living in trees.

Riddle of the Feathered Dragons
Page 150

The [Scansoriopteryx] pelvis is still like that of a reptile (as opposed to a theropod) 

http://link.springer.com/article/10.1007%2Fs10336-014-1098-9#page-1

the head of [Scansoriopteryx] femur lacks a distinctive neck and is instead more proximally oriented as in reptiles with sprawling limbs

The reason that the dino to bird folk are confused about this, is that they see the "correction" made at the knee (to make the stance "erect") in basal paraves and birds, and equate this "erect" stance with the quite different "erect" stance of dinosaurs.


http://books.google.ca/books?id=Z0YWn5F9sWkC&pg=PA46&lpg=PA46&dq=parasagittal+gait&source=bl&ots=5IpEhI42X4&sig=zVn7UBHxK7EYwqz95O7uGIjrIuI&hl=en&sa=X&ei=vLxsVO3cIsOOiwK3mICYBw&ved=0CDwQ6AEwBQ#v=onepage&q=parasagittal%20gait&f=false

He also pointed out the distinction between a parasagittal gait and a sprawling gait, later understood in terms of fundamentally different mechanics.

Note the two different meanings:
http://en.wikipedia.org/wiki/Sagittal_plane

The term parasagittal is used to describe any plane parallel to the sagittal plane.

http://evolutionwiki.org/wiki/Parasagittal

Parasagittal refers to a gait in which the legs are oriented entirely under the body, like columns. Seen in mammals, and birds.

http://dml.cmnh.org/2008Sep/msg00563.html

Avialan characters of scansoriopterygids include- - Hyposphene -hypantrum articulations in trunk vertebrae absent (according to Senter).

Very interesting thoughts from Andrea Cau:

http://theropoda.blogspot.ca/2012/07/il-ritorno-del-paraviano-pterosauro.html

In an old post, I have argued, along with Lukas Panzarin, the hypothesis that the highly unorthodox Scansoriopterygidae are of theropodi "pterosaur-like" , converging with small pterosaurs grade rhamphorhynchoide, as they have a short skull, tall and with teeth prominent and projected rostrally, forelegs stretched, the outer finger of the hand hypertrophic but apparently unsuitable to prehension (the penultimate phalanx is not significantly longer than the other), the possible (even if you need a confirmation) of a track patagium nell'olotipo of Scansoriopteryx , attached to the third finger of the hand instead of showing the obvious feathering inserted the second finger as in other maniraptori. The recently described rhamphorhynchoide German Bellubrunnus (Hone et al. 2012) shows a tail very similar to that of Scansoriopteryx . Randomness? Or yet another evidence of convergent evolution? It should be emphasized that the tail of rhamphorhynchidi derivatives is very similar to that of dromaeosauridi derivative, while that of Bellubrunnus is more similar to that of the basal paraviani, such as scansoriopterygidi. Perhaps, the two lineages developed their tails like follow similar evolutionary trajectories, which Scansoriopteryx and Bellubrunnus are both intermediate stages. Although I agree that it is very risky to postulate such a narrow adaptive convergence between baseline and dromaeosauridi rhamphorhynchidi (but Microraptor is potentially arboreal and planatore , then ecologically similar, if not to a flying animal, at least in the same region adaptive in that would place the scansoriopterygidi planatori) I consider it possible that the still little-known faunas of the Middle Jurassic basal paraviani might reserve surprises in the future.
If the third finger of hypertrophic Scansoriopteryx is already so stretched in the sample holotype, which is very immature, why not imagine that in the adult finger that was even longer? And if so, what good would such an extended finger? Perhaps to support a patagium, especially if it was confirmed the absence of flight feathers in the wing of Epidexipteryx ?
I admit the many "ifs", but also the many enigmatic aspects of scansoriopterygidi, which probably require explanations heterodox: the assumptions are born so reckless ...

