Restoring the tip of the Pteranodon rostrum in UALVP 24238

Another short one today.
Everyone in pterosaur land has wondered what shape and length the missing tip of the rostrum took on in the UALVP 24238 specimen of Pteranodon (improperly called ‘Dawndraco‘, Fig. 1). That’s because, unlike most other specimens, the upper and lower margins don’t converge to form a triangle in lateral view, but remain essentially parallel for most of their length.

Figure 1. The missing tip of the UALVP 24238 skull can be restored like the similar Tanking-David specimen skull tip. The dorsal rim is straight. The ventral rim is convex.

Figure 1. The missing tip of the UALVP 24238 skull can be restored like the similar Tanking-Davis specimen skull tip. The dorsal rim is straight. The ventral rim is convex.

The privately held
Tanking-Davis specimen of Pteranodon is the most similar to the UALVP 24238 rostrum. Yes, it converges, but at a very shallower angle. Anteriorly the ventral margin curves up to meet the very straight dorsal margin. The same can be imagined/restored for the missing UALVP 24238 rostrum, and the result appears to be reasonable.

Figure 2. The UALVP 24238 specimen of Pteranodon is largely complete and here the tip of the rostrum is restored like that of a related specimen.

Figure 2. The UALVP 24238 specimen of Pteranodon is largely complete and here the tip of the rostrum is restored like that of a related specimen.

As in most pterosaur genera,
no two specimens are identical. See the panoply of known relatively complete Pteranodon skulls to scale and in phylogenetic order here.

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Flugsaurier 2018: Los Angeles County Museum

Flugsaurier
is a meeting of those interested in pterosaurs that happens in another part of the world every few years. I went to the first few. Saw a lot of specimens. Met a lot of colleagues. Produced a few abstracts and gave some presentations.

Over the next few days
there’s a Flugsaurier meeting taking place in Los Angeles. Many well-known and not-so-well known speakers are giving presentations this year. I will not be among them. Why?

So far as I know,
all of the conveners and many of the presenters continue to ignore a paper I wrote 18 years ago on the origin of pterosaurs from fenestrasaurs, not archosaurs. Other papers followed on wing shape, trackmaker identification and other topics, all supporting that phylogenetic hypothesis of relationships. Evidently workers would prefer to hope that pterosaurs arose from archosaurs close to dinosaurs. This is not where the data takes anyone interested in the topic who is not a party to taxon exclusion.

In addition, several of the conveners

  1. subscribe to the invalid quad-launch hypothesis
  2. the bat-wing reconstruction of the brachiopatagium.
  3. they believe that pedal digit 5 framed a uropatagium.
  4. They refuse to add tiny Solnhofen pterosaurs to their cladograms.
  5. They refuse to add several specimens of each purported genus to cladograms—and because of this they don’t recognize the four origins of the pterodactyloid-grade (not clade).
  6. They still don’t recognize that pterosaurs grew isometrically.
  7. They still don’t accept that pterosaur mothers retained their egg/embryo within the body until just before hatching (a lepidosaur trait).
  8. They still don’t accept that pterosaur bone fusion patterns follow lepidosaur, rather than archosaur patterns.
  9. They accept the idea that giant eyeballs filled the anterior skulls of anurognathids, not realizing that the supposed ‘scleral ring’ on edge of the flathead anurognathid is actually the mandible and tiny teeth.
  10. They reject any notion that all basal and some derived pterosaurs were bipedal, despite the footprint and morphological evidence proving bipedal locomotion.
  11. They all hold out hope that the largest azhdarchids could fly.
  12. I was going to say that all workers believe that crest size and hip shape identify gender, when the evidence indicates these are both phylogenetic markers, but then I found an abstract in 2018 that casts doubt on the gender/crest/pelvis hypothesis. So there’s hope.

That’s a fairly long list of ‘basics’
that most pterosaur workers ‘believe in’ despite the fact that there is no evidence for these false paradigms — but plenty of evidence for the lepidosaur origin of pterosaurs, from which most of the above hypotheses follow.

