Another long-necked embryo tritosaur: Li et al. in press

This appears to be
yet another Tanystropheus-like and Dinocephalosaurus-like taxon, yet not closely related to either. Earlier we looked at another similar embryo, still within its mother.

Li, Rieppel and Fraser in press (2017)
bring us a new curled up (as if in an egg, but without a shell) embryo from the Guanling Formation (Anisian), Yunnan province, China (Figs. 1, 2). The specimen is unnamed and not numbered. It appears to combine the large head and eyes of langobardisaurs with the short limbs and many cervical vertebrae of Dinocephalosaurus. Please remember, in this clade, juveniles do not have a short rostrum and large eyes unless their parents also had these traits.

Figure 1. The unnamed and not numbered Triassic embryo Li et al. assign to a new species close to Dinocephalosaurus.

Figure 1. The unnamed and not numbered Triassic embryo Li et al. assign to a new species close to Dinocephalosaurus. At 72 dpi monitor resolution, this image is 2.5x life size. Here bones are colorized, something Li et al. could have done, but avoided. I’m happy to report that the line drawing was traced by Li et al. on their own photo. The two are a perfect match.

Unfortunately
Li et al. have no idea what they’re dealing with phylogenetically. They relied on old invalidated hypotheses of relationships. They report the specimen:

  1.  is a marine protorosaur and an archosauromorph – actually it is a marine tritosaur lepidosaur. Taxon exclusion and traditional bias hampered the opinion of Li et al. They did not perform a phylogenetic analysis.
  2. is closely related to Dinocephalosaurus – actually it is more closely related to the much smaller, but longer-legged Pectodens (Figs. 4, 5). In the large reptile tree (LRT, 1036 taxa) 8 steps are added when the embryo is force-nested with Dinocephalosaurus. The embryo is distinct enough that the new specimen deserves a new genus.
  3. confirms viviparity – probably not (but see below). The specimen is confined within an elliptical shape (Fig. 1), as if bound by an eggshell or membrane, which was not preserved. Perhaps, as in pterosaurs and many other lepidosaurs, the embryo was held within the mother’s body until just before hatching, within the thinnest of egg shells and/or membranes.
  4. is too immature to describe it as a new taxon – not so. Tritosaur lepidosaurs (from Huehuecuetzpalli to Pterodaustro) develop isometrically. Thus, full-term embryos and hatchlings have adult proportions.
Figure 2. The specimen from figure 1 unrolled for clarity. This specimen most closely resembles the basal langobardisaur, Pectodens, not Dinocephalosaurus. Remember, tritosaurs develop isometrically. Embryos closely resemble adults. That's why three scale bars are included.

Figure 2. The specimen from figure 1 unrolled for clarity. This specimen most closely resembles the basal langobardisaur, Pectodens, not Dinocephalosaurus. Remember, tritosaurs develop isometrically. Embryos closely resemble adults. That’s why three scale bars are included. This specimen has feeble limbs but a strong swimming tail, distinct from that of Dinocephalosaurus.

Li et al. report
“In the fossil record only oviparity and viviparity can be distinguished, Ovoviviparity of different intermediate stages, which is often observed in modern squamates would then be referred to the category of viviparity, whatever the stages of maturity and nutritional patterns are.” Yes, they correctly report ovoviviparity in squamates, which are the closet living relatives of tritosaur lepidosaurs. That’s exactly what we have here.

Figure 1. The new Dinocephalosaurus has traits the holotype does not, like a longer neck with more vertebrae, a robust tail with deep chevrons and a distinct foot morphology with an elongate pedal digit 4.

Figure 3. The new Dinocephalosaurus has traits the holotype does not, like a longer neck with more vertebrae, a robust tail with deep chevrons and a distinct foot morphology with an elongate pedal digit 4.

Li et al. report,
“[The] skeleton is preserved tightly curled so as to produce an almost perfect circular outline, which is strongly indicative of an embryonic position constrained by an uncalcified egg membrane.”

Figure 2. Pectodens skull traced using DGS techniques and reassembled below.

Figure 4. Pectodens skull traced using DGS techniques and reassembled below. No sclerotic ring here. 

Distinct from Pectodens the new genus embryo has:

  1. 24 cervicals
  2. 29 dorsals
  3. 2 sacrals
  4. and about 64 caudals
Figure 1. Pectodens reconstructed using the original tracings of the in situ fossil in Li et al. 2017.

Figure 5. Pectodens reconstructed using the original tracings of the in situ fossil in Li et al. 2017. The skull shown here is the original reconstruction. Compare it to figure 4.