http://scienceblogs.com/tetrapodzoology/2008/10/23/epidexipteryx-at-last/

In my large-scale analysis of theropods (in preparation), Scansoriopterygidae are placed sligtly more basal than Avialae: they’re basal paravians, sister-group of Eumaniraptora (Avialae+Deinonychosauria). This different position explains more the “incisivosaur-like” skull and the absence of some pelvic features widespread among basal avialans, dromaeosaurids and troodontids (in particular the scapular, ischial and pubic features).
In my blog, I suggested an alternative and very heterodox interpretation of Scansoriopterygids: given the absence of evidence for remiges in Epidexipteryx, is it possible that the “feather impressions” seen in the forelimb of the “Scansoriopteryx heilmanni specimen” are not feathers, but a different tegument: it is interesting to note that in remige-bearing maniraptorans, the remiges are inserted on the second finger, whereas in Scasoriopteryx these impression are close to the hyper-elongated third finger. This very long lateral finger is similar to the pterosaurian fourth digit. So, it is possible that the “feather impressions” of Scansoriopteryx were remnant of a patagium. In my opinion, the presence of this structure may explain the elongation of the lateral digit in these small theropods better than the Aye-Aye hypothesis.

2015:
http://phenomena.nationalgeographic.com/2015/04/29/chinese-dinosaur-had-bat-like-wings-and-feathers/

These wings were mutually exclusive: dinosaur or pterosaur, feathery or leathery. But Yi went for both options! It had membrane wings with a feathery covering on the leading edge. It shows that at least some dinosaurs had independently evolved the same kind of wings as pterosaurs—an extraordinary example of convergent evolution.

http://blogs.scientificamerican.com/tetrapod-zoology/2015/05/05/yi-qi-is-neat/

Membranous-winged scansoriopterygids were predicted in 2008. Moving on, there are several neat things that I think should be discussed here (scarcely none of which have been mentioned in existing online discussions of this find). The first one is that the concept of a scansoriopterygid with gliding membranes is (while radical and shocking) not novel if you’ve been paying attention.Exactly such a creature was predicted by my colleague Andrea Cau (and illustrated by excellent palaeoartist Lukas Pankarin) way back in October 2008 after the publication of Epidexipteryx* (Zhang et al. 2008). Like Yi qi and other scansoriopterygids, Epidexipteryx lacks vaned feathers yet has a hyper-long digit that looks suitable – Andrea proposed – for the support of a patagial membrane. Yi qi is, therefore, yet another of those fossil animals predicted to exist prior to its discovery. SpecBio fans might like to know that C. M. Kosemen took this suggestion and ran with it, but that’s a story for another time.

http://dinosaur-museum.org/featheredinosaurs/Are_Birds_Really_Dinosaurs.pdf

Had Scansoriopteryx been discovered before the use of cladistics became so prevalent some two decades ago, the cursorial theory of how birds evolved from theropod dinosaurs could not have progressed as it has. Scansoriopteryx is known from strata believed to be from the Middle Jurassic, possibly 165 to 180 million years ago, and much older than when Archaeopteryx is known to have lived during the Late
Jurassic. So there is no time paradox in its being ancestral to birds, as there is for dromaeosaurs of other bird-like dinosaurs. Scansoriopteryx simply does not represent a ground dwelling dinosaur, but it clearly does have a significant ability for climbing while at the same time is not as well developed for flight as was Archaeopteryx. The shoulder/chest complex of the scapula and coracoid are more primitive than
that of Archaeopteryx. The furcula is not present, but is instead represented by separate clavicles.

http://dinosaur-museum.org/featheredinosaurs/arboreal_maniraptoran.pdf

The systematic description of Scansoriopteryx
depends upon whether certain characters are
considered as truly plesiomorphic, or as derived
reversals that only resemble primitive conditions
secondarily. The main distinction between the two
interpretations is that Scansoriopteryx was derived
either from a pre-theropod saurischian [archosaur] ancestor, or
from a theropod. The first scenario suggests that
the ancestral forms which led to Scansoriopteryx
were basal saurischians from the Middle Triassic,
or earlier, before theropods had appeared. The
second option would suggest that Scansoriopteryx
appeared much later in time from a theropod lineage
which, in becoming arboreal, developed massive
reversals secondarily resembling primitive
characteristics. The basal saurischian relationship
is seen here as being the more parsimonious
interpretation. 

