Yi qi and Occam’s razor

Yesterday and earlier we looked at the new dino-bird, Yi qi. This is the third and final note.

Figure 1. Identification errors (in red) on the original Yi qi diagram from Xu et al. 2015.

Figure 1. Identification errors (in red) on the original Yi qi diagram from Xu et al. 2015. LSE = left styliform element. RSE = right styliform element.

To reiterate: 
In Xu et al. (2015) the authors noted a new long bony element in the fore limb of Yi qi (Fig. 1) never before seen on any bird, dinosaur, or archosaur. They labeled the new bone a ‘styliform element’ in accord with similar elements in flying squirrels. Chemical analysis determined that the new bone was indeed bone. This long bone, as long as (or in the opinion of the authors, longer than) any other forelimb element appeared to be attached at one end to the wrist, and at the other end, to nothing else. In certain flying squirrels, the styliform extends laterally to expand the width of the gliding membrane. Xu et al. also compared the unfamiliar bone to similar bones in gliding marsupials, bats and pterosaurs (the pteroid), all of which arise from different parts of the anatomy, frame extradermal membranes and are employed during aerial exercises.

Xu et al. wrote:
“The most striking feature of Yi is the presence of an anomalous, slightly curved, distally tapered, rod-like structure whose length considerably exceeds that of the ulna, associated with each wrist and apparently extending from the ulnar side of the carpus. However, the rod-like bone of the forelimb of Yi is morphologically unlike any normal theropod skeletal element. Indeed, no equivalent of the rod-like bone is known in any other dinosaur even outside Theropoda,” So Yi qi had potentially analogous structures among other tetrapods, but not among birds or other dinosaurs.

The big question is:
was the new bone indeed longer than the ulna? If you trace the elements as they are (Figs. 1, 2), the new bone is not longer than the ulna.

yi-qi-recon1000b

Figure 2. Yi qi tracing of the in situ specimen using DGS method and bones rearranged, also using the DGS method, to form a standing and flying Yi qi specimen. Note the lack of a styliform element, here identified as a left radius and right ulna.

It’s easy to make errors here
The right radius is incomplete and splintered (Figs. 1-3) so that it looks like a radius and ulna. The distal right radius is lost beyond the edge of the matrix. The right so-called ‘styliform element’ extends beyond the elbow and was presumed to also extend beyond the end of the matrix. However, taken ‘as is’ the two so-called ‘styliform elements’ are actually similar in length to each other and similar to the virtually complete left ulna.

Taphonomic churning
Evidently the elements were churned prior to burial with medial digits of the left hand now lateral and the right foot high kicking like a Rockette. Thus Padian’s comment, “Yi qi’s body is not preserved below the ribcage, so reconstructions of the pelvis, hindlimbs and tail must be conjectured from what is known of other scansoriopterygids,” is incorrect or an oversight (see Figs. 1, 2).

Figure 3. Right ulna (former styliform process) of Yi traced using DGS techniques. See Figure 4 for comparison to Epidendrosaurus.

Figure 3. Right ulna (former styliform process, amber color here) of Yi traced using DGS techniques. See Figure 5 for comparison to Epidendrosaurus. The newly identified ulna was flipped prior to burial so that the thicker proximal end is now closer to the lost wrist. The radius (pink and purple) is cracked lengthwise with toothy ridges marking the break. This gave the impression of a radius and ulna, but no intact radius and ulna have tooth-like cracks and ridges. This is a shattered hollow bone. 

On the left forelimb,
the left so-called ‘radius’ (Figs. 1, 4) is barely present. The left so-called ‘styliform element’ gives the erroneous impression of continuing beneath the left humerus. It does not.

Figure 3. Closeup of the former 'styliform element' here identified as a radius in Yi qi.

Figure 4. Closeup of the former ‘styliform element’ here identified as a left radius in Yi qi.

The authors never considered the possibility
that the ulna and radius could have been splintered during crushing prior to burial. Both forearm elements are as hollow as long bones from other dino-birds. And they have been subject to torsion and churning prior to burial. In this scenario the purported left ‘radius’ (Fig. 4) is simply a splinter of the ulna, separated during axial torsion. Similarly, the purported right ‘ulna’ is a splinter of the right radius (Fig. 3).