Li et al overlooked:

  1. strap-like coracoids, strip-like clavicle and T-shaped interclavicle
  2. scattered manual elements
  3. pelvic girdle
  4. ectopterygoid, jugal, articular, angular, surangular

Li et al. report:
“The fewer cervical vertebrae (24 as opposed to 33 (based on an undescribed specimen kept in the IVPP)), and the presence of sclerotic plates are features inconsistent with Dinocephalosaurus.This embryo therefore documents the presence of at least one additional dinocephalosaur-like species swimming in the Middle Triassic of the Eastern Tethys Sea.

“Scleral ossicles have previously not been described in any protorosaur.”
– but they are common in tritosaur lepidosaurs, like pterosaurs.

Figure 6. Pectodens adult compared to today's embryo and its 8x larger adult counterpart after isometric scaling.

Figure 6. Pectodens adult compared to today’s embryo and its 8x larger adult counterpart after isometric scaling. Looks more like Pectodens than Dinocephalosaurus, doesn’t it? See taxon inclusion WORKS! Sclerotic rings were omitted here to show skull bones. The ring would have had a smaller diameter if if were surrounding a sphere, rather than crushed flat. 

A word to traditional paleontologists:
Don’t keep digging yourself deeper into invalidated hypotheses and paradigms. Use the LRT, at least for options.

Don’t give up on naming embryos
and adding them to phylogenetic analysis, especially if they are tritosaur lepidosaurs. Hatchlings nest with adults so you can used hatchlings in analysis.

Don’t avoid creating reconstructions.
That’s a great way to discover little splinters of bone, like clavicles and coracoids, that would have been otherwise overlooked.

The LRT is here for you.
BETTER to check this catalog prior to submission rather than have your work criticized for being unaware of the latest discoveries or overlooking pertinent taxa AFTER publication.

References
Li C, Rieppel O, Fraser N C, in press. Viviparity in a Triassic marine archosauromorph reptile. Vertebrata PalAsiatica, online here.

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Pectodens: basal to tanystropheids and pterosaurs

It’s always good
to see another tritosaur. That’s the lineage that gave rise to a menagerie of taxa, including pterosaurs. That’s a heretical hypothesis of relationships recovered by the large reptile tree (LRT, 997 taxa).

Figure 1. Pectodens reconstructed using the original tracings of the in situ fossil in Li et al. 2017.

Figure 1. Pectodens reconstructed using the original tracings of the in situ fossil in Li et al. 2017.

Li et al. 2017 conclude:
“A new, small terrestrial tetrapod is described from the Middle Triassic of Yunnan, China. Pectodens zhenyuensis n. gen. n. sp. bears very characteristic elongate teeth forming a comb-like marginal dentition. The elongate cervicals of Pectodens zhenyuensis n. gen. n. sp. with low neural spines together with the morphology of the cervical ribs are features consistent with protorosaurs, such as Macrocnemus. However, the imperforate puboischiadic plate, simple rounded proximal tarsals, and a straight 5th metatarsal are primitive characteristics. Unlike tanystropheids, but in common with Protorosaurus (personal observation, N.C. Fraser, 2013), both lack a thyroid fenestra in the pelvis.”

Figure 2. Pectodens skull traced using DGS techniques and reassembled below.

Figure 2. Pectodens skull traced using DGS techniques and reassembled below. Here a quadratojugal process of the jugal is identified and other parts are assembled with greater accuracy than a freehand sketch (Fig. 1).

Pectodens zhenyuensis (Li et al. 2017; IVPP V18578; Anisian, Middle Triassic; 38cm in length) was originally considered a diapsid and a possible protorosaur. Here Pectodens nests between Macrocnemus and Langobardisaurus (Fig. 3). Originally the interclavicle, sternum and quadratojugal were overlooked.

Note the large orbit, the long metarsal 5 and the perforated pubis. The elongate caudal transverse processes anchor powerful leg muscles.

Occasionally within the Tritosauria
metatarsal 5 is not short, but elongate. It is always axially twisted. The pubis and ischium typically angle away from one another, but sometimes produce a thyroid fenestra. Tritosaurs have a sternum, like many other lepidosaurs do. Protorosaurs do not have a sternum.

Li et al. did not attempt a phylogenetic analysis.
Instead they made educated guesses as to the affinities of Pectodens, overlooking the variation present in related taxa revealed in a cladogram. Pulling a Larry Martin (highlighting or letting yourself get confused by one or two traits) is never a good idea. Better to let hundreds of traits determine the exact nesting of a taxon without bias. Let the taxa nest themselves. Let the convergent traits simply be convergent traits.

