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

Advertisements

New phylogeny of Stegosauria

A few problems here.
Raven and Maidment 2017 have produced a phylogeny of the clade Stegosauria (Fig. 1). Unfortunately it splits stegosaur proximal outgroups (in the large reptile tree (LRT, subset in Fig. 2) from stegosaurs. It splits stem or basal ankylosaurs from derived ankylosaurs. And it supports a clade, the Thyreophora, that was found to be paraphyletic in the LRT. Finally, it nests Laquintasaurus with Scutellosaurus, contra the LRT.

Figure 1. Phylogeny of Stegosauria according to Ravena and Maidment 2017. Yellow/green taxa are stegosaurs and their ancestors in the LRT. Gray taxa are nodosaurs and ankylosaurs. Blue taxon is a basal ceratopsian. Magenta taxon is lost. The LRT nests stegosaurs apart from ankylosaurs, thus the Thyreophora is paraphyletic and invalid.

Figure 1. Phylogeny of Stegosauria according to Ravena and Maidment 2017. Yellow/green taxa are stegosaurs and their ancestors in the LRT. Gray taxa are nodosaurs and ankylosaurs. Blue taxon is a basal ceratopsian. Magenta taxon is lost. The LRT nests stegosaurs apart from ankylosaurs, thus the Thyreophora is paraphyletic and invalid.

Raven and Maidment appear to have chosen outgroups
for Stegosauria instead of letting a larger gamut analysis choose them. So, once again, taxon exclusion lessens the effectiveness of and confidence in a hypothesis.

Figure 2. Phytodinosauria with a focus on Stegosauria (yellow green).

Figure 2. Subset of the LRT: Phytodinosauria with a focus on Stegosauria (yellow green).

References
Raven TJ and Maidment SCR 2017. A new phylogeny of Stegosauria (Dinosauria, Ornithischia). Palaeontology 2017:1–8.
Barrett PM, Butler RJ, Mundil R, Scheyer TM, Irmis RB, Sánchez-Villagra MR (2014) A palaeoequatorial Ornithischian and new constraints on early dinosaur diversification. Proceedings of the Royal Society B 281(1791): 20141147. http://dx.doi.org/10.1098/rspb.2014.1147

The skull of Lesothosaurus revisited

Lesothosaurus diagnostics (Galton 1978, Early Jurassic, 2.2m) was originally considered a basal ornithopod. Sereno (1991) suggested it may be one of the most primitive of all ornithischian dinosaurs. Butler et al. (2008) proposed that Lesothosaurus was a basal member of Neornithischia (pachycephalosaurs, ceratopsians and ornithopods), or Thyreophora (stegosaurs + ankylosaurs). The large reptile tree nests Lesothosaurus with quadrupedal Scutellosaurus and stegosaurs, not ankylosaurs. Galton, Sereno, Butler et al. published before the discoveries of more basal ornithischians like Daemonosaurus and Chilesaurus, but Wikipeidia needs another excuse for these deletions.

Just slight differences here (Fig. 1)
from traditional imagery and interpretations, chiefly those of Sereno 1991.

Figure 1. The skull of Lesothosaurus (BMNH 8501) traced and reconstructed.

Figure 1. The skull of Lesothosaurus (BMNH 8501) traced and reconstructed. Lower teeth imagined.

This new interpretation clears up
some of the problems in the large reptile tree (now 694 taxa) that were preventing high bootstrap scores at certain nodes. A bit of posterior palpebral appears above the fused postorbital/postfrontal. You also find that in Stegosaurus. The naris is close to the ventral rim, atypical for basal phytodinosaurs and basal ornithischians, but convergently appearing in certain derived ornithischians. Likewise a toothless premaxilla developed several times in the Ornithischia.

References
Butler RJ, Upchurch P, Norman, DB 2008. The phylogeny of the ornithischian dinosaurs. Journal of Systematic Palaeontology 6 (1): 1–40. doi:10.1017/S1477201907002271.
Galton PM 1978. Fabrosauridae, the basal family of ornithischian dinosaurs (Reptilia:
Ornithopoda). Paläontolgische. Zeitschrift 52:138–59.
Knoll F, Padian K and de Ricqles A 2009. Ontogenetic change and adult body size of the early ornithischian dinosaur Lesothosaurus diagnosticus: implications for basal ornithischian taxonomy”. Gondwana Research online preprint: 171. doi:10.1016/j.gr.2009.03.010.
Sereno PC 1991. Lesothosaurus, “fabrosaurids,” and the early evolution of Ornithischia. Journal of Vertebrate Paleontology 11(2):168-197

wiki/Lesothosaurus