I am not attending Flugsaurier 2018
because the convening pterosaur workers deny and suppress the data listed above. Plus, I can more actively and thoroughly test assertions made during the conference from ‘my perch’ here in mid-America.

Good luck to those attending. 
Test all assertions and hypotheses, no matter their source.

Flugsaurier 2018: ‘Young istiodactylid’ nests with tall pterodactylids in the LPT

Flugsaurier 2018 opens today, August 10,
and the abstract booklet is out. So it’s time to take a look at some of the news coming out of that Los Angeles pterosaur symposium. Since the purpose of the symposium is increase understanding of pterosaurs, I hope this small contribution helps.

Figure 1. The Erlianhaote specimen attributed by Hone and Xu 2018 to istiodactylidae nests in the LPT with the large derived pterodactylids.

Figure 1. The Erlianhaote specimen attributed by Hone and Xu 2018 to the clade Istiodactylidae (within Ornithocheiridae) nests in the LPT with the large derived pterodactylids. Note the un-warped deltopectoral crest and lack of a deep cristospine, along with the long legs and short wings.

Hone and Xu at Flugsaurier 2018
describe, “An unusual and nearly complete young istiodactylid from the Yixian Formation, China (Fig. 1). The specimen shows the characteristic istiodactylid cranial features of tooth shape and enlarged nasoantorbital fenestra. However, it has proportionally large hindlimbs and wing proportions that are similar to those of azhdarchids. This has led to suggestion that the specimen may be a composite and that only the cranial material is istiodactylid. Preparation work around some key parts revealed no inconsistencies in the matrix or evidence of glue. The specimen is held in the Erlianhaote Dinosaur Museum, Erlianhote, China.”

Figure 2. The Erlianhaote specimen nests with these pterodactylids in the LPT, not with Istiodactylus (Fig. 3).

Figure 2. The Erlianhaote specimen nests with these pterodactylids in the LPT, not with Istiodactylus (Fig. 3). Note the antorbital fenestra becomes longer with larger size in this clade. The teeth are similar to those in istiodactylids.

Reconstructed as is
(Fig. 2) and added to the large pterosaur tree (LPT, 233 taxa, not yet updated due to no museum number nor genus name) the young ‘istiodactylid’ nests as a large derived pterodactylid. 13 steps separate this taxon from the Istiodactylus clade.

Ornithocheirids,
like Istiodactylus (Figs. 3, 4) and the SMNL PAL 1136 specimen (Fig. 5), share a very large wing finger, a short metacarpus, a warped deltopectoral crest, small free fingers and deeply keeled sternal complex not found in the Erlianhote specimen.

Figure 3. Istiodactylus has a shorter neck, longer wing finger and deep cristospine, among other traits not found in the new Erlianhaote specimen.

Figure 3. Istiodactylus has a shorter neck, longer wing finger and deep cristospine, among other traits not found in the new Erlianhaote specimen.

Figure 4. Istiodactylus sinensis is an istiodactylid from China sharing few traits with the new Erlianhaote specimen. Note the warped deltopectoral crest not warped in the new specimen.

Figure 4. Istiodactylus sinensis is an istiodactylid from China sharing few traits with the new Erlianhaote specimen. Note the warped deltopectoral crest not warped in the new specimen. Manual 4.1 is shorter than in other well-known istiodactylids.

The largest ornithocheirid

Figure 5. The unnamed largest ornithocheirid, SMNK PAL 1136, nests with Istiodactylus.

Figure 6. The Erlianhaote pterodactylid reconstructed in several views.

Figure 6. The Erlianhaote pterodactylid reconstructed in several views. The imagined (gray) areas of the skull here were imagined as an istiodactylid, but the better restoration is shown in figure 2.

It’s better not to eyeball certain specimens.
Sometimes you have to run them through a phylogenetic analysis to find out what they are. That’s what the LPT is for. It minimizes taxon exclusion and handles convergence.

Pterosaurs are still lepidosaurs.
So they follow lepidosaur fusion patterns, which follow phylogeny. Hone and Xu made the mistake of imagining pterosaurs might have archosaur fusion patterns that follow ontogeny.