In this hypothesis
the curved right ‘styliform element’ is in reality an ulna and the straighter left ‘styliform element’ is a radius. As preserved the new radius and ulna are about the same length and similar in diameter to their counterparts. In birds and sister taxa, like Epidendrosaurus (Fig. 5), the radius is typically straighter than the ulna. And the ulna tapers distally.

In the Xu et al. 2015 Supplemental Data,
the ‘styliform element’ lengths each have an asterisk associated with them: 133.5* and 91.3* mm. The ulna likewise has an asterisk measurement, 88.5* mm. The length of the radius was not estimated or shown. The authors note: “* indicates estimated value; the preserved length of the right styliform element is 91.3 mm, but taphonomic information and morphological comparisons between the right and left styliform elements lead us to estimate that the total length of the styliform element is 133.5 mm.” In this case, they overestimated. Perhaps a more precise tracing would have been helpful.

Occam’s razor
As everyone knows, “The principle states that among competing hypotheses that predict equally well, the one with the fewest assumptions should be selected.” Xu et al. struggled with the identity of the odd long bones in Yi qi. Unfortunately they did not select the identity with the fewest assumptions (and autapomorphies). Instead they made headlines around the globe and more than a few eyebrows rise with the invention of a new styliform element, which is really just a misidentified common fore arm element.

In this case
the left ‘styliform element’ articulates with the wrist because the left radius likewise articulates with the wrist. That the distal end no longer articulates with the elbow can be ascribed to taphonomic torsion prior to burial.

The right radius still articulates with the humerus, but the right ulna has flipped lengthwise, such that the narrow distal end (Fig. 4) is now proximal, far behind the elbow. While more difficult to visualize how this may have happened, this bone flip also must be the product of taphonomic churning prior to burial. Perhaps it goes along with the high kick of the foot. Comparisons to the articulated specimen of Epidendrosaurus (Fig. 4) are instructive here. The ulna shapes are virtually identical.

Figure 4. Right ulna of Yi (former 'styliform element') compared to right ulna of Epidendrosaurus.

Figure 5. Right ulna of Yi (former ‘styliform element’) compared to right ulna of Epidendrosaurus.

References
Padian K. 2015. Paleontology: Dinosaur up in the air. Nature (2015) doi:10.1038/nature14392
Xu X, Zheng X-T, Sullivan C, Wang X-L, Xing l, Wang Y, Zhang X-M, O’Connor JK, Zhang F-C and Pan Y-H 2015.
 A bizarre Jurassic maniraptoran theropod with preserved evidence of membranous wings.Nature (advance online publication)
doi:10.1038/nature14423

Notes on Limusaurus (Dinosauria, Theropoda) and its odd little hand

Figure 1. Limusaurus in situ. The associated croc has been painted red here.

Figure 1. Limusaurus in situ. The associated croc has been painted red here. Their marriage was legal in only 16 states.  : )

Limusaurus inextricabilis (Xu et al. 2009; earliest Late Jurassic, Oxfordian; 1.7m in est. length; IVPP V 15923; Figs. 1, 2) is an herbivorous theropod with very tiny arms and hands in the lineage of proto-birds. Coincidentally, Limusaurus was buried next to the skeleton of a tiny Jurassic croc (in red above). Pol and Rauhut (2012) nested Limusaurus with Elaphrosaurus and Spinotropheus. Those taxa have not been added yet to the large reptile tree where Limusaurus nests between Sinocalliopteryx and Aurornis. Limusaurus shares a ventral pelvis, but not long arms and large hands with these bird-like taxa. Like birds, Limusaurus has a pair of sternae. The coracoids appear to be transitional between disc-like and stem-like.

Figure 2. Limusaurus reconstructed. Both hands are shown and colorized. Digit 1= purple. Digit 2= pink. Digit 3 = Green. Digit 0=pale yellow. Digit 0 goes back to basal tetrapods, like Ichthyostega, and only appear here due to the vestigial nature of the manus in which it matured at an embryologically immature state compared to sister taxa.

Figure 2. Limusaurus reconstructed. Both hands are shown and colorized. Digit 1= purple. Digit 2= pink. Digit 3 = Green. Digit 0=pale yellow. Digit 0 goes back to basal tetrapods, like Ichthyostega, and only appear here due to the vestigial nature of the manus in which it matured at an embryologically immature state compared to sister taxa.

