Another look at a possible pterosaur wingtip ungual

Figure 1. The Yale specimen of Rhamphorhynchus phyllurus with preserved wingtip ungual highlighted. See figure 2 for closeup.

Figure 1. The Yale specimen of Rhamphorhynchus phyllurus with preserved wingtip ungual highlighted. See figure 2 for closeup.

The Yale specimen of Rhamphorhynchus phyllurus (Figs. 1, 2; VP 1001778) has one painted wing tip and one that may include another wingtip ungual.

Figure 2. Closeup of Rhamphorhynchus phyllurus in figure 1 focusing on the preserved wingtip ungual.

Figure 2. Closeup of Rhamphorhynchus phyllurus in figure 1 focusing on the preserved wingtip ungual. Was this carved in? Or is it real? Note the cylindrical tip of the penultimate wing phalanx (m4.4). The wingtip was buried deep within the matrix and had to be exposed.

The wingtip
was buried deep within the matrix and had to be exposed. So the question is: was it carved? Or is it real? If it was carved, why was it carved? Traditionally pterosaurs are not supposed to have wing tip unguals, but I’ve found them in several specimens.

You might remember
we looked at this wing tip earlier with a different provided image. The present one appears to offer more clues.

The paint-by-numbers analogy to phylogenetic analysis

Figure 1. owl paint-by-numbers

Figure 1. Owl paint-by-numbers. If you accurately add color to each little shape, pretty soon a picture will emerge. You don’t have to compose it. That’s already been done for you. Follow this method and your result will echo the original composition,  lighting and subject matter to a great degree. 

Phylogenetic analysis is like a paint-by-numbers kit.
You fill in each little color by following the instructions. Or you fill in each little matrix box (taxon/character) with the correct score. Only afterwards do you see the big picture. Or only afterwards does the software produce the resulting cladogram, the big picture of hypothetical relationships.

By contrast, in traditional painting
the master artist starts with a loose sketch, then arranges elements in a composition to fit a triangle, a golden rectangle, or some other substructure. The colors, tints and shadows are added in large blocks to a canvas of the right size to fit a certain wall. Finally the details (lace, highlights, eyelashes, etc. are added.

Like a paint-by-numbers canvas,
the big picture in evolution has already happened. The “instructions” or “clues” come to us in the form of preserved and exposed traits in fossils and living taxa. We don’t have all the clues, and never will, but with what we do have we fill them in until a complete picture begins to emerge, blank spaces and all.

Likewise, the large reptile tree
(LRT) and large pterosaur tree (LPT) are large gamut cladograms that will never be completed. However, as new taxa are added the details and transitions between established taxa become finer and finer blends. The big picture, or tree topology, has been pretty steady for several years and hundreds of additions.

Make sure your taxa 
are all species or specimens. Those provide good data. Avoid suprageneric taxa. By combining traits from several genera you’ll muddy the canvas. The tiny features will be lacking. You’ll cherry-pick favorites and overlook obscure details that might be Important.

Science is for everyone
Not just for PhDs. If they can create a cladogram, so can you. They test published work for validity. So do I and so can you. Along the way, you will make mistakes. I do too. Others will point out mistakes. Defend your decisions where appropriate. Fix problems at every opportunity. Follow this method and your result will echo the original tree topology. Then keep adding taxa as they become available to fill in any blank spaces.

The first time an idea is proposed
it is rarely accepted. As time goes by, some hypotheses disappear. And some should disappear. Others, whether valid or not, get headlines because the PR machinery is tilted in their favor. Still others slowly grow in acceptance and are ultimately embraced because they reflect the original tree topology we’re all trying to see more clearly.

Good luck on all your endeavors.

Coincidence? Or Discovery?

A recent reply (see below) to an earlier post bears noting:

Diandongosuchus nests as a basal phytosaur when choristoderes and basal younginoids are included, far from Qianosuchus, which also does not nest with poposaurs, which are all bipedal (or formerly bipedal) herbivores, a far cry from Diandongosuchus.

