Mirasaura grauvogeli: a new fenestrasaur now closer to Oculudentavis
Yesterday we celebrated the publication and description
of Mirasaura grauvogeli, a Middle Triassic pre-pterosaur = fenestrasaur = miniature tanystropheid tritosaur lepidosaur miscast as a drepanosauromorph (Spiekman et al 2025) due to taxon and citation exclusion.
Drepanosauromorpha is the sister clade to the Huehuecuetzpalli through Quetzalcoatlus clade = Tritosauria in the large reptile tree (LRT, 2339 taxa, subset Fig 1). So not far off.
Figure 1. Subset of the LRT focusing on the Tritosauria, including Drepanosauromorpha and Tanystropheidae.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg?w=177″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg?w=584″ class=”size-full wp-image-93849″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg” alt=”Figure 1. Subset of the LRT focusing on the Tritosauria, including Drepanosauromorpha and Tanystropheidae. ” width=”584″ height=”990″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg?w=584&h=990 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg?w=88&h=150 88w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg?w=177&h=300 177w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/tritosauria2025cladogram588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 1. Subset of the LRT focusing on the Tritosauria, including Drepanosauromorpha and Tanystropheidae, now with the inclusion of Mirasaura among the basal Fenestrasauria.
Today we’ll nibble at some of the crumbs left behind.
The authors wrote, “The early evolution of amniote integumentary appendages, however, remains poorly understood because of the absence of fossil evidence.”
Rarity? Yes. Absence? No.
The authors did not include Oculudentavis, Sharovipteryx and Cosesaurus (Fig 2) in their text or cladogram. Longisquama (Fig 2) is mentioned in text and thus breaks their rule of “absence”.
Phylogenetically both Iguana and Sphenodon are as early = ancient, yet their homologous dorsal frills were ignored by the authors.
Figure 1. Longisquama and Cosesaurus compared to the most complete specimen of Mirasaura after tracing elements using DGS colors.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg?w=300″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg?w=584″ class=”size-full wp-image-93801″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg” alt=”Figure 1. Longisquama and Cosesaurus compared to the most complete specimen of Mirasaura after tracing elements using DGS colors.” width=”584″ height=”444″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg?w=584&h=444 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg?w=150&h=114 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg?w=300&h=228 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/longisquama-reconstruction588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 2. Longisquama and Cosesaurus compared to the most complete specimen of Mirasaura after tracing elements using DGS colors.
“Despite its superficially bird-like skull, Mirasaura is not closely related to avemetatarsalians but instead belongs to the exclusively Triassic reptilian clade Drepanosauromorpha.”
‘Avemetatarsalia’ includes dinos and pteros, which had their last common ancestor in the LRT with the last common ancestor of all reptiles (amniote is a junior synonym), Silvanerpeton. That makes ‘Avemetatarsalia’ a junior synonym for Reptilia when more taxa are added to analysis, as in the LRT.
Cosesaurus also had a ‘superficially bird-like skull’ to such an extent that paleontologist Ellenberger 1993 considered it a proto-bird without considering its pterosaurian homologs. The same mistake was made by the more recently described Oculudentavis, now widely acknowledged as a lepidosaur. The LRT nests it with Cosesaurus and kin.
“the morphology of the integumentary appendages and phylogenetic placement of Mirasaura indicate that they are not structurally homologous to feathers or other integumentary appendages in living amniotes.”
Those plumes are great elongated homologs to the single-row dorsal frills = spines
of the lepidosaurs, Iguana (Fig 3) and Sphenodon.
Figure 1. The spines of Iguana.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg?w=300″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg?w=584″ class=”size-full wp-image-5056″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg” alt=”The spines of Iguana.” width=”584″ height=”339″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg?w=584&h=339 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg?w=150&h=87 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg?w=300&h=174 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/03/588px-cuban_iguana.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 3. The spines of Iguana.
“Our findings show that complex integumentary appendages are not restricted to avemetatarsalians and mammaliaforms among amniotes and evolved in a lineage basal to all extant reptiles,”
Silvanerpeton, the taxon had scales, but no integumentary appendages.
