SVP abstracts 27: Plesiosaur breathing

Wintrich and Vanoefer 2020
bring us a look at plesiosaur breathing, but do not consider the vertical feeding configuration (Fig. 1) and bubble-net blowing hypothesis.

Figure 3. Click to enlarge. Albertonectes reconstructed. This 11 m elasmosaur is the longest thusfar recorded. This may be the breathing pose, swallowing air, then submerging the neck. When horizontal the air could be passed back to the lungs, as hypothesized for Dinocephalosaurus.

Figure 3. Click to enlarge. Albertonectes reconstructed. This 11 m elasmosaur is the longest thusfar recorded. This may be the breathing pose, swallowing air, then submerging the neck. When horizontal the air could be passed back to the lungs, as hypothesized for Dinocephalosaurus.

From the Wintrich and Vanoefer 2020 abstract:
“Plesiosaurs are enigmatic marine reptiles known from the Late Triassic to the Late Cretaceous and represent the most derived group of sauropterygians.

Why ‘enigmatic’? Plesiosaurs are readily identified without argument.

“Among plesiosaurs, there are several lineages showing an extremely long neck, which raises different biomechanical questions dealing with use and function, up to the breathing mechanism. Furthermore, for aquatic tetrapods, buoyancy control is an important adaptation to support the body in the water column. The respiratory system and its influence on buoyancy control have been discussed only briefly, and no mathematical approach has been taken so far. However, the breathing mechanism and therefore the respiratory system of highly aquatic tetrapods has to be specialized in different ways to enable life in a pelagic environment.”

“Here, we follow different mathematical approaches based on the metabolism (work of breathing), the trachea, and the morphology of the skull and trunk, in order to reconstruct the breathing mechanism, respiratory system, and lung volume in plesiosaurs, and then discuss the most plausible respiratory anatomy.”

“Furthermore, we find support for the hypothesis of a functional secondary palate from the reconstructed respiratory system as well as for the use of gastroliths, especially in the Elasmosauridae.”

“In addition to this, we calculated the center of mass to reconstruct buoyancy control in plesiosaurs.”

“In general, we studied four different long-necked plesiosaurs (Cryptoclidus, Albertonectes, Rhaeticosaurus, Rhomaleosaurus) and included Augustasaurus as the most derived pistosaur for which the entire neck is known.”

“Our results demonstrate that the lung volume was larger than suspected for an aquatic tetrapod. However, plesiosaurs showed an adaption similar to that of marine turtles, which have shifted the lung to the dorsal side of the trunk. The influence of the long trachea on breathing is not as great as suggested before. However, especially in the elasmosaurid, the long neck influences the center of mass. This supports the hypothesis of gastroliths functioning in buoyancy control in elasmosaurs. Furthermore, based on an ancestral state reconstruction, we show that the specialized plesiosaurian respiratory systems probably evolved in early sauropterygians.”

‘Influences’. ‘Supports’. ‘Probably’. Conclusions?
I didn’t see any here. Did I miss something? Seems like this is all old news. Earlier we looked at the possibility that plesiosaurs were vertical hunters (Fig. 1), expressing bubble nets as they rose beneath fish schools, as in modern mysticetes. Let’s see that ‘specialized’ hypothesis tested in the 2021 abstracts.

Wintrich T and Vanhoefer J 2020. A specialized respiratory system in plesiosaurs (Sauropterygia): breathing with the long neck. SVP abstracts.

Flugsaurier 2018: Los Angeles County Museum

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.

Something new in Eudimorphodon revealed by DGS

Some people are still having trouble with DGS as a technique. They think of it as something that is virtually guaranteed to spook a reconstruction. Instead of increasing confidence that parts have been correctly identified, they have no confidence in work that has the taint of DGS.

Here’s a step-by-step run through DGS on a familiar specimen, Eudimorphodon ranzii. Using DGS enabled the recognition of some oddly long posterior ribs (that were always visible, just ignored) and a wider than deep torso in a pterosaur for which these traits were not otherwise recorded.

Eudimorphdon ranzii (Zambelli 1973, Wild 1978) s a Late Triassic pterosaur known from an articulated crushed skeleton missing feet, tail and most of each wing (Figs. 1-3). Some parts are easy to see and trace, like the skull and sternal complex. Some parts are more difficult like the two pubes (Wild 1978 only found one by combining the two into an oddly broad prepubis),  the pelvis, and the odd arrangement of the posterior ribs.

Eudimorphdon ranzii with post cranial bones colorized.

Figure 1. Eudimorphdon ranzii with post cranial bones colorized.

Step one: Colorize the bones (Fig. 1)
Darren Naish seems to think this is okay if you know which bone is which ahead of time when looking at the specimen and you’re just making a visual presentation. I like to take it one step further and use DGS to segregate bones that are more difficult to identify. Here the pelvis is found. The dorsal ribs will precisely transferred to the reconstruction, not generically applied. As we’ve learned earlier, sometimes pterosaurs have the cross section of a horned lizard.