The passage above is very good, but there is no reason to think the ancestor was a saurischian dinosaur. It was not any kind of dinosaur. It was actually a pterosaur.


http://www.nature.com/nature/journal/v498/n7454/full/nature12168.html

Three main references:

http://www.ivpp.ac.cn/qt/papers/201206/P020120604508520389814.pdf (2002)

A juvenile coelurosaurian theropod [Epidendrosaurus] from China indicates arboreal habits

http://dinosaur-museum.org/featheredinosaurs/arboreal_maniraptoran.pdf

http://link.springer.com/article/10.1007/s10336-014-1098-9/fulltext.html  (2014)


Epidexipteryx

http://en.wikipedia.org/wiki/Epidexipteryx

The skull of Epidexipteryx is also unique in a number of features, and bears an overall similarity to the skull of Sapeornis, oviraptorosaurs and, to a lesser extent, therizinosauroids. It had teeth only in the front of the jaws, with unusually long front teeth angled forward, a feature only seen in Masiakasaurus among other theropods. The rest of the skeleton bore an overall similarity to the possibly closely related Scansoriopteryx, including a hip configuration unusual among other dinosaurs: the pubis was shorter than the ischium, and the ischium itself was expanded towards the tip. The tail of Epidexipteryx also bore unusual vertebrae towards the tip which resembled the feather-anchoring pygostyle of modern birds and some oviraptorosaurs.[2]

Consensus

https://www.cfa.harvard.edu/~scranmer/SPD/crichton.html

"I want to pause here and talk about this notion of consensus, and the rise of what has been called consensus science. I regard consensus science as an extremely pernicious development that ought to be stopped cold in its tracks. Historically, the claim of consensus has been the first refuge of scoundrels; it is a way to avoid debate by claiming that the matter is already settled. Whenever you hear the consensus of scientists agrees on something or other, reach for your wallet, because you're being had.
Let's be clear: the work of science has nothing whatever to do with consensus. Consensus is the business of politics. Science, on the contrary, requires only one investigator who happens to be right, which means that he or she has results that are verifiable by reference to the real world. In science consensus is irrelevant. What is relevant is reproducible results. The greatest scientists in history are great precisely because they broke with the consensus.
There is no such thing as consensus science. If it's consensus, it isn't science. If it's science, it isn't consensus. Period.
In addition, let me remind you that the track record of the consensus is nothing to be proud of." 

A lecture by Michael Crichton Caltech Michelin Lecture January 17, 2003

Characters

Here is an accumulation of information related to characters analyzed in Nesbitt (2011) and Nesbitt et al (2009). These support the pterosaur to bird theory and contradict the dino to bird theory. This section is a work in progress.


NESBITT (2011) 

Nesbitt characters 212, 213, 214, 223, 230, 231 and 370.

212. Forelimb–hind limb, length ratio: (0) more than 0.55; (1) less than 0.55 (Gauthier,

1984; Sereno, 1991a; Juul, 1994; Benton, 1999).

Humerus + radius [forelimb] : Femur + tibia [hindlimb]

SEE REFERENCE IN DINOSAURIA (page 204):

Dromaeosaurid forelimbs are among the longest in theropods, the ratio of forelimb length to hindlimb length being about 65% in Velociraptor, 70% in Deinonychus and 80% in Sinornithosaurus.

It seems that Nesbitt scored the Velociraptor incorrectly as (1).
Note that Velocirptor and Deinonychus are terrestrial, secondarily flightless members of Paraves.