Limusaurus is notable for two main reasons:

  1. It is an herbivorous theropod from the earliest Late Jurassic
  2. It has four fingers when all sister taxa have three. This fact has added credence and confusion to the chick embryo digit identification issue we looked at earlier here

It’s well worth looking at both sides of this extra finger issue:
From the Xu et al. abstract: “Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156–161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.”

There were a long list of authors (see below) that agreed with the above, some of whom have come to our attention earlier for publishing various errors, along with their otherwise excellent work.

Unfortunately the Xu et al. abstract ignores the fact that basal tetrapods, like Acanthostega, had an additional digit medial to digit #1 (we’ll call this digit #0). Therefore digit #0 is part of the phylogenetic history of all tetrapods, despite the fact that it is almost never expressed in tetrapod adults, only embryos. Digit #0 is expressed in Limusaurus, in which the hand is a small vestige relative to those of sister taxa. Like other vestiges the hand of Limusaurus did not continue to develop normally as a hatchling and into adulthood. Rather the hand retained a shape found at a certain embryological stage, a stage that included digit #0.  That’s why the big metacarpal (in purple) has the morphology of metacarpal 1 in sister taxa (Fig. 3) and metacarpal #0 continues the shape of metacarpal #1 in cross section. In most tetrapods digit #0 is fused to metacarpal #0 or otherwise disappears before birth or hatching.

Figure 3. The manus of several theropods including Limusaurus. Here digit 1 is purple, digit 2 is pink and digit 3 is green. Note the presence of digit 0 in this vestigial hand, a holdover from basal tetrapods that has not been correctly identified by Xu et al. and others.

Figure 3. The manus of several theropods including Limusaurus. Here digit 1 is purple, digit 2 is pink and digit 3 is green. Note the presence of digit 0 in this vestigial hand, a holdover from basal tetrapods that has not been correctly identified by Xu et al. and others. Click to enlarge.

That’s why otherwise you only see digit #0 in embryos. Thus there is no “phase shift” of digit identity. There is only loss, fusion or absorption of digit #0, a factor missed by earlier workers.

References
Xu X, Clark JM. Mo J, Choiniere J, Forster CA, Erickson GM, Hone DWE, Sullivan C, Eberth DA, Nesbitt S, Zhao Q, Hernandez R, Jia C-K, Han F-L. and Guo Y 2009. A Jurassic ceratosaur from China helps clarify avian digital homologies. Nature, 459(18): 940–944.

Chilesaurus, new dinosaur: not so ‘enigmatic’ after all…

Chilesaurus diegosuarezi (Novas et al. 2015; Late Jurassic, 150 mya, Fig. 1) is the current media darling. Described as an ‘enigmatic’ and ‘bizarre’ theropod, Chilesaurus was nested  with Velociraptor, Tawa and kin, and Elaphrosaurus using various prior cladograms in the supplementary data. So that’s an issue (no internal agreement).

Several articulated specimens are known at distinct ontogenetic stages.

Unfortunately taxon exclusion raises its ugly head again…
The large reptile tree nests Chilesaurus outside of the Theropoda, near the base of the Phytodinosauria, at the base of the Ornithischia and at the base of the clade that also includes Daemonosaurus and Jeholosaurus (Fig. 1), two taxa that were unfortunately ignored by the Novas et al. study. Hate to see that happen yet again.

Figure 1. Chilesaurus and kin, including Damonosaurus and basal phytodinosauria.

Figure 1. Chilesaurus and kin, including Damonosaurus and basal phytodinosauria to scale.

Figure 2. Look familiar? Here are the pelves of Jeholosaurus and Chilesaurus compared.  As discussed earlier, this is how the ornithischian pelvis evolved from that of Eoraptor and basal saurorpodomorpha.

Figure 2. Look familiar? Here are the pelves of Jeholosaurus and Chilesaurus compared. As discussed earlier, this is how the ornithischian pelvis evolved from that of Eoraptor and basal saurorpodomorpha..

Folks,
Chilesaurus is not bizarre.

It is simply a descendant from an unknown Late Triassic transitional taxon at the base of the Ornithischia, a hypothesis overlooked by Novas et al. Chilesaurus is not a theropod, but a phytodinosaur (Fig. 3). The fact that fossils of Chilesaurus were found much later than the original split is not a cause for concern. That happens all the time.