Figure 1. Diandongosuchus nests as a basal phytosaur when choristoderes and basal younginoids are included, far from Qianosuchus, which also does not nest with poposaurs, which are all bipedal (or formerly bipedal) herbivores, a far cry from Diandongosuchus.

David Marjanović on April 12, 2017 at 3:16 am said: 
“The redescription of Diandongosuchus (Fig. 1) has now been published in open access. I’m afraid I can’t congratulate you. The new paper, and the SVP abstract before it, uses data you didn’t (and couldn’t) use – you were right for the wrong reasons. No congratulations for coincidences. :-|  “

Reply ↓
davidpeters1954 on May 22, 2017 at 8:31 pm said:
“So, phylogenetic analysis and expanding the inclusion set are the wrong reasons? Tsk, tsk, David. Your bias is showing.”

Back story:
Diandongosuchus (Li et al. 2012) was originally nested with poposaurs. Within a few days of its publication, Diandongosuchus was added as a taxon to the large reptile tree (LRT) and it nested not with poposaurs, but at the base of the phytosaurs. Several other blog posts here, here and here further illustrated the link.

Recently 
Stocker et al. 2016 also nested Diandongosuchus with phytosaurs and shortly thereafter news of that publication was posted here,

Botton line:
Stocker et al. did not recognize the earlier discovery. It was easy to Google. It would have been appropriate to add the original discoverer to the list of authors. This is common practice, even when that person is deceased. More recently Dr. Marjanović withheld congratulations and demeaned the scientific method by which the discovery was attained (an expanded taxon list employed in phylogenetic analysis) as “the wrong reasons.”

 

Carl Sagan once wrote:
“In a lot of scientists, the ratio of wonder to skepticism declines in time. That may be connected with the fact that in some fields—mathematics, physics, some others—the great discoveries are almost entirely made by youngsters.”

“The suppression of uncomfortable ideas may be common in religion or in politics, but it is not the path to knowledge; it has no in the endeavor of science. We do not know in advance who will discover fundamental insights.”

“There are many hypotheses in science which are wrong. That’s perfectly all right; they’re the aperture to finding out what’s right. Science is a self-correcting process. To be accepted, new ideas must survive the most rigorous standards of evidence and scrutiny.”

The hypothesis
that Diandongosuchus is more closely related to phytosaurs than to poposaurs originally appeared here in 2012 and was confirmed four years later by Stocker et al. That Dr. Marjanović does not approve of the earlier discovery tell us more about professional biases against ‘outsiders’, which we’ve seen before, than it does about the ‘coincidence’ he conjures.

 

References
Li C, Wu X-C, Zhao L-J, Sato T and Wang LT 2012. A new archosaur (Diapsida, Archosauriformes) from the marine Triassic of China, Journal of Vertebrate Paleontology, 32:5, 1064-1081.
Stocker MR, Nesbitt SJ, Zhao L-J, Wu X-C and Li C 2016. Mosaic evolution in phytosauria: the origin of longsnouted morphologies based on a complete skeleton of a phytosaur from the Middle Triassic of China. Abstracts of the Society of Vertebtate Paleontology meeting 2016.

 

Entelognathus: revisions

Yesterday we looked at Entelognathus (Figs. 1-3; Zhu et al. 2013), a Silurian placoderm transitional to bony fish. That was my first placoderm and I made some errors that have since been corrected. Those errors were corrected when I realized the frontal (pineal in placoderms and Cheirolepis) originated as a tiny median (purple) triangle that included the pineal opening. I was also confused by the splitting of the parietal in Osteolepis, which I thought gave rise to the parietal/postparietal split, but instead that is an autapomorphy arising only in certain Osteolepis specimens. Further confusion comes from the fusion of bones, the splitting of bones and the different names given to the same bone in Silurian to Devonian taxa. Because of this, today and today only I will call the bones by the colors provided by Zhu et al. A key to their various names is provided (Fig. 1).