Why am I not at Flugsaurier 2018?
In addition to about a dozen reasons that I can list later, or your can guess now, I can be more helpful and timely here.

References
Andres B and Ji Q 2006. A new species of Istiodactylus (Pterosauria, Pterodactyloidea) from the Lower Cretaceous of Liaoning, China. Journal of Vertebrate Paleontology, 26: 70-78.
Bowerbank JS 1846. On a new species of pterodactyl found in the Upper Chalk of Kent P. giganteus). Quarterly Journal of the Geological Society 2: 7–9.
Bowerbank JS 1851. On the pterodactyles of the Chalk Formation. Proceedings of the Zoological Society, London, pp. 14–20 and Annals of the Magazine of Natural History (2) 10: 372–378.
Bowerbank JS 1852. On the pterodactyles of the Chalk Formation. Reports from the British Association for the Advancement of Science (1851): 55.
Hone DWE and Xu 2018. An unusual and nearly complete young istiodactylid from the Yixian Formation, China. Flugsaurier 2018: the 6th International Symposium on Pterosaurs. Los Angeles, USA. Abstracts: 53–56.
Hooley RW 1913. On the skeleton of Ornithodesmus latidens. An ornithosaur from the Wealden shales of Atherfield (Isle of Wight)”, Quarterly Journal of the Geological Society, 69: 372-421
Howse SCB, Milner AR and Martill DM 2001. Pterosaurs. Pp. 324-335 in: Martill, D. M. and Naish, D., eds. Dinosaurs of the Isle of Wight, The Palaeontological Association
Wang X, Rodrigues T, Jiang S, Cheng X and Kellner AWA 2014. An Early Cretaceous pterosaur with an unusual mandibular crest from China and a potential novel feeding strategy. Scientific Reports 4 : 6329, pp. 1-9. | DOI: 10.1038/srep06329
Witton MP 2012. New Insights into the Skull of Istiodactylus latidens (Ornithocheiroidea, Pterodactyloidea). PLoS ONE 7(3): e33170. doi:10.1371/journal.pone.0033170

wiki/Istiodactylus

Juvenile Rhamphorhynchus and flightless pterosaur abstracts

Part 4
The following manuscripts are independently published online without peer-review at the DavidPetersStudio.com website. http://www.davidpetersstudio.com/papers.htm

Better to put them out there this way
than to let these works remain suppressed. Hope this helps clarify issues.


Peters D 2018g. First flightless pterosaur
PDF of manuscript and figures

Pterosaur fossils have been discovered all over the world, but so far no flightless pterosaurs have been reported. Here an old and rarely studied pterosaur fossil (Sos 2428) in the collection of the Jura Museum in Eichstätt, Germany, was re-examined and found to have a reduced pectoral girdle, small proximal wing elements (humerus, radius and ulna), three vestigial distal wing elements, the relatively longest pelvis of any pterosaur and the widest gastralia, or belly ribs. This discovery represents a unique morphology for pterosaurs. The Jura specimen lacked the wing size, forelimb muscularity and aerodynamic balance necessary to sustain flapping flight. It was a likely herbivore.


Peters D 2018h. First juvenile Rhamphorhynchus recovered by phylogenetic analysis
PDF of manuscript and figures
Standing seven to 44 centimeters in height, a growing list of 120+ specimens assigned to the pterosaur genus Rhamphorhynchus are known chiefly from the Solnhofen Limestone (Late Jurassic, southern Germany). An early study recognized five species and only one juvenile. A later study recognized only one species and more than 100 immature specimens. Phylogenetic analyses were not employed in either study. Workers have avoided adding small Solnhofen pterosaurs to phylogenetic analyses concerned that these morphologically distinct specimens were juveniles that would confound results. Here a large phylogenetic analysis that includes tiny Solnhofen pterosaurs tests that concern and seeks an understanding of relationships and ontogeny within the Pterosauria with a focus on Rhamphorhynchus. 195 pterosaurs were compiled with 185 traits in phylogenetic analysis. Campylognathoides + Nesodactylus were recovered as the proximal outgroups to the 25 Rhamphorhynchus specimens. The ten smallest of these nested at the clade base demonstrating phylogenetic miniaturization. Two Rhamphorhynchus had identical phylogenetic scores, the mid-sized NHMW 1998z0077/0001, and the much larger, BMNH 37002. These scores document a juvenile/adult relationship and demonstrate isometry during pterosaur ontogeny, as in the azhdarchid, Zhejiangopterus, and other pterosaurs. Rather than confounding results, tiny Solnhofen pterosaurs illuminate relationships. All descended from larger long-tailed forms and nested as transitional taxa at the bases of the four clades that produced all of the larger Late Jurassic and Cretaceous pterodactyloids. No long-tailed pterosaurs survived into the Cretaceous, so miniaturization was the key to pterosaur survival beyond the Jurassic.