213. Clavicles: (0) present and unfused; (1) fused into a furcula (modified from Gauthier, 1986; Sereno, 1991a; Benton, 1999; Benton and Walker, 2002). Clavicles are present in non-archosaurian archosauriforms and basal crocodylian-line archosaurs. Clavicles are not present in crocodylomorphs (e.g., Hesperosuchus ‘‘agilis,’’ CM 29894; Protosuchus richardsoni, AMNH FR 3024) and, therefore, they are scored as inapplicable. Like the interclavicle, the clavicles of the pterosaur Eudimorphodon are separate ossifications in a small specimen and incorporated into the sternum (Wild, 1993). All other pterosaurs seem to lack distinct ossifications of the clavicles. Within Dinosauria, clavicles are present, but do not contact in some ornithischians (e.g., Psittacosaurus) and are unossified in others (Butler et al., 2008a). The clavicles of some nonsauropod sauropodomorphs (e.g., Massospondylus) may contact each other at the midline, but do not fuse (Yates and Vasconcelos, 2005). A furcula (fused clavicles) is present in nearly all theropods known from complete skeletons including Coelophysis bauri (AMNH FR 30647; Rinehart et al., 2007; Nesbitt et al., 2009d) and Allosaurus fragilis (UUVP 6102; Chure and Madsen, 1996). This character has been employed by various datasets exploring theropod relationships (e.g., Norell et al., 2001; Clarke, 2004).

Pterosaurs have clavicles and an interclavicle that are fused into the sternum.



214. Interclavicle: (0) present; (1) absent
(fig. 30) (Gauthier, 1986; Sereno, 1991a; Juul,
1994; Benton, 1999).
The interclavicle is present in archosauriforms
plesiomorphically (Sereno, 1991a) and
persists through Pseudosuchia. In Pterosauria,
an interclavicle appears to be present
in young individuals of Eudimorphodon
(MCSNB 8950), but fuse to the pectoral
elements in larger individuals (Wild, 1993). A
distinct interclavicle is not present in all other
pterosaurs. Ornithischians and saurischians
lack an interclavicle. However, the pectoral
girdles in the successive sister taxa to
Dinosauria (Silesaurus, Marasuchus, Lagerpeton)
do not have the pectoral region
completely preserved. As a result, the optimization
of this character within Dinosauromorpha
is not clear.

SEE REFERENCE IN DINOSAURIA (page 204)

Also from Nesbitt et al (2009) (page 872):

The evolutionary transformation of the furcula
from separate clavicles is nicely illustrated in
archosaurs and their close relatives. Euparkeria
capensis and early pseudosuchians retain both the
interclavicle and clavicles. The interclavicle is lost
at the dinosaur node or at an unknown node
among early dinosauromorphs. From there, the origin
of the furcula is well understood with the
addition of the work on basal sauropodomorphs of
Yates and Vasconcelos (2005) and the discovery of
furculae in early coelophysoid theropods (Tykoski
et al., 2002; Rinehart et al., 2007). Some ornithischians
have two small clavicles that do not
contact each other (Osborn, 1924a; Brown and
Schlaikjer, 1940; Sternberg, 1951; Chinnery and
Weishampel, 1998). In contrast, saurischians
retain clavicles with the clavicles contacting at the
midline.

Birds and pterosaurs have an interclavicle. Dinosaurs do not.

223. Coracoid, postglenoid process: (0) short; (1) elongate and expanded posteriorly only 

The significance of this is that a longer coracoid allows the bird scapula (attached to the coracoid) to be positioned high enough on the body (horizontally), to allow flapping (by the wings being raised high enough up).

Birds are like pterosaurs and not like dinosaurs.

230. Humerus, apex of deltopectoral crest situated at a point corresponding to: (0) less than 30% down the length of the humerus; (1) more than 30% down the length of the humerus (fig. 31) (modified from Bakker and Galton, 1974; Benton, 1990a; Juul, 1994; Novas, 1996; Benton, 1999).
Langer and Benton (2006) thoroughly discussed the distribution of the character states of this character and find that state (1) is restricted to dinosaurs within Archosauria. Here, I follow the conclusions and scorings of Langer and Benton (2006). 