I was also surprised
to see that Zhu et al. 2013 found no trace of a purple/orange division in Entelognathus (Fig. 1f). This is odd for a transitional taxon, but still possible. Worth looking into. Equally odd, Zhu et al. did not color the purple bone consistently (Fig. 1).

The pineal opening drift
from the purple to the orange bones attends the lengthening of the rostrum and perhaps the brain and olfactory regions. The purple bone invades the paired orange bones and at the posterior tip of the parietal is the pineal opening. So the purple bone more or less delivers the pineal opening more or less in the middle of the orange bones.

Figure 1. From Zhu et al. 2013 SuppData showing placoderm and other basal vertebrate skull roofs. Note: Entelognathus is the only taxon without frontals, which I found in the photos of the fossil, figure 2.

Figure 1. From Zhu et al. 2013 SuppData showing placoderm and other basal vertebrate skull roofs. Note: Entelognathus is the only taxon without a frontal/parietal split, which I found in the photos of the fossil, figure 2 and corrected at the tip of the long arrow.

I traced bone sutures on photos of the specimen
and found that purple/orange division. So now Entelognathus has a complete set of skull roofing bones from the nasal to the frontal to the parietal and post parietal. I may have even seen where the yellow green intertemporal splits from the orange parietal.

Figure 2. Entelognathus fossil. Scale bar = 1 cm. Here the frontal/parietal division is shown.

Figure 2. Entelognathus fossil. Scale bar = 1 cm. Here the frontal/parietal division is shown. Rather than a median uture, one finds a medial ridge.

I hope to never do another fish.
But happy that I was able to resolve some earlier questions and move on. Feelings aside, mistakes that go on unnoticed are worse than mistakes you, or others, find and correct.

Figure 1. Entelognathus drawings from Zhu et al. 2013, with colors and homologous tetrapod bone. abbreviations added.

Figure e. Entelognathus drawings from Zhu et al. 2013, with colors and homologous tetrapod bone. abbreviations added. Corrected from an earlier version.

References
Zhu M, Yu X-B, Ahlberg PE, Choo B and 8 others 2013. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature. 502:188–193.

Cheirolepis fossil images
wiki/Cheirolepis
wiki/Eusthenopteron
wiki/Entelognathus

 

Placoderm Entelognathus skull bones re-identified with tetrapod homologies

Images repaired May 18, 2017 after studying photos of the specimen, comparing related taxa and dispensing with false paradigms. Click here for more details. 

Barford 2013 wrote: 
“It may be hard to see, but you seem to share a family resemblance with Entelognathus primordialis. The fish, which lived 419 million years ago in an area that is now part of China, is the earliest known species with a modern jaw.” Here (Fig. 1) one can identify a complete set of homologous tetrapod skull bones understood by the original authors, who identified the bones with traditional placoderm names. (Ala, placoderms, bony fish and sacropterygians, including tetrapods, have different names for the same bone). And they made a mistake or two along the way, none of which negate their conclusions, but cement them.

I never thought I’d be featuring any placoderm fish in this blog
or in ReptileEvolution.com, but Entelognathus, as everyone already knows — and I just learned, is something very special. A major discovery. And this was my first day studying placoderms.

Barford 2013 reported, “Palaeontologists have traditionally believed that the fishes’ features bore no relation to ours. They assumed that the placoderm face was lost to evolutionary history, and most thought that the last common ancestor of living jawed vertebrates had no distinct jawbones — that it was similar to a shark, with a skeleton made mostly of cartilage and at most a covering of little bony plates. The theory went that the bony fishes evolved later, independently developing large facial bones and inventing the ‘modern’ jaw. Such fishes went on to dominate the seas and ultimately gave rise to land vertebrates. [Entelognathus] has what looks like a bony fish’s jaw, even though it is older than the earliest known sharks and bony fishes.”