“Until now, the only fossil evidence for complex integumentary appendages in reptiles outside Avemetatarsalia comes from Longisquama insignis, a small diapsid reptile from the Middle or Late Triassic epoch of Kyrgyzstan.”
Cosesaurus was earlier. Sharovipteryx was a coeval.
Ignoring published data is one way to make your own discovery that much more exciting. But then you run the risk. Longisquama was nested with these taxa among the small, fenestrasaur, tanystropheid, tritosaur lepidosaurs earlier (Peters 2000).
Though they share a diapsid skull architecture with diapsids like Petrolacosaurus and its desendants. By contrast, diapsid lepidosaurs are not related except through Silvanerpeton the last common ancestor of Lepidosauromorpha and Archosauromorpha.
Figure 4. DGS tracing of the Mirasaura holotype with the best dorsal frill. Note the lack of overlap, as in Longisquama, Sphenodon and Iguana.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif?w=130″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif?w=443″ class=”size-full wp-image-93805″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif” alt=”Figure 4. DGS tracing of the Mirasaura holotype with the best dorsal frill. Note the lack of overlap, as in Longisquama, Sphenodon and Iguana.” width=”584″ height=”1351″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif?w=584&h=1351 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif?w=65&h=150 65w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif?w=130&h=300 130w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura-plumes588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 4. DGS tracing of the Mirasaura holotype with the best dorsal frill. Note the lack of overlap, as in Longisquama, Sphenodon and Iguana.
“Investigation of the integumentary appendages shows a distally expanded bilateral morphology and preserved melanosomes that are similar in geometry to melanosomes in feathers.”
Not bilateral. Only a single row of integumentary appendages is present (Fig 4), as in extant Iguana (Fig 3) and Sphenodon along with extinct Longisquama and Cosesaurus (Fig 2).
“The absence of branching in the appendages of the new taxon, however, indicates that these bilateral structures formed independently of avemetatarsalian feathers.”
Feathers? No. Spines? Yes. See Iguana photo above (Fig 3).
This identification is not news (Peters 2000, 2018).
“comparisons of the skeletal and integumentary anatomy between the new species and Longisquama show that both are closely related taxa.”
Oddly Longisquama is never figured in this paper. Conspicuous by its absence.
Figure 5. The skulls of Mirasaura, Longisquama, Oculudentavis to scale. Bergamodactylus, the basalmost pterosaur, is half size here.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg?w=124″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg?w=424″ class=”size-full wp-image-93811″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg” alt=”Figure 5. The skulls of Mirasaura, Longisquama, Oculudentavis to scale. Bergamodactylus, the basalmost pterosaur, is half size here.” width=”584″ height=”1409″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg?w=584&h=1409 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg?w=62&h=150 62w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg?w=124&h=300 124w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.skull588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 5. The skulls of Mirasaura, Longisquama, Oculudentavis to scale. Bergamodactylus, the basalmost pterosaur, is half size here. Note the curled quadrate (red) in Oculudentavis and Cosesaurus. The quadrate is straight in pterosaurs, Longisquama and Mirasaura.
Diagnosis
“Mirasaura grauvogeli is a drepanosauromorph [not true] distinguished by the following combination of character states (autapomorphies among non-saurian diapsids are marked with an asterisk*): elongate integumentary appendages extending dorsally from the anterior part of the trunk [as in Cosesaurus, and some pterosaurs like Scaphognathus and Jeholopterus were soft tissue is preserved]; long, narrow snout [as in pterosaurs and Oculudentavis]; dome-like skull roof [as in pterosaurs and Oculudentavis]; anterior portions of the jaws completely edentulous [yes, but note how tiny and slender the jaw tips were]*; dorsoventrally low maxilla [as in Oculudentavis]; frontal strongly constricted anteriorly [as in Oculudentavis]; forward-facing orbits [exaggerated, actually about 35º from parasagittal]; transversely wide parietal [as in Oculudentavis]; elongate, barrel-shaped trunk [as in Longisquama]; seven cervical vertebrae [I count nine, fig x, as in pterosaurs and kin]; 24 dorsal vertebrae [that number may change given the presence of 9 cervicals and 5+ sacrals]*; hypapophyses on cervical vertebrae [those are cervical ribs]; cervical ribs and gastralia absent [actually both present]; moderately elongate caudal haemal spines; narrow and elongate scapular blade [a displaced trachea = windpipe]; carpals lacking proximal elongation [the plesiomorphic condition]; ilium with tall, anterodorsally oriented blade [actually anteriorly-oriented framiing 5+ sacrals]; and large, curved pedal unguals [plesiomorphic].”