Pelvis  changes
Along with Jeholosaurus, Chilesaurus demonstrates the changes that were happening to the dinosaurian pelvis at the genesis of the ornithischian pelvis. As a plant eater, Chilesaurus and kin were expanding their gut volume to digest less digestible plant matter.

Manus and tooth changes
The hands of Chilesaurus are not as primitive and plesiomorphic as might be hoped, but then Chilesaurus is a descendent of that Late Triassic transitional taxon, appearing tens of millions of years after the split. Things evolve! While the teeth remain large and robust, Chilesaurus had flat teeth, rather than the pointed ones of its Triassic sister, Daemonosaurus or its Cretaceous sister, Jeholosaurus.

Figure 3. Cladogram of basal dinosaurs. Note that Chilesaurus nests near the base of the Phytodinosauria and at the base of the Ornithischia, both far from the Theropoda.

Figure 3. Cladogram of basal dinosaurs. Note that Chilesaurus nests near the base of the Phytodinosauria and at the base of the Ornithischia, both far from the Theropoda.

Clues
to the largely missing post-crania of Daemonosaurus are provided by its sister Chilesaurus.

Now let’s talk about the PR barrage
This is where the science reporters separate themselves from the scientists. All those who reported without testing the results of Novas et al. … you may have to do some backtracking.

References
Novas FE, Salgado, Suárez LM, Agnolín FL, Ezcurra MND, Chimento NSR.,de la Cruz R, Isasi MP, Vargas AO, Rubilar-Rogers D. 2015. An enigmatic plant-eating theropod from the Late Jurassic period of Chile. Nature. doi:10.1038/nature14307

Male and Female Stegosaurus?

Usually I leave dinosaurs to the dinosaur experts…
but this new paper seems to be appropriate fodder.

Figure 1. Click to enlarge. According to Saitta 2015, male and female Stegosaurus can be differentiated by their plates.

Figure 1. Click to enlarge. According to Saitta 2015, male and female Stegosaurus can be differentiated by their plates.

 

A recent PlosOne paper by Saitta (2015) claims Stegosaurus sexual dimorphism can be determined by plate shape (Fig. 1).

Unfortunately others disagree.
Drs. Kevin Padian and Ken Carpenter raised serious issues here.

And part of the problem,
perhaps a major part of the problem, is the lack of articulation in the specimens (Fig. 2).

Figure 2. An insitu plot of the Stegosaurus material. Deep blue colors indicate plates, provided by Saitta. Three pelves of different shapes and sizes are marked. Arrows point anteriorly. This is a jumble. And the plates are disarticulated.

Figure 2. An insitu plot of the Stegosaurus material. Deep blue colors indicate plates, provided by Saitta. Three pelves of different shapes and sizes are marked. Arrows point anteriorly. This is a jumble. And the plates are disarticulated.

A plot
of the in situ specimens (Fig. 2)  indicates that at least three individuals are shown in disarray here, (five were mentioned and likely the others are from other parts of the site). One is smaller than the others. How is it possible to match plates to pelves? And how do all the other bones fit herel? I would not want to attempt a reconstruction with this scattering of at least three individuals.

This is a hard hypothesis to substantiate. 
Not only do different stegosaurs have different shaped plates, but nearly every plate on every stegosaur is distinct, even in articulated specimens.

What I find most interesting…
How did Drs. Padian and Carpenter get their comments published online at ScienceMag.org on the same day the PlosOne paper came online? Only Carpenter is listed in the acknowledgments (for providing specimen photos). Both are listed in the references.  I assume they were not referees, but must have been granted access to the paper prior to publication.

And, why are their no comments in the COMMENTS section for this paper (at the time of this publication)? That’s the standard operating procedure for PlosOne papers.

References
Saitta ET 2015. Evidence for Sexual Dimorphism in the Plated Dinosaur Stegosaurus mjosi (Ornithischia, Stegosauria) from the Morrison Formation (Upper Jurassic) of Western USA. PLoS ONE 10(4): e0123503. doi:10.1371/journal.pone.0123503

Marasuchus skull restored

Updated March 13 with text and figure corrections and overlooked references.

Very few skull bones are known for Marasuchus, the tiny theropod-like dinosaur or proto-dinosaur. Here’s a shot at a restoration of the skull (Fig. 1).

Figure 1. Marasuchus skull restored. My what big teeth you have! Based on the maxilla and occiput, this appears to be a long, low skull. Looks like a little basal theropod, like Tawa. Line drawing from Theropod Database in which M. Mortimer moved the 'quadrate' to the postorbital, based on Bonaparte 1975.