According to Wikipedia
Entelognathus
 primordialis
 (Zhu et al. 2013; Late Ludlow, Silurian, 419 mya; IVPP V18620) “is a genus of placoderm fish with dermal marginal jaw bones (premaxilla,
maxilla and dentary), features previously restricted to Osteichthyes (bony fish).”

More than that,
all of the skull bones find homologies in tetrapods and bony fish (Figs. 1, 2) when certain bones are correctly identified or homologized. It just takes a few colors here and there to make it all clear.

Figure 1. Entelognathus drawings from Zhu et al. 2013, with colors and homologous tetrapod bone. abbreviations added.

Figure 1. Entelognathus drawings from Zhu et al. 2013, with colors and homologous tetrapod bone. abbreviations added. This revised image adds a small triangular frontal between the anterior processes of the parietal and the rest of the bones follow suit. 

All of the bones in the skull of Entelognathus
find homologies with those in Cheirolepis (Whiteaves 1881; Fig. 2) and also with tetrapods. Entelognathus lived 59 million years before the appearance of tetrapods like Ichthyostega. and is someday going to be a part of the story behind those Middle Devonian footprints.

Here new labels and colors
repair original errors and indicate tetrapod homologies in Entelognathus (Zhu et al. 2013).

  1. Three purported sclerotic bones are circumorbital bones (prefrontal, postfrontal, jugal)
  2. The purported jugal is the dorsal half of the maxilla before these bones fused.
  3. The purported quadratojugal is the posterior of the maxilla
  4. The rostral is the nasal
  5. The triangular frontal was overlooked
  6. The pineal plate is a pair of parietals
  7. The central plate is a pair of postparietals
  8. The marginal plate is the supratemporal
  9. The anterior paranuchal plate is the tabular
  10. The opercular is the quadratojugal
Figure 2. Cheirolepis skull (left) with skull bones colorized as in Osteolepis (right) and Enteognathus, figure 1. Colors make bone identification much easier. Note the post opercular bone differences between Osteolepis and Cheirolepis indicating separate and convergent derivation, based on present data.

Figure 2. Cheirolepis skull (left) with skull bones colorized as in Osteolepis (right) and Enteognathus, figure 1. Colors make bone identification much easier. Note the post opercular bone differences between Osteolepis and Cheirolepis indicating separate and convergent derivation, based on present data.

On the subject of nomenclature
Zhu et al. 2013 (SuppData) list the various names given to fish skull bones and their homologies in other fish clades. Some of the more confusing include:

  1. The parietal in sarcopterygians is the frontal in actinopterygians and the preorbital in placoderms.
  2. The postparietal in sarcopterygians is the parietal in actinopterygians and the central in placoderms.
  3. The supratemporal in sarcopterygians is the intertemporal in actinopterygians and the marginal in placoderms.
  4. The tabular in sarcopterygians is the supratemporal in actinopterygians and the anterior paranuchal in placoderms.
  5. And there are others…

Where is the authority that can fix this problem?
But if we fix it, then what? Then all prior literature will have to be translated. Either way, we’re hosed. Maybe we should just colorize homologous bones and leave it at that, as Zhu et al. did in their SuppData.

Entelognathus precedes Cheirolepis by 29 million years.
Preopercular and opercular bones do not appear in Entelognathus, but are present in Cheirolepis. So they are new bones in osteichythys.

The ‘al’ bone in Entelognathus (Fig. 1) is the cleithrum, supporting the pectoral fin.

The split (spiracle) between the skull roofing bones (intertemporal. supratemporal, tabular) and cheek bone (squamosal) do not appear in Entelognathus, but do so in Cheirolepis.

Sclerotic rings are not necessary in such small and well-protected eyes as in Entelognathus and if present, would have been very tiny and fragile.

Comparisons of the circumorbital bones in Entelognathus and Cheirolepis are strikingly similar down to the small post-orbit depression in the jugal in Entelognathus that becomes a notch in Cheirolepis.

Comparisons of the postopercular bones
of Cheirolepis and Osteolepis (Fig. 2) show little to no homology, suggesting a possible separate but convergent derivation.