Earlier we looked at the pectoral girdle and sternum of Langobardisaurus, Huehuecuetzpalli and other tritosaurs. Pectodens fits right in.

The posterior maxillary teeth in Pectodens
are wider at their base presaging the grinding teeth found in Cosesaurus, basal pterosaurs and Langobardisaurus.

Note the way the fingers and toes
bend anteriorly during use. That’s a lepidosaur trait. Pectodens would have had sprawingling hind limbs given its simple femoral head. Tracks matching such curved toes are known from the Middle Triassic.

Li et al. considered Pectodens to be the first terrestrial taxon
from the its locality. And that’s definitely a probability. However, given that Tanystropheus and others may have been underwater bipedal predators (squid parts were found in their torso), let’s leave open the possibility that Pectodens was maybe dipping its toe in the water.

Figure 1. Subset of the LRT focusing on Tritosauria. Pectodens nests here basal to the Characiopoda (Tanystropheids + Fenestrasauria including pterosaurs).

Figure 1. Subset of the LRT focusing on Tritosauria. Pectodens nests here basal to the Characiopoda (Tanystropheids + Fenestrasauria including pterosaurs).

Let’s not continue to nest tanystropheids
with protorosaurs. Sure they share several traits by convergence, but they are not related to one another as determined by a large gamut analysis, the LRT.

References
Li C, Fraser NC, Rieppel O, Zhao L-J and Wang L-T 2017. A new diapsid from the Middle Triassic of southern China. Journal of Paleontology.7 pp. doi: 10.1017/jpa.2017.12

 

Live birth in ‘Dinocephalosaurus’? Maybe. Maybe not.

Yesterday Liu et al. 2017 reported on
a pregnant Dinocephalosaurus (Figs. 1-5). This is wonderful and exciting news. However, the embryo is NOT in the process of passing through the cloaca, as we’ve seen in ichthyosaurs. The embryo is much higher in the abdomen, still in the uterus. So the headline “Live birth in an archosauromorph reptile” is… at best… premature. Live birth is still a possibility. A critical examination of the data reveals a few more major and minor problems.

Dinocephalosaurus in resting, feeding and breathing modes.

Figure 1. The holotype (not the new specimen) of Dinocephalosaurus in resting, feeding and breathing modes. In breathing mode the throat sac would capture air that would not be inhaled until the neck was horizontal at the bottom of the shallow sea. Orbits on top of the skull support this hypothesis. Image from Peters 2005. The new specimen has a longer neck, a more robust tail, and a different pedal morphology.

Unfortunately
the authors nested Dinocephalosaurus within the Archosauromorpha (Fig. 2). That is incorrect. Dinocephalosaurus nests within the new Lepidosauromorpha in the large reptile tree (LRT, 952 taxa), which minimizes the taxon exclusion problem suffered by the much smaller taxon list in the Liu et al. 2017 tree.

Figure 6. Cladogram from Liu et al. 2017 with colors added based on results from the LRT. Taxon exclusion is a major problem here.

Figure 2. Cladogram from Liu et al. 2017 with colors added based on results from the LRT. Taxon exclusion is a major problem here. Note in the Liu et al. cladogram members of the Protorosauria are divided into three clades. In sympathy, members of the Tritosauria and Protorosauria do indeed converge with one another. More taxa clears up the problem shown here of cherry-picking taxa.

Dinocephalosaurus actually nests
within the lepidosaur clade, Tritosauria, a clade that also includes Tanystropheus, pterosaurs and several other taxa (Fig. 7) that had been mistaken for protorosaur relatives in the Liu et al and other prior studies.

As a lepidosaur, 
Dinocephalosaurus would have been able to retain embryos within the mother far longer that in extant archosauromorphs. And based on the extreme thinness of pterosaur eggshells (closest known relatives with embryos, Fig. 7), those leathery eggshells only develop just prior to egg laying. So live birth is only one of a spectrum of options for Dinocephalosaurus. As in pterosaurs, the eggs could have hatched shortly after the female laid them on the shoreline.

Dinocephalosaurus. Note the very narrow cranial portion of the skull and the very wide cheeks. That, by it self, opens the orbits dorsally. Sure there's some lateral exposure, but those eyes are looking up!

Figure 3. The holotyype of Dinocephalosaurus. Although extremely similar, the new specimen is different in several ways. See below.