The pterosaur deltopectoral crest is like birds. The dinosaur deltopectoral crest is not like birds. (See details below).



231. Humerus, length: (0) longer than or
subequal to 0.6 of the length of the femur; (1)
shorter than 0.6 of the length of the femur
(modified from Novas, 1996; Langer and
Benton, 2006).
Langer and Benton (2006) thoroughly
discussed the distribution of the character
states and find that state (1) is restricted to
Herrerasaurus (PVSJ 373), Eoraptor (PVSJ
512), and neotheropods.

Humerus length compared to femur length is similar for pterosaurs to birds but different than dinosaurs.



370. Astragalus-calcaneum, articulation:
(0) free; (1) coossified (fig. 46) (Sereno and
Arcucci, 1994a; Irmis et al., 2007a).
In most archosauriforms, save avians and
close relatives, the astragalus and calcaneum
are separate elements. In pterosaurs (e.g.,
Dimorphodon, YPM 9182), Lagerpeton (PVL
4619), and Dromomeron romeri (GR 223), the
astragalus and calcaneum are coossified.
Among basal dinosaurs, the proximal tarsals
are coossified in Heterodontosaurus (SAMPK-
1332) and coelophysoids (Rowe and
Gauthier, 1990; Tykoski, 2005b).

More info on the deltopectoral crest.

http://palaeos.com/vertebrates/glossary/glossaryD.html

Deltopectoral crest: a longitudinal ridge or crest on the (proximal) humerus. See figure. Cursorial forms typically do nothave a large crest. It is typically an important attachment point for adductors, rather than retractors.

http://pterosaurheresies.wordpress.com/category/pterosaur-evolution

"Both tiny birds and tiny pterosaurs dispensed with their long stiff tail. In birds it became a pygostyle. In pterosaurs the long stiff tail became a reduced, string-like tail with bead-like verts. Note the similarities in the pectoral girdles. Both could stand with their toes beneath their shoulder glenoids. Both had retroverted pedal digits but of two distinct designs. The anterior ilium of both taxa supported large thigh muscles. A large deltopectoral crest supported large flight adductors anchored to the sternum."

http://palaeos.com/vertebrates/glossary/glossaryD.html

Deltopectoral crest: a longitudinal ridge or crest on the (proximal) humerus. Cursorial forms typically do not have a large crest. It is typically an important attachment point for adductors, rather than retractors.

 http://en.wikipedia.org/wiki/Evolution_of_dinosaurs

Dinosaurs evolved within a single lineage of archosaurs 232-234 Ma (million years ago) in the Ladinian age, the latter part of the middle Triassic. Dinosauria is a well-supported clade, present in 98% of bootstraps. It is diagnosed by many features including loss of the postfrontal on the skull and an ELONGATE deltopectoral crest on the humerus.[1]

The Dinosauria: Second Edition:

The humerus is slender and twisted. It has a caudally deflected proximal end and a moderately developed deltopectoral crest that is restricted to the PROXIMAL third of the humerus in Deinonychus and to the PROXIMAL quarter in Velociraptor.

https://pterosaurheresies.wordpress.com/2015/03/06/problem-the-scansoriopteryx-pelvis-and-humerus/

The deltopectoral crest on most dinosaurs, including Archaeopteryx (Fig. 1), is prominent and extends down a quarter of the humerus. We don’t see this in Scansioropteryx (Fig. 3). But then again, we don’t see this in Aurornis (Fig. 4) either. That doesn’t delete them from the theropod clade because every other aspect of their anatomy says: theropod!

https://pterosaurheresies.files.wordpress.com/2015/03/aurornis-humerus.jpg

http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0001-37652009000400017&lng=es&nrm=iso&tlng=es

Pterosaur humerus:

Description: The specimen MB.R. 2828 (Fig. 2; Table I) has a slightly dorso-ventral compression, showing several fractures. It possesses the characteristic saddle-shaped proximal articular head, common to the pterosaur humeri. The deltopectoral crest is well developed and inclined proximoventrally. It is tongue-shaped with a rounded distal end. In lateral view the proximal margin of the deltopectoral crest is markedly concave, while the distal margin is straight. There is an elongated ridge on the medial side of the crest running from the distal to the proximal edge, and is likely an attachment of a flight muscle (m. pectoralis, see Bennett 2003). There is an elongated concavity on the medial side, close to the distal margin, whose function is unknown. The ulnar crest is blunt and slightly crushed laterally. The dorsal margin shows a well developed pneumatic foramen, located close to the lateral side.

https://books.google.ca/books?id=8CKYxcylOycC&pg=PA117&lpg=PA117&dq=rhamphorhynchus+deltopectoral+crest&source=bl&ots=SqlXdvwI83&sig=2eO8UZBiUTQPDTye_KvTu59iYQw&hl=en&sa=X&ei=MQuQVd_BKpGGNsCtifgC&ved=0CB0Q6AEwAA#v=onepage&q=rhamphorhynchus%20deltopectoral%20crest&f=false
See page 117.

It appears that the pterosaur deltopectoral crest differed between the long-bony-tailed pterosaurs and the short-bony-tailed pterosaurs.
In both cases they were confined to close to the proximal end of the humerus. But the deltopectoral crest in the long-bony-tailed pterosaurs was much smaller.

The deltopectoral crest in the primitive birds was close to the proximal end and relatively small.
In contrast the deltopectoral crest in dinosaurs was elongate. It was not confined to the proximal end of the humerus.



http://courses.washington.edu/chordate/453photos/skeleton_photos/bird-humeri.jpg

Miscellaneous bird humeri (proximal heads to the left). The top 2 face anteriorly & the bottom 3 face posteriorly.

Notice the bird deltopectoral crest is confined to the proximal end as in pterosaurs.


http://en.wikipedia.org/wiki/Dinosaur#Distinguishing_anatomical_features

apex of deltopectoral crest (a projection on which the deltopectoral muscles attach) located at or more than 30% down the length of the humerus (upper arm bone)

Notice the dinosaur deltopectoral crest is not confined to the proximal end as in birds and pterosaurs. It is elongate.

NESBITT (2009) 

11. Furcula shape in anterior view: asymmetrical (0) or symmetrical/nearly symmetrical (1). The furculae of most nonparavian theropods are markedly asymmetrical (e.g., Allosaurus, Citipati). The furculae of paravians, with the exception of Buitreraptor, are nearly symmetrical. It is unclear if the asymmetry of the furcula of Buitreraptor was the result of taphonomy or represents morphological asymmetry.

We see that the dinosaur furcula is not like the Paravian furcula.
That supports the conclusion that they are not related.

Also see (page 77)
http://books.google.ca/books?id=8QRKV7eSqmIC&pg=PA77&lpg=PA77&dq=oviraptor+furcula&source=bl&ots=fqQ1dMcCAm&sig=Mh6VWAuDyk92AmCENy_FFdCRvn0&hl=en&sa=X&ei=g-t4U96CENKdyASEzYDYAw&ved=0CGAQ6AEwCQ#v=onepage&q=oviraptor%20furcula&f=false

THE EARLY EVOLUTION OF ARCHOSAURS:
RELATIONSHIPS AND THE ORIGIN OF
MAJOR CLADES 92011)

The character states supporting pterosaurs
as members of Archosauria and Ornithodira
are not restricted to character states related
to locomotion as suggested by Bennett
(1996). As demonstrated in the list above,
the character states cover features present all
over the body, not just in the hind limb.
Furthermore, it is difficult to argue that the
restricted number of tarsals, the size of the
distal tarsals, and the shape of the proximal
tarsals in pterosaurs would be convergent
with those of dinosauromorphs based on
function alone (Sereno 1991a). In summary,
Pterosauromorpha is well supported as the
sister taxon to Dinosauromorpha.