Note some skull bones
later split apart at the median, while others fuse together. It’s their shapes and locations that identify them. “The large hexagonal central plate seems to have a single ossification centre, whereas most placoderms have paired centrals,” reports Zhu et al, making a case in point. A pineal opening is not present in the pineal plate (fused parietals) of Enteleognathus. This is further evidence that the pineal opening migrated from the frontals to the parietals over tens of millions of years. More on that tomorrow.

Barford 2013 concludes
“There remains a chance that E. primordialis evolved its jaw independently from the bony fish, so that we did not inherit it, and the resemblance is an illusion.” I don’t agree with this conclusion. The evidence for homology elsewhere overwhelms any competing hypotheses.

Friedman and Brazeau (2013) also comment on this discovery.
First, Entelognathus alwaybranches outside the radiation of living jawed vertebrates, meaning that key components othe osteichthyan face are no longer unique innovations of that group. Second, acanthodians — that pivotal assortment of extinct shark-like fishes — are shifted, en masse, tthe branch containing the cartilaginous fishes. This triggers a cascade of implications. If all acanthodians are early cartilaginous fishes, then their shark-like features are not generalities of jawed vertebrates, but specializations of the cartilaginous-fish branch. The most recent common ancestor of jawed vertebrates was thus probably clad in bonarmor othe sort common to both placoderms anbony fishes. This inversion of a classic scenario in vertebrate evolution raises an obvious question: how did we get it so wrong?”

In summary
Even when someone gets it right, some of the details may still be correctable – and the present corrections do not overturn the conclusion, but support it. As usual, I have not seen the fossil firsthand. I have not added Entelognathus to the LRT. I simply make comparisons to published figures of Cheirolepis, which was one source of the earlier problems I had, no hopefully settled.

Thanks to David M.
for directing me to the Entelognathus paper. : – )

Please let me know
if someone else has drawn the same insight in the last 4 years since the publication of Zhu et al. 2013. If so, I am unaware of it.

References
Barford E 2013. Ancient fish face shows roots of modern jaw. Nature News. online here.
Friedman M and Brazeau 2013. A jaw-dropping fossil fish. Nature 502:175-177. online here.
Whiteaves JF 1881. On some remarkable fossil fishes from the Devonian rocks of Scaumenac Bay, in the Province of Quebec. Annals and Magazine of Natural History. 8: 159–162.
Zhu M, Yu X-B, Ahlberg PE, Choo B and 8 others 2013. A Silurian placoderm with osteichthyan-like marginal jaw bones. Nature. 502:188–193.

wiki/Cheirolepis
wiki/Entelognathus

Prorastomus/Pezosiren: when sirenians still had legs

Nothing heretical today.
We haven’t looked at any sirenians yet. And this one adds one more taxon to the LRT.

Figure 1. Prorastomus is a pro-sirenian with legs. All four feet remain unknown.

Figure 1. Prorastomus (or is this Pezosiren) is a pro-sirenian with legs. All four feet remain unknown. Elements from Pezosiren are also shown.

Prorastomus sirenoides (Owen 1855; Middle Eocene, 40 mya; 1.5m in length; Fig. 1) and Pezosiren are basal sirenians with four legs, a short tail and more teeth. They nest with the recenly extinct dugong, Dusisiren, in the large reptile tree (LRT, 1006 taxa).

Figure 2. Sirenian skulls, including Dusisiren, Prorastomus, and Eotheroides.

Figure 2. Sirenian skulls, including Dusisiren, Prorastomus, and Eotheroides. Note the loss of many teeth in Dusisiren.