Figure 2. The colorized bones on a fresh canvas.

Figure 2. The colorized bones on a fresh canvas. Most tetrapods have shorter posterior dorsal ribs, but not here in Eudimorphodon. Lighter tones on the pelvis represent overlying bones, in this case vertebrae. It is important to put a numeral on each vert and rib because it is otherwise easy to become confused.

Step two: Transfer the colorized bones onto a fresh white background (Fig. 2)
Here we’re just trying to put the bones on a fresh canvas. You’ll note some bones are estimates based on vague clues as they appear beneath the sternal complex.

Figure 3. Moving colorized bones into a rough reconstruction.

Figure 3. Moving colorized bones into a rough reconstruction or Eudimorphodon. Here both pelves are shown as they appeared in situ. In figure 1 I jumped the gun and put the parts together.

Step three: Move the colorized bones into a rough assembly (Fig. 3)
Here we’re just trying estimate a body shape to make tracing the colored bones easier.

Figure 4. Lateral, dorsal and cross-sectional views of Eudimorphodon ranzii. Note the overlap of the posterior ribs over the hind limbs and the very wide torso. The cross section shows the 2nd dorsal ribs and the 23rd. Note the small ischium which could only produce small eggs. A little taller and wider than we thought before. The forelimbs are pretty short relative to the torso.

Figure 4. Lateral, dorsal and cross-sectional views of Eudimorphodon ranzii. Note the overlap of the posterior ribs over the hind limbs and the very wide torso. The cross section shows the 2nd dorsal ribs and the 23rd. Note the small ischium which could only produce small eggs. A little taller and wider than we thought before. The forelimbs are pretty short relative to the torso.

Step four: Tracing the colorized bones for the final reconstruction. (Fig. 4)
If I just attempted a lateral view I would have missed out on the very broad posterior torso based on the length of the posterior ribs. So I create both a dorsal view and a cross section view. Note that the sternal ribs, rarely found on most pterosaurs, extend laterally to meet the dorsal rib tips in Eudimorphodon. This give it a slightly wider body anteriorly, increasingly wider posteriorly. This is an odd autapomorphy, but it is based on many ribs, so it can’t be ignored. As you can see from the in situ image (Fig. 1) those long posterior ribs were there all the time. They were simply ignored by myself and others.

Eudimorphodon: a little odder than we thought
That torso is odd. Rather than tapering toward the pelvis, as in many other pterosaurs and tetrapods in general, the posterior torso is flat and wide, roofing the femora. My guess it provides a greater volume for eggs or respiration. With such small eggs, more eggs could have been carried by the mother. Note that the predecessor of E. ranzii, MPUM 6009, has a much deeper pelvic opening, likely to produce one large egg at a time. Note the reduction of the pelvis is also reflected in the reduction of the number of sacrals to four or five depending on the connection to the posterior pelvis.

If there is anything wrong with the results here, please let me know. If not feel free to use the technique yourself. I think it works pretty well.

I also don’t make these identifications without entering the taxa into a phylogenetic analysis that typically finds the same traits in sister taxa. Unfortunately posterior ribs are virtually unknown among Triassic and Early Jurassic sisters.

Pterosaur workers haven’t produced too many Eudimorphodon reconstructions, and certainly none that have recovered the oddly long posterior ribs. My earlier reconstructions were given generic ribs. So I did a bad thing. I went along with the paradigm of a tubular pterosaur body without testing that paradigm. While it takes a lot of work for small discoveries such as this, and the results are minor changes, well, I had nothing better to do on a quiet Sunday.

Wild R 1978. Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen Trias von Cene bei Bergamo, Italien. Bolletino della Societa Paleontologica Italiana 17(2): 176–256.
Zambelli R 1973. Eudimorphodon ranzii gen.nov., sp.nov. Uno Pterosauro Triassico. Rendiconti Instituto Lombardo Accademia, (rend. sc.) 107: 27-32.

Breathing in a box – Respiration in pterosaurs (Geist et al. 2013)

A new paper on pterosaur breathing has arrived.
Unfortunately, Geist et al. (2013) follow Claessens et al. (2009) in hoping that the prepubis was mobile in order to drive a thoracic lung pump. It’s not. (BTW, we looked at Claessens et al earlier here).

Geist et al. report, “we note that many of the large pterodactyloid taxa had relatively rigid trunks due to a high degree of fusion of the thoracic skeleton, a condition we describe as a synthorax… but also appear to have severely constrained ribcage movement for respiration.”

How is this shown?
Geist et al. report, “Although their morphology is variable, they [pterosaurs] always have a narrow, rod-like caudoproximal peduncle, but widen and flatten cranially. Their general appearance and orientation is often strikingly similar to that of the pubic bones of extant crocodilians.”

This is an error.
Prepubes are oriented ventrally with a non-moving butt joint, and only sometime slightly cranially. Thus they are not similar to extant crocodilians, which rotate their pubes uniquely.

