Little Pulaosaurus nests with Changchunsaurus and Dryosaurus in the LRT
Yang, Kng nd Xu 2025 described
a little neornithischian from the Middle Jurassic, Pulausaurus (Figs 1–6, IVPP V30936) and nested it close to Agilisaurus (Fig 5) in two previously published cladograms of ornithischian dinosaurs.
The authors wrote, “A phylogenetic analysis places P. qinglong as one of the earliest-diverging neornithischians yet described.”
Figure 1. Pulaosaurus in situ. This is a complete, articulated specimen with shades of internal contents and soft tissue.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg?w=130″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg?w=444″ class=”size-full wp-image-93729″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg” alt=”Figure 1. Pulaosaurus in situ. This is a complete, articulated specimen with shades of internal contents and soft tissue. ” width=”584″ height=”1348″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg?w=584&h=1348 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg?w=65&h=150 65w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg?w=130&h=300 130w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.insitu588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 1. Pulaosaurus in situ. This is a complete, articulated specimen with shades of internal contents and soft tissue.
Yang, King and Xu wrote,
“The first analysis [based on Han et al 2018) produced 26,901 most parsimonious trees.”
So, not much resolution in that cladogram, which excludes sauoropods and other basal phytodinosaurs (Fig 7), but inappropriately. cherry picks Silesaurus. There Pulausaurus nested within the base of the clade Ornithischia = close to the outgroup taxa.
Figure 2. Skull of Pulaosaurus insitu and reconstructed major elements.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg?w=216″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg?w=584″ class=”size-full wp-image-93738″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg” alt=”Figure 2. Skull of Pulaosaurus insitu and reconstructed major elements.” width=”584″ height=”810″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg?w=584&h=810 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg?w=108&h=150 108w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg?w=216&h=300 216w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.skull588-1.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 2. Skull of Pulaosaurus insitu and reconstructed major elements.
Yang, King and Xu wrote,
“The reduced consensus tree is from 572 of the most parsimonious trees.”
That analysis nested Pulausaurus with Agilisaurus, not as close to the base of the Ornithischia.
Figure 3. Pulaosarus manus with PILs. Those manual unguals are still sharp.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif?w=195″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif?w=584″ class=”size-full wp-image-93733″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif” alt=”Figure 3. Pulaosarus manus with PILs. Those manual unguals are still sharp.” width=”584″ height=”897″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif?w=584&h=897 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif?w=98&h=150 98w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif?w=195&h=300 195w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus.manus588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 3. Pulaosarus manus with PILs. Those manual unguals are still sharp.
Yang, King and Xu wrote,
“The second phylogenetic analysis with weight equal based on Fonseca et al. (2024)
dataset produces 10,000 most parsimonious trees.”
In this second analysis more taxa were tested and Pulausaurus nested with Agilisaurus again.
Editors note: More taxa are not helpful if they are based on scraps and if valid outgroup taxa are ignored.
Figure 4. Pes of Pulaosaurus with PILs added here.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif?w=170″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif?w=580″ class=”size-full wp-image-93735″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif” alt=”Figure 4. Pes of Pulaosaurus with PILs added here. ” width=”584″ height=”1031″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif?w=584&h=1031 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif?w=85&h=150 85w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif?w=170&h=300 170w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/pulaosaurus_pes588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 4. Pes of Pulaosaurus with PILs added here. Digits 1 and 5 are essentially fused to the metatarsals 2 and 4, so are easy to overlook.
Timeout: Which taxa belong in Neornithischia?
Cooper 1985 defined this clade as “all genasaurians more closely related to Parasaurolophus than to Ankylosaurus or Stegosaurus .”
In the LRT (Fig 7) Neornithischia includes Ornithopoda + Ceratosauria. So the LRT recovers this same clade and Pulaosaurus is a member. Agilisaurus is not.
Sereon 1986 named the same clade Cerapoda, which was preoccupied by a moth clade and arrived a year too late. Nevertheless, “In 2021, Cerapoda was given a formal definition under the PhyloCode: “The smallest clade containing Iguanodon bernissartensis Boulenger in Beneden, 1881, Pachycephalosaurus wyomingensis (Gilmore, 1931), and Triceratops horridus Marsh, 1889.
In the LRT (Fig 7) that clade also includes stegosaurs and anklyosaurs,
which was not the intention. Those clades were intended to be excluded.
Baron, Norman and Butler 2017 reported that Lesothosaurus was the most basal member of Neornithischia.