Compared to its phylogenetic predecessor,
Moeritherium, Prorastomus/Pezosiren demonstrates the reduction in sacral vertebrae, the reduction in the cranial crest and the enlargement of the tail (what little is known). Pezosiren portelli (Domning 2001) is a related genus

According to Domning 2001
“Modern seacows (manatees and dugongs; Mammalia, Sirenia) are completely aquatic, with flipperlike forelimbs and no hindlimbs. Here I describe Eocene fossils from Jamaica that represent nearly the entire skeleton of a new genus and species of sirenian—the most primitive for which extensive postcranial remains are known. This animal was fully capable of locomotion on land, with four well-developed legs, a multivertebral sacrum, and a strong sacroiliac articulation that could support the weight of the body out of water as in land mammals. Aquatic adaptations show, however, that it probably spent most of its time in the water. Its intermediate form thus illustrates the evolutionary transition between terrestrial and aquatic life. Similar to contemporary primitive cetaceans3, it probably swam by spinal extension with simultaneous pelvic paddling, unlike later sirenians and cetaceans, which lost the hindlimbs and enlarged the tail to serve as the main propulsive organ. Together with fossils of later sirenians elsewhere in the world, these new specimens document one of the most marked examples of morphological evolution in the vertebrate fossil record.”

References
Domning DP 2001. The earliest known fully quadrupedal sirenians. Nature. 413 (6856): 625–627. online.
Owen R 1855.
 On the fossil skull of a mammal (Prorastomus sirenoïdes, Owen) from the island of Jamaica. The Quarterly Journal of the Geological Society of London 11:541-543.
Self-Sullivan C 2006. Evolution of the Sirenia.

wiki/Dusisiren
wiki/Prorastomus
wiki/Evolution_of_sirenians
wiki/Pezosiren

Liaoningosaurus: perhaps not an ankylosaur

This one goes back several years… with several updates!
Xu, Wang and You 2001 described what they thought was a juvenile ankylosaur, Liaoningosaurus paradoxes (Early Cretaceous, Yixian Formation) featuring “a large bony plate (somewhat shell-like) shielding the abdomen.” They tested it against only 13 other taxa and nested it outside the nodosaurs and outside the ankylosaurs…with no taxa between it and Stegosaurus.

Figure 1. Several specimens of Liaoningosaurus crushed flat plus a lateral view based on original holotype tracings.

Figure 1. Several specimens of Liaoningosaurus crushed flat plus a lateral view based on original holotype tracings.Note the lizard-like sprawling limbs in situ, a product of taphonomic crushing. Like all dinos, this one also had vertical limbs. Only a few small osteoderms are identified.

Xu et al. report:
“Diagnosis. An ankylosaurian that differs in having: shell-like ventral armour, trapezoidal sternum with slender and distally pointed posterolateral process and short medial articular margin, and pes more than twice as long as manus.”

Perhaps Xu et al. focused on ankylosaurs and nodosaurs
because all the specimens of Liaoningosaurus I have seen in publications or online (Fig. 1) have been crushed flat, with ribs spread out like ankylosaur ribs. Moreover, the pelvis was very wide, with limbs beneath the ilia, like those in ankylosaurs.

A closer look
(Fig. 1) reveals the ribs would have enclosed a deeper chest, not a wider one, though not as relatively deep as in Stegosaurus. Other more primitive stegosaurs likewise had a shorter, rounder torso cross-section.

Note,
the limbs are preserved sprawling, like those of the horned lizard, Phrynosoma. No dinosaur had sprawling limbs, so it’s okay to bring in both the limbs and the ribs (Fig. 1).

Finally,
basal stegosaurs also have a very broad pelvis with limbs rotating beneath the ilium. Considering how closely ankylosaurs and stegosaurs match each other in so many traits, it is a tribute to the LRT that it recovers them in separate clades, separated by bipedal agile taxa like Lesothosaurus and Heterodontosaurus. It is unlikely that ankylosaurs ever reared up on their hind limbs, but stegosaurs appear to be able to do this.

Osteoderms are rare in Liaoningosaurus,
which is odd for an armored ankylosaur.

Ankylosaur teeth and stegosaur teeth greatly resemble one another and also resemble Liaoningosaurus teeth (Fig. 1), despite the great difference in size.