These manuscripts benefit from
ongoing studies at the large reptile tree (LRT, 1256 taxa) in which taxon exclusion possibilities are minimized and all included taxa can trace their ancestry back to Devonian tetrapods.

Largest ‘flying reptile’ from the Crato formation? Maybe not.

Cheng et al. 2018
report on a partial wing finger (MPSC R 1221, Fig. 1) that they say represents, “The largest flying reptile from the Crato Formation, Lower Cretaceous, Brazil.”

But is it? 

Figure 1. The as yet undescribed SMNS PAL 1136 specimen is much larger than comparable bones in the new specimen, MPSC R 1221.

Figure 1. The as yet undescribed SMNS PAL 1136 specimen is much larger than comparable bones in the new specimen, MPSC R 1221. If the scale bars are correct, the SMNS specimen is much larger.

No…
if the scale bars are correct. The larger, as yet undescribed, and very impressive SMNS PAL 1136 specimen (Fig. 1) is not mentioned in the text. I do not know if the SMNS specimen is from the Crato or Roualdo formation (I have not gone back to look up that datum). In any case, the authors overlooked this specimen, because it is not mentioned in the text or charts that list a few dozen other large pterosaurs. It should have been included. Of course, then the headline would have read, “…second largest…” and no one wants that.

So was this oversight intentional?
We’ll never know. The SMNS specimen has been in the literature for 24 years (Frey and Martill 1994).

Addendum several days later
The Crato Formation was not erected until 13 years after the 1994 paper by Martill, Bechly and Loveridge. Therefore all layers were considered Santana Formation in 1994. So the SMNS specimen from the Santana formation might have come from the upper or lower layers. It should have been included in the 2018 survey.

The authors conclude
“Based on the fusion of the extensor tendon process and the first wing phalanx and bone histology, MPSC R 1221 presents a subadult individual of a late ontogeny stage (OS5) at time of death, whichmeans that the final maximized wingspan might have been larger. This is corroborated by the osteohistological sections since this individual did not present an external fundamental system.” Look how eager the authors are to hang on to that superlative, ‘largest’, even though we know of at least one that is so much larger.

The authors do not realize
or continue to deny data, that pterosaurs do not follow archosaur fusion patterns during ontogeny—because pterosaurs are not archosaurs, and their fusion patterns follow phylogenetic patterns.

I never heard the term,
“external fundamental system” before. So, I looked it up: “A closely spaced series of lines of arrested growth that is called the External Fundamental System (EFS) indicates that adult size has been reached.” Now we all know!

I hope this blog post
will one day turn out dozens of young paleontologists who will read every paper they see with a seasoned and skeptical eye. If so, a few of you may someday become editors of academic journals or manuscript referees. When that happens, don’t let mistakes like this slip out. Having a website, like ReptileEvolution.com, that is full of data and illustrations, makes it easy to fact-check superlative claims, like this one, with just a few clicks.

On that note:
here (Fig. 2) is a published illustration of a pterosaur wrist from Duque and Barret 2018 with labels that were a little mixed up with regard for the ulna and radius. The referees should have caught this.

Figure 1. From figure 9 from Duque and Barreto 2018 with corrections noted and digit 5 colorized

Figure 2. From figure from Duque and Barreto 2018 with corrections noted and digit 5 colorized. This mistake should have been caught by the authors and referees, not me.