Campylognathoides (CM 11424), the earliest pterosaur with a reduced pedal digit 5.

Figure 1. Campylognathoides (CM 11424). Note the prepubis, perpendicular to the spinal column and butt-joined to the pubis is immobile. Think of it like the booted pubis in T-rex, which was not used in reparation. Note the way Geist et al orient the prepubis is figure 2.

After just reporting that some pterosaurs fuse their prepubes medially, Geist et al. report, “The hinge-like [medial] articulation with the pubic bones indicates that the prepubic bones were highly mobile in the vertical plane, but were restricted in transverse movements.”

This is also bad reporting, based on bad drawings in the literature, (like Claessen 2009), not based on manipulating 3D prepubes on pubes, as I have done over several dozen taxa.

The prepubes act as immobile pubic extensions. And in that capacity they anchor adduction muscles to the femur, like the long booted pubis of T-rex. They have nothing to do with respiration. Nor were they co-oped for respiration from their original function.

Sure prepubes had large muscle scars. They were attached to large leg muscles!

The most mobile parts of the pterosaur thorax were the posterior dorsal ribs. They were slender and single-headed to retain their mobility. Lizards breathe with their ribs. Here’s a short video and another video for anyone who can’t picture when a lizard holds still, their rib cage keeps pumping. Holding still, btw, is the same as having a fused backbone, in terms of methodology.

Geist et al report, “Claessens et al. (2009) presented a detailed model for an avian-like mechanism of sternal excursion as the primary lung ventilation mechanism for both small and large pterosaurs—the “skeletal breathing pump” model.” In essence the sternal complex rotated on the coracoid joint with some movement from the scapulocoracoid. This pulled the ribs  and gastralia forward increasing thorax volume.”

Geist et al were critical of Claessens noting, “In pterosaurs the orientation of the rib articulations on the thoracic vertebrae are very unlike those of birds (Fig. 3), and likely would not have permitted the degree of fore-aft movement of the caudal ribs as proposed by Claessens et al. (2009).”

Geist et al continue, “Three dimensionally preserved specimens indicate that pterosaur ribs project more or less perpendicular to the long axis of the body (e.g., see Fig. 4A–E). A caudal inclination of these ribs similar to the orientation of those in extant birds cannot be confirmed here.”

That would be odd. Pterosaurs are now flying pancakes? No. Perhaps that’s just an odd choice of words because their illustrations don’t follow that morphology. Sharovipteryx was the flying pancake!

Geist et al. report, “the morphology of the synthorax of large pterosaurs is permissive of extracostal mechanisms that resembled the visceral pumps [diaphragms] found in mammals and crocodilians.” After considering turtle, mammal, bird and croc respiration, they never once mentioned lizards. Unfortunate.

Figure 1. Click to animate. Pterosaur breathing using a liver pump as envisioned by Geist et al. 2013.

Figure 2. Click to animate. Pterosaur breathing using a liver pump as envisioned by Geist et al. 2013. Unfortunately, the prepubis was immobilized due to a butt joint at the pubis, so this isn’t accurate.

Ultimately, Geist et al. report, “Accordingly, we propose a model for large pterosaurs in which a more or less crocodilian-like visceral displacement pump drove the inhalation phase of the respiratory cycle, and contraction of flank muscles acting on the gastralia and prepubic bones restored the viscera to their initial positions to drive exhalation (Fig. 9). In our model, sheets of diaphragmaticus-like skeletal muscle originated on the pubis, prepubic bones, and/or caudal-most gastralia and inserted on a transverse septum or the fascia of the liver.”

The Triebold Pteranodon, one of the most complete ever found. The metacarpals are quite a bit longer here. So is the beak.

Figure 3. The Triebold Pteranodon. Note the orientation of the prepubes, ventrally, in line with the standing femora. The Geist orientation is based on Claessens et al. (2009) which was based on a mistake.

Of course, as they admit, there is no evidence for this. And I’ll add some serious evidence against it.

Some pterosaur pubes are much shorter than their ischia. What would this mean for a cranially directed prepubis? Take a look at Figure 1 and put it together for yourself. Unfortunately, Geist et al. looked at only those pelves that had a long pubis.

If pterosaurs had a avian-like or monitor-like air-sac system associated with their lungs, then the posterior ribs could have pumped air in and out of those sacs, driving air through the lungs. Very typical of lizards, not crocs or birds.

To their credit, Geist et al. touch on the fact that sternal ribs are rarely ossified, noting they were likely often cartilage-based.

Claessens LPAM, O’Connor PM, Unwin DM 2009. Respiratory Evolution Facilitated the Origin of Pterosaur Flight and Aerial Gigantism. PLoS ONE 4(2): e4497. doi:10.1371/journal.pone.000449
Geist NR, Hillenius WJ, Frey E, Jones TD and Elgin RA 2013. Breathing in a box: Constraints on lung ventilation in giant pterosaurs. The Anatomical Record. 013 Dec 10.