Figure 1. Marasuchus skull restored. My what big teeth you have! Based on the maxilla and occiput, this appears to be a long, low skull. Looks like a little basal theropod, like Tawa. Line drawing from Theropod Database in which M. Mortimer moved the ‘quadrate’ to the postorbital, based on Bonaparte 1975.

Marasuchus would make a pretty good little basal theropod/basal dinosaur EXCEPT,

  1. each cervical is too short
  2. no cervicals have epipophyses
  3. the pubis is too short
  4. and it has no boot
  5. the ischium is too deep and V-shaped along its entire length.
  6. the femur lacks some grooves and bumps found in sister taxa
  7. the proximal tibia has a lateral bump does not reach the posterior rim
  8. distal tarsal 4 is not flat
  9. the astragalus has a larger facet for a larger fibula

Marasuchus is also smaller than basal dinosaur/theropod sisters (Fig. 2) and, considering this list, one wonders if some of these traits are due to neotony, the juvenilization of traits when a taxon experiences generational miniaturization.

Figure 1. To scale compared to Marasuchus, Agnosphitys cromhallensis (Fraser et al. 2002) is known from a selection of uncrushed bones, all of which resemble those from Marasuchus, but slightly larger with a relatively longer rostrum and shorter arms. These two represent a separate and distinct lineage of theropods.  Click to enlarge.

Figure 2. To scale compared to Marasuchus, Agnosphitys cromhallensis (Fraser et al. 2002) is known from a selection of uncrushed bones, all of which resemble those from Marasuchus, but slightly larger with shorter arms. These two represent a separate and distinct lineage of theropods.  Click to enlarge.

If not as a basal theropod close to the odd theropods, Procompsognathus and Segisaurus, then where else could Marasuchus more parsimoniously nest? Most of the above traits can be found individually far from bipedal dino-types, but the suite cannot be found elsewhere. I think we have to rely on maximum parsimony here.

Your thoughts?

References
Bonaparte JF 1975. Nuevos materiales de Lagosuchus talampayensis Romer (Thecodontia – Pseudosuchia) y su significado en el origen de los Saurischia, Chañarense Inferior, Triasico Medio de Argentina [New materials of Lagosuchus talampayensis Romer (Thecodontia – Pseudosuchia) and its significance on the origin of the Saurischia, Lower Chañares, Middle Triassic of Argentina]. Acta Geológica Lilloana 13:5-90.
Romer AS 1971. The Chanares (Argentina) Triassic reptile fauna X. Two new but incompletely known long-limbed pseudosuchians: Brevoria 378: 1-10.
Romer AS 1972. The Chanares (Argentina) Triassic reptile fauna. XV. Further remains of the thecodonts Lagerpeton and Lagosuchus: Breviora 394: 1-7.
Sereno PC and Arcucci AB 1994. Dinosaurian precursors from the Middle Triassic of Argentina: Marasuchus lilloensis gen. nov. Journal of Vertebrate Paleontology, 14: 53-73

Theropod Database

wiki/Marasuchus

Daemonosaurus, the Phytodinosauria and the Persistence of the Postfrontal

Earlier
we nested the Triassic saber-tooth, Daemonosaurus, at the base of the Ornithischia and near the base of the Phytodinosauria. The addition of the sauropodomorph, Leyesaurus (Apaldetti et al. 2014, Triassic/Jurassic boundary) to the large reptile tree supports that nesting with a very similar skull (Fig.1).

Figure 1. Click to enlarge. Sisters to Daemonosaurus, including Leyesaurus and Jeholosaurus. The postfrontal (in light red) is not fused in most of these taxa (Heterodontosaurus is the exception), contra current dinosaur paradigms. Note the resemblance of Daemonosaurus to the basal sauropodomorph, Leyesaurus. The increase in tooth size in Daemonosaurus was not derived from theropods, but was a unique character trait, shared, more or less with its sister, Jeholosaurus and to a lesser extent in Heterodontosaurus.