References
Cheng X, Bantim RAM, Sayão JM, Kellner AWA, Wang X and Saraiva AAF 2018. The largest flying reptile from the Crato Formation, Lower Cretaceous, Brazil. Historical Biology. https://doi.org/10.1080/08912963.2018.1491567
Duque RRC and Barret AMF 2018. New exceptionally well-preserved Pterosauria from the lower Cretaceous Araripe Basin, Northeast Brazil. Cretaceous Research 10.1016/j.cretres.2018.05.004
Frey E and Martill DM 1994. A new Pterosaur from the Crato Formation (Lower Cretaceous, Aptian) of Brazil. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 194: 379–412.

Giant flying arboreal mammal-killer in the Jehol (Early Cretaceous, China)

So… this one has been under the radar since 2004
And you’ll see why.

Like a prehistoric eagle,
this was the largest flying predator in the Jehol biota (Early Cretaceous, China). It had no feathers. And it has gone unrecognized as a giant flying predator since Wang and Zhou 2004 announced it in Nature for other reasons.

At this time the only evidence
for this taxon comes in the form of a giant embryo anurognathid pterosaur, IVPP V13758 (Fig. 1) the size of other adult anurognathids. As an adult it would have been 8x larger (if similar to other pterosaur and based on the pelvic opening). The skull retains traits of the related, but more basal Dimorphodon from the Early Jurassic of England, but the giant anurognathid was coeval and similar in size to another Jehol predator, the pre-tyrannosauroid, Tianyuraptor, and larger than a coeval four-winged, flight-feathered ornitholestid, Microraptor (Fig. 1). It was also larger than the modern bald eagle (Haliaeetus leucocephalus). All the early Cretaceous toothed birds, like Yanornis, and Hongshanornis, were smaller.

Figure 1. Adult scaled version of the IVPP anurognathid pterosaur, with a skull similar in size to those attributed to Dimorphdodon. Bergamodactylus and Preondactylus are ancestral to Dimorphodon. Other Jehol predators are shown in white.

Figure 1. Adult scaled version of the IVPP anurognathid pterosaur, with a skull similar in size to those attributed to Dimorphdodon. Bergamodactylus and Preondactylus are ancestral to Dimorphodon. Other Jehol predators are shown in white.

If early Cretaceous mammals thought they were safe up in the trees,
think again. This giant anurognathid kept their numbers in check by going after them in the trees. That’s a big guess, but if you’re looking for a predator capable of snatching mammals out of the trees, there are no other candidates in the Early Cretaceous of China. Just look at those teeth!

Most anurognathids were small
because they ate small insect prey. Ask yourself if something as large as the IVPP embryo as an adult would have been satisfied eating insects. No, it was going after larger prey.

Figure 1. Large anurognathids and their typical-sized sisters. Here the IVPP embryo enlarged to adult size is larger than D. weintraubi and both are much larger than more typical basal anurognathids, Mesadactylus and MCSNB 8950.

Figure 2. Large anurognathids and their typical-sized sisters. Here the IVPP embryo enlarged to adult size is larger than D. weintraubi and both are much larger than more typical basal anurognathids, Mesadactylus and MCSNB 8950.

Unfortunately
Wang and Zhou 2004 (Fig. 3) didn’t know what sort of pterosaur their first embryo/egg was. Back then they thought pterosaur babies had a shorter rostrum that adults. Wrong. Back then they thought anurognathids were all small taxa. Wrong. Back then they didn’t spend much time tracing traits (Fig. 3) and reconstructions were largely guesswork. We fix all those problems here and at ReptileEvolution.com

The IVPP embryo pterosaur

Figure 3. Click to enlarge DGS tracing. The IVPP embryo pterosaur (far left) as originally traced, (near left) as originally reconstructed as a baby ornithocheirid, (near right) traced using the DGS method, (far right) adult reconstructed at 8x the embryo size.

We first looked at the IVPP embryo
here, several years ago and several times since.

Figure 4. The IVPP embryo anurognathid compared to other basal pterosaurs.