Figure 2. CT scans of Oculudentavis from Xing et al. 2020 and colored here, plus a comparison of Cosesaurus to scale.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg?w=191″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg?w=584″ class=”size-full wp-image-43506″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg” alt=”Figure 2. CT scans of Oculudentavis from Xing et al. 2020 and colored here, plus a comparison of Cosesaurus to scale.” width=”584″ height=”917″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg?w=584&h=917 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg?w=96&h=150 96w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg?w=191&h=300 191w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2020/03/oculudentavis-khaungraae4views588-1.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 6. CT scans of Oculudentavis from Xing et al. 2020 and colored here, plus a comparison of Cosesaurus to scale.
The authors wrote,
“We incorporated Mirasaura into the most detailed available character–taxon matrix for drepanosauromorphs, closely related diapsid taxa and early members of lepidosauromorpha and Archosauromorpha in both relevant taxon and character sampling.” (Pritchard, Sues, Scott and Reisz 2021).
Unfortunately, that paper also suffered from taxon exclusion. See link below.
Figure x. Mirasaura cervicals. More than 7 are present here.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif?w=300″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif?w=584″ class=”size-full wp-image-93837″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif” alt=”Figure x. Mirasaura cervicals. More than 7 are present here. ” width=”584″ height=”360″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif?w=584&h=360 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif?w=150&h=92 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif?w=300&h=185 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.cervicals588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 7. Mirasaura cervicals. More than 7 are present here.
Mirasaura is a tiny taxon.
So bones like gastralia (Fig 8) and other slender elements are represented by impressions at best.
Figure 7. Mirasaura gastralia and other elements.
” data-medium-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif?w=300″ data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif?w=584″ class=”size-full wp-image-93840″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif” alt=”Figure 7. Mirasaura gastralia and other elements.” width=”584″ height=”289″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif?w=584&h=289 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif?w=150&h=74 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif?w=300&h=148 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2025/07/mirasaura.gastralia588.gif 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Figure 8. Mirasaura gastralia and other elements impressions.
Mirasaura
adds to the list of pterosaur ancestors that have gone unrecognized as pterosaur ancestors by members of Academia – always due to taxon and citation exclusion. It’s almost as if they are avoiding adding taxa that were added twenty-five years ago in Peters 2000 and updated in Peters 2018.
Fortunately, taxon exclusion is easily remedied by simply adding taxa.
Not sure why academics have resisted doing that simple exercise.
References
Ellenberger P 1993. Cosesaurus aviceps. Vertébré aviforme du Trias Moyen de Catalogne. Étude descriptive et comparative. Mémoire Avec le concours de l’École Pratique des Hautes Etudes. Laboratorie de Paléontologie des Vertébrés. Univ. Sci. Tech. Languedoc, Montpellier (France). Pp. 1-664.
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2018 unpublished. Cosesaurus aviceps, Sharovipteryx mirabilis and Longisquama insignis Reinterpreted. ResearchGate.net
Pritchard AC, Sues H-D, Scott D and Reisz RR 2021. Osteology, relationships and functional morphology of Weigeltisaurus jaekeli (Diapsida, Weigeltisauridae) based on a completeskeleton from the Upper Permian Kupferschiefer of Germany. PeerJ 9, e11413 (2021).
Reisz RR and Sues H-D 2000. The ‘feathers’ of Longisquama. Nature 408, 428–428.
Spiekman SNF et al (12 co-authors) 2025. Triassic diapsid shows early diversification of skin apprendages in reptiles. Nature https://doi.org/10.1038/s41586-025-09167-9
Publicity
Mirasaurua in sciencealert.com
Source: https://pterosaurheresies.wordpress.com/2025/07/25/mirasaura-grauvogeli-a-new-fenestrasaur-now-closer-to-oculudentavis/
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