In the LRT (Fig 7) Lesothosaurus is the proximal outgroup taxon, basal to stegosaurs.
Problem is
some workers link pachycephalosaurs with ceratopsia.
In the LRT (Fig 7) pachycephalosaurs nest with heterodontosaurs outside the Neornithischia.
Fonseca et al 2024
in their ‘comprehensive phylogenetic analysis on early ornithischian evolution’ added Silesaurus to the Ornithischia, ignoring Chilesaurus, Daemonosaurus and Sauropodomorpha as basal and outgroup taxa respectively.
So, this was not a ‘comprehensive phylogenetic analysis on early ornithischian evolution’ due to taxon exclusion among outgroup taxa.
The LRT (Fig 7) minimizes taxon exclusion. So there is not cherry-picking.
Figure 5. Agilisaurus and Pulausaurus to scale.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg?w=300″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg?w=584″ class=”size-full wp-image-93740″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg” alt=”Figure 5. Agilisaurus and Pulausaurus to scale. ” width=”584″ height=”308″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg?w=584&h=308 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg?w=150&h=79 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg?w=300&h=158 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/agilisaurus-overall.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 5. Agilisaurus and Pulausaurus to scale.
Here,
in the large reptile tree (LRT, 2339 taxa, Fig 7) Pulausaurus nests with larger Early Cretaceous Changchunsaurus and Late Jurassic Dryosaurus (Fig 5).
Getting the tree topology correct should be the number one goal of vertebrate paleontology.
I know of not other workers who are testing as many vertebrate taxa from such a wide gamut in phenomic = trait-based analysis. Such a study needs to be replicated several times and ultimately accepted in consensus before more focused studies continue to support myths promoted by cherry-picking outgroup taxa and omitting pertinent taxa.
Figure 6. Pulausaurus compared to scale with Dryosaurus and Changchunsaurus.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg?w=239″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg?w=584″ class=”size-full wp-image-93742″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg” alt=”Figure 6. Pulausaurus compared to scale with Dryosaurus and Changchunsaurus. ” width=”584″ height=”732″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg?w=584&h=732 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg?w=120&h=150 120w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg?w=239&h=300 239w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/changchunsaurus-588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 6. Pulausaurus compared to scale with Dryosaurus and Changchunsaurus.
The LRT minimizes taxon exclusion
reducing the need to trust tradition and authority that cherry-picks outgroup taxa.
Figure 6. Subset of the LRT focusing on the Phytodinosauria with the addition of Pulausaurus basal to Ornithopoda.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg?w=143″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg?w=488″ class=”size-full wp-image-93744″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg” alt=”Figure 6. Subset of the LRT focusing on the Phytodinosauria with the addition of Pulausaurus basal to Ornithopoda. ” width=”584″ height=”1227″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg?w=584&h=1227 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg?w=71&h=150 71w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg?w=143&h=300 143w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/phytodinosauria2025.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 7. Subset of the LRT focusing on the Phytodinosauria with the addition of Pulausaurus basal to Ornithopoda. Agilisaurus nests here with Stegoceras and Heterodontosaurus.
The LRT appears to present a novel hypothesis of interrelationships.
If not, please provide a citation so I can promote it here
References
Baron MG, Norman DB and Barrett PM 2016. Postcranial anatomy of Lesothosaurus diagnosticus (Dinosauria: Ornithischia) from the Lower Jurassic of southern Africa: implications for basal ornithischian taxonomy and systematics. Zoological Journal of the Linnean Society. doi:10.1111/zoj.12434
Butler RJ, Upchurch P and Norman DB 2008. The phylogeny of the ornithischian dinosaurs. Journal of Systematic Palaeontology, 6(1): 1–40.
Cooper MR 1985. A revision of the ornithischian dinosaur Kangnasaurus coetzeei Haughton, with a classification of the Ornithischia. Annals of the South African Museum, 95: 281-317.
Fonseca AO et al (5 co-authors) 2024. A comprehensive phylogenetic analysis on early ornithischian evolution. Journal of Systematic Palaeontology. 22 (1): 2346577.
Yang Y, King JL and Xu X 2025. A new neornithischian dinosaur from the Upper Jurassic Tiaojishan Formation of northern China. PeerJ 13:e19664 http://doi.org/10.7717/peerj.19664
wiki/Pulaosaurus
wiki/Neornithischia
Source: https://pterosaurheresies.wordpress.com/2025/07/19/little-pulaosaurus-nests-with-changchunsaurus-and-dryosaurus-in-the-lrt/
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