Figure 4. The IVPP embryo anurognathid enlarged to adult size and compared to other basal pterosaurs.

References
Wang X-L and Zhou Z 2004. Palaeontology: pterosaur embryo from the Early Cretaceous. Nature 429: 623.

http://reptileevolution.com/dimorphodon.htm
http://reptileevolution.com/ivpp-embryo.htm

https://pterosaurheresies.wordpress.com/2011/07/26/what-do-those-pterosaur-embryos-really-look-like/

The pterosaur tongue bone… and others, too

I did not give the pterosaur tongue much thought
until Li, Zhou and Clarke 2018 discussed it. They report, “Pterosaurs show convergent evolution of traits linked to tongue protrusion and mobility in birds (narrow midline element [achieved through fusion] and elongate, paired and rostrally positioned ceratobranchials).”

Figure 1. Scaphognathus (holotype) with hyoid traced in magenta.

Figure 1. Scaphognathus (holotype) with hyoid traced in magenta.

“We find pterosaurs similarly show lightly-built single pairs of ceratobranchial elements.

“Here, we bring together evidence from preserved hyoid elements from dinosaurs and outgroup archosaurs, including pterosaurs, with enhanced contrast x-ray computed tomography data from extant taxa.” (That means crocs and birds. Pterosaurs are not outgroup archosaurs, but fenestrasaur tritosaur lepidosaurs. Only taxon exclusion and academic suppression prevents this from being widely accepted.)

“The absence of direct and cranially-extensive support from bony elements make crocodilian tongue incapable of significant independent motion. Relative to outgroup lepidosaurs and other tetrapods the bony structure in crocodilians and surveyed basal archosaurs is uniformly simple and small with a single pair of ceratobranchials and no well-mineralized midline element or fusion.” 

“Elongation of hyobranchial elements co-occurs with increased ossification of a midline element (i.e., in paravians) and ceratobranchial fusion on the midline (i.e. pterosaurs). It is also well mineralized in some primarily quadrupedal, herbivorous ornithischians dinosaurs (e.g., ankylosaurids and hadrosauroids.) Within testudines, increase ossification of the midline element is seen in terrestrial taxa with an increase role for intraoral manipulation of food by the tongue.” Quoted verbatim. Ornithischia hyoids from Pinacosaurus are shown in figure 7.

Ludodactylus.

Figure 2. Click to enlarge. Ludodactylus, with rare pterosaur hyoid.

Li, Zhou and Clarke conclude
“In lepidosaurs, which show remarkable diversity in hyoid shape, there remains a primary respiratory function for the hyoid elements. The hyobranchial elements (multiple sets of ceratobranchials) show a primarily dorsoventral movement that is deployed during buccal pumping. The hyoid structure shows strong muscular links to the pectoral girdle that are lost in archosaurs and any tongue protrusion is via attached fleshy extensions rather than bony components. Perhaps not true in fenestrasaurs, which have a bird-like hyoid. See below.

“In Archosauria, the evolution of novel respiratory mechanisms apparently drove a simplification of the tongue that was retained in most taxa. Only with the evolution of flight (birds and pterosaurs) and in select quadrupedal herbivores was tongue structure elaborated.” By convergence, one must assume. Otherwise, what is the common thread?

Li, Zhou and Clarke did not realize
pterosaurs are lepidosaurs, not archosaurs. Anyone can test this by adding more relevant taxa to any relevant cladogram.

Figure 4. Sphenodon hyoids in two views.

Figure 3. Sphenodon hyoids in two views from Jones et al. 2009

Phylogenetic bracketing
using Sphenodon (Schwenk 1986) and Iguana (both with a fleshy tongue and posteriorly branching hyoids) should give tritosaur pterosaurs a fleshy tongue, too… except pterosaurs are highly derived and volant tritosaurs. Only the juvenile Huehuecuetzpalli, otherwise identical to the adult except in size, preserves hyoids. These are only visible posterior to the jaw. “According to their position, the anterior element was identified as the first ceratobranchial and the posterior element as the epihyal. The latter one, however, may be the hyoid cornu.” (Reynoso 1998). Tanystropheus and Macrocnemus have simple slender hyoids that approach one another anteriorly.