There are five digits on the manus
in Liaoningosaurus (Fig. 1) and metacarpal #5 is as long as #4. Unfortunately, ankylosaurs and kin in the LRT lack a preserved manus, but a look through the Princeton Field Guide to Dinosaurs (Paul 2010) finds no similar ankylosaur manus.

Arbour et al. 2014 report, “Examination of the holotype of Liaoningosaurus paradoxus, IVPP V12566, indicates that the ventral “plastron” is better interpreted as epidermal scales, because the broken edges do not reveal any bony histology.” Readers will note that the odd ventral plate (closeup in Fig. 2) does not appear in other Liaoningosaurus specimens (Fig. 1), but they are exposed dorsally.

Figure 2. Liaoningosaurus ventral patch. Note the scales.

Figure 2. Liaoningosaurus ventral patch. Note the scales.

The large reptile tree (LRT, 1005 taxa) includes several more ornithischian taxa, though fewer taxa with armor. In the LRT Liaoningosaurus nests between Scutellosaurus and Stegosaurus, several nodes away from the other armored ornithischians, Minmi and Scelidosaurus. No skull traits were tested in Liaoningosaurus due to the low resolution of the available images.

The armored ornithischians (stegosaurs and ankyosaurs) are so similar
to one another they are traditionally nested in one clade: Thyreophora. By contrast, the LRT separates ankylosaurs from stegosaurs. Here the few hind limb traits that separate Liaoningosaurus from Scelidosaurus and/or Minmi and ally it with Scutellosaurus and/or Stegosaurus include the following:

  1. Tibia/femur ratio not less than 1:1
  2. Fibula not appressed to tibia
  3. Fibula diameter not > half tibia diameter
  4. Metatarsus not compact
  5. Metatarsal 1 < half metatarsal 3
  6. Metatarsal 1 not > half metatarsal 4
  7. Metatarsals 2 and 3 align beyond p1.1
  8. Pedal 4 length <  metatarsal 4

Perhaps better imagery
of the skull and other parts will add to or modify this list and nesting.

The addition of a basal ankylosaur
with these traits would nudge Liaoningosaurus toward the ankylosaurs. In the LRT ankylosaurs were derived from large, armored, lumbering Scelidosaurus. By contrast, the stegosaurs were derived from small, agile Lesothosaurus and Scutellosaurus. So finding a small armored dinosaur with the above list of traits, even if it is a juvenile, should suggest taking a close look at its stegosaur affinities, despite the initial appearance of a wide round horned-lizard-like torso.

PS
Ji et al. 2016 found fish within the torso (but not restricted to the gut) of a Liaoningosaurus suggesting a fish diet, rather than an herbivorous one.

PPS
Xu et al. 2001 reported, “Liaoningosaurus has an unusual combination of characters and it might (for example) represent a third ankylosaur lineage.” Perhaps one closer to stegosaurs. Xu et al. 2001 also report, “all manual and pedal unguals claw-shaped.” At present the manual unguals do not appear to be claw-shaped, with the the exception of #3, as in stegosaurs… AND as in ankylosaurs.

References
Xu X, Wang XL and You HL 2001. A juvenile ankylosaur from China. Naturwissenschaften 88:297. doi:10.1007/s001140100233
Ji Q, Wu X, Cheng Y, Ten F, Wang X and Ji Y 2016. Fish-hunting ankylosaurs (Dinosauria, Ornithischia) from the Cretaceous of China. Journal of Geology, 40(2).
Thompson RS, Parish JC, Maidment SCR and Barrett PM 2011. Phylogeny of the ankylosaurian dinosaurs (Ornithischia: Thyreophora). Journal of Systematic Palaeontology. 10 (2): 301–312. doi:10.1080/14772019.2011.569091
Arbour VM, Burns ME, Bell PR and Currie PJ 2014. Epidermal and dermal integumentary structures of ankylosaurian dinosaurs. Journal of Morphology, 275(1): 39-50. doi:10.1002/jmor.20194

wiki/Liaoningosaurus