Figure 1. Click to enlarge. Sisters to Daemonosaurus, including Leyesaurus and Jeholosaurus. The postfrontal (in light red) is not fused in most of these taxa (Heterodontosaurus is the exception), contra current dinosaur paradigms. Note the resemblance of Daemonosaurus to the basal sauropodomorph, Leyesaurus. The increase in tooth size in Daemonosaurus was not derived from theropods, but was a unique character trait, shared, more or less with its sister, Jeholosaurus and to a lesser extent in Heterodontosaurus.

Along the way
it appeared that the postfrontal was retained in basal dinosaur taxa and outgroups, contra traditional dinosaur thinking.

Leyesaurus and Daemonosaurus
share greatly elongate cervicals and other traits (Fig. 1). Daemonosaurus was not included in the original analysis, but then Leyesaurus did not nest as a basal sauropodomorph as it does here. Leyesaurus has quite a wide skull, distinct from most basal dinosaurs. It nests with Massospondylus, which earlier we noted looked a lot like Daemonosaurus.

References
Apaldetti C, Marinez RN, Alcober OA and Pol D 2014. A New Basal Sauropodomorph (Dinosauria: Saurischia) from Quebrada del Barro Formation (Marayes-El Carrizal Basin), Northwestern Argentina. PLoS ONE 6(11): e26964. doi:10.1371/journal.pone.0026964

.

Daemonosaurus has two sister taxa: Haya and Jeholosaurus

Updated February 28, 2015 with a new skull for Daemonosaurus.

Earlier we talked about the Late Triassic saber-toothed dinosaur, Daemonosaurus (Sues et al. 2011, Fig. 1, CM 76821) originally considered a weird basal theropod between Eoraptor and Tawa. That’s due to taxon exclusion. The real sisters of Daemonosaurus were not tested. The authors also mistakenly nested Eoraptor within the Theropoda when it is actually an outgroup, a phytodinosaur closer to Sauropodomorpha, which were only included as a suprageneric taxon, along with Ornithischia. Unfortunately that’s the same suprageneric/taxon exclusion/inclusion problem that happens so often it’s not funny anymore.

Figure 1. Skulls of Daemonosaurus, Haya and Jeholosaurus to scale.

Figure 1. Skulls of Daemonosaurus, Haya and Jeholosaurus to scale nest as sister taxa. And it’s easy to see why. Somewhere in this clade lies the origin of the predentary bone and the retroverted pubis. Long premaxillary teeth and a short rostrum are key traits. Note the infilling of the mandibular fenestra.

Solution: add taxa and avoid suprageneric taxa
In the large reptile tree Daemonosaurus did not nest with theropods, but at the base of the Ornithischia between basal phytodinosaurs like Eoraptor and the basal ornithischian, Pisanosaurus. It’s been three years since that post.

Today Jeholosaurus (Han et al 2012) and Haya (Figs. 1,2), two widely acknowledged basal ornithischians, nest with Daemonosaurus. One look at the three of them together pretty much sums up the rest of this post. Note their chronology. This is a basal clade that lasted through all three periods of the Mesozoic.

That they nest together tells me the post-crania of Daemonosaurus likely had at least a proto-ornithischian pelvis and supports my earlier observation of a proto-predentary.

Figure 1. Haya skull and post-crania.

Figure 1. Haya skull and post-crania. At present this specimen gives us the best approximation of the post-crania of Daemonosaurus, although the neck vertebrate were longer. Note the stub of a fifth toe on the pes.

Fossils of Coelophysis were present on the same block that contained the skull of Daemonosaurus, Wonder if there was a predator/prey relationship? Skull lengths were similar. Overall size was likely similar too.

References
Han F-L, Barrett PM, Butler RJ and Xu X 2012. Postcranial anatomy of Jeholosaurus shangyuanensis (Dinosauria, Ornithischia) from the Lower Cretaceous Yixian Formation of China. Journal of Vertebrate Paleontology 32:1370-1395.
Makovicky PJ, Kilbourne BM, Sadleir RW and Norell MA 2011. A new basal ornithopod (Dinosauria, Ornithischia) from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology 31: 626–640.
Sues H-D, Nesbitt SJ, Berman DS and Henrici AC 2011. A late-surviving basal theropod dinosaur from the latest Triassic of North America. Proceedings of the Royal Society Bpublished online 
Xu, Wang and You, 2000. A primitive ornithopod from the Early Cretaceous Yixian Formation of Liaoning. Vertebrata PalAsiatica 38(4)318-325.

wiki/Daemonosaurus
wiki/Haya
wiki/Jeholosaurus