Figure 3. Longisquama in situ with hyoids identified.

Figure 4. Longisquama in situ with straight lateral and Y-shaped central hyoids identified.

Overlooked by Li, Zhou and Clarke,
a flightless lepidosaur and pterosaur outgroup taxon, Longisquama (Fig. 4) shares the pterosaur hyoid morphology. Sharovipteryx (Fig. 5) and Kyrgyzsaurus may as well. However in the lateral two taxa the lateral hyoids are so large they appear to have been able to spread laterally and so produce a cobra-like fleshy strake from otherwise loose neck skin and so introduce another aerodynamic membrane. Skull material prevents seeing a central hyoid if present.

Figure 4. Sharovipteryx in situ with hyoids identified. Note the expansive neck skin. Much has been said about the wasp-like insect in the left orbit, but Sharov was an insect collector first and there are many other insects, particularly beetles, all over the matrix.

Figure 5. Sharovipteryx in situ with hyoids identified. Note the expansive neck skin. Much has been said about the wasp-like insect in the left orbit, but Sharov was an insect collector first and there are many other insects, particularly beetles, all over the matrix.

Very few pterosaurs preserve hyoids.
The few that do have bird-like, y-shaped central hyoids (Figs. 1, 2) and sometimes straight lateral hyoids (Fig. 4), as Li, Zhou and Clarke correctly reported.

Yet another bunch of overlooked lepidosauriforms,

  1. Megalancosaurus has a Y-shaped central hyoid, unknown in other drepanosaurs.
  2. Stem chameleon (Early Cretaceous from amber) has a large central element split anteriorly
  3. Chlamydosaurus and kin use the hyoid elements for stretching the dermal skin, whether neck, like the Triassic Sharovipteryx (see above) or throat, as in the anole, Polychrus.
  4. Earliest reptile hyoids I have found: Diplovertebron (Fig. 6, a basal archosauromorph (in the LRT) from the Westphalian, but with its genesis in the Viséan.
Figure 3. Reconstruction of G. watsoni as a distinctly different genus, nesting with Eldeceeon rather than G. bohemicus.

Figure 6. Reconstruction of Diplovertebron (= Gephyrostegus watsoni) showing paired hyoids, the earliest I have seen on a reptile (Westphalian, Late Carboniferous, 300 mya)

Conclusions:
Hyoids vary greatly in size and shape in tetrapods. The basal/typical tetrapod tongue is boneless, fleshy and anchored by hyoids (see Sphenodon (Schwenk 1986), Caiman, Homo or Iguana). A few tongues are modified with an internal Y-shaped element for prey apprehension or reduced mobility, as in Melanerpes and Trioceros. This is found in various lepidosaurs (including pterosaurs) and birds by convergence. Mammals that use their tongues for prey/nectar apprehension do not have hyoids inside the tongue.

Figure 7. A selection of hyoids from Hill et al. 2015 with a focus on the ornithischian, Pinacosaurus.

Figure 7. A selection of hyoids from Hill et al. 2015 with a focus on the ornithischian, Pinacosaurus.

References
Hill RV, D’ Emic MD, Bever GS and Norell MA 2015. A complex hyobranchial apparatus in a Cretaceous dinosaur and the antiquity of avian paraglossalia. Zoological Journal of the Linnean Society 175: 892–909.
Jones et al. 2009. The head and neck muscles associated with feeding in Sphenodon (Reptilia: Lepidosauria: Rhynchocephalia). Palaeontologia Electronica 12(2).
Li Z, Zhou Z and Clarke JA 2018. Convergent evolution of a mobile bony tongue in flghted dinosaurs and pterosaurs. PLoS ONE 13(6):e0198078. https://doi.org/10.1371/journal.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.
Schwenk K 1986. Morphology of the tongue in the tuatara, Sphenodon punctatus (Reptilia: Lepidosauria), with comments on function and phylogeny. J Morphol. 1986; 188: 129–156. pone.0198078