Liu et al. report that live birth is unknown in the Archosauromorpha.
However, in the LRT mammals and enaliosaurs (sauropterygians + ichthyosaurs) are both archosauromorphs that experience live birth. Hyphalosaurus, a choristodere archosauromorph, had extremely thin eggshells and retained developing embryos inside the mother until laying those eggs.

Figure 5. Hypothetical Tanystropheus embryo compared to Dinocephalosaurus embryo.

Figure 4. Hypothetical Tanystropheus embryo compared to part of an embryo of the new specimen attributed to Dinocephalosaurus.

More about that embryo
What little is preserved of the Dinocephalosaurus embryo (Fig. 4) is curled up in its amniotic sac, as one would expect for any reptile embryo still in utero. For comparison, note the hypothetical Tanystropheus embryo alongside it. That long neck has to go somewhere and Dinocephalosaurus provides further evidence that juvenile tritosaurs were isometric duplicates of their adult parents. That long neck did not develop with maturity. Among other tritosaurs we see juveniles similar in proportion to adults in the basal form, Huehuecuetzpalliand all pterosaur embryos.

Liu et al. further report. “Despite the complexity of this transition, viviparity has evolved at least 115 times in extant squamates (lizards and snakes), in addition to a single time in the common ancestor of therian mammals. Moreover, viviparity is a common reproductive mode in extinct aquatic reptiles including eosauropterygians, ichthyosaurs, mosasauroids, some choristoderans and likely mesosaurs.” Since mosasauroids are extinct squamates that makes at least 116 times for lepidosaurs.    Some living squamates produce eggs that hatch shortly after they are expelled, a sort of transition from oviparity to viviparity. That’s where pterosaurs fall and perhaps Dinocephalosaurus.

More cladogram issues
The Liu et al. figure 1 cladogram shows a polytomy of most reptilian clades arising during the Permian. No such polytomy appears in the LRT in which Archosauromorpha diverged from the Lepidosauromorpha tens of millions of years earlier in the Viséan (Lower Carboniferous). Liu et al. mistakenly report that trilophosaurs, rhynchosaurs and pterosaurs are archosauromorph reptiles. They are lepidosauromorph reptiles in the LRT.

Figure 1. The new Dinocephalosaurus has traits the holotype does not, like a longer neck with more vertebrae, a robust tail with deep chevrons and a distinct foot morphology with an elongate pedal digit 4.

Figure 5. The new Dinocephalosaurus has traits the holotype does not have, like a longer neck with more vertebrae, a robust tail with deep chevrons and a distinct foot morphology with a more elongate pedal digit 4. The partial embryo is in magenta at left.

The new specimen looks like a Dinocephalosaurus, but is it one?
Distinct from the holotype, the new specimen has a deep robust tail with deep chevrons (Fig. 5) as in Litorosuchus (Fig. 6), also from China, but from higher strata. They all share a common ancestor in one of the highly variable Macrocnemus specimens (Fig. 7). The toes of the new specimen are more asymmetric. The neck probably has more vertebrae (several are lost, but note the longest ones are NOT at the base of the neck in the holotype). Unfortunately little more can be said with so much of the mother lacking at present. We’ve already seen a Chinese Tanystropheus similar to, but not identical to the European Tanystropheus. We can imagine even greater variation within the available gamut of the present sparse fossil evidence.

Figure 3. Litorosuchus compared to Macrocnemus and kin at two scales.

Figure 6. Litorosuchus compared to Macrocnemus and kin at two scales. Litorosuchus has deep chevrons and a robust tail, as in the new specimen, lacking in Dinocephalosaurus.

Figure 2. Given the scrappy, skull-less data, the putative Dinocephalosaurus either nests with its namesake or by convergence with Litorosuchus which shares deep chevrons and a robust tail, among other traits.

Figure 7. Given the scrappy, skull-less data, the putative Dinocephalosaurus either nests with its namesake or by convergence with Litorosuchus which shares deep chevrons and a robust tail, among other traits.

It really is too much
to expect identical specimens to come from distinct fossil bearing strata. So variation within Dinocephalosaurus is a possibility. But so is convergence from the lineage of Litorosuchus.

Next steps
The paleo-community needs to include more specimen-based taxa in their cladograms or the Liu et al. problem (not restricted to them!) is going to continue ad infinitum. I know that’s a lot of work. But it can be done (I’ve done it!) and it needs to be done. Just start with a large gamut analysis and keep adding taxa to it. That will make the current phylogenetic problems go away.

Final note
Images of tanystropheids and dinocephalosaurs swimming horizontally through open waters (Liu et al. 2017 their figure 3) may not be an accurate portrayals of their daily lives. Other options have been published (Fig. 1) or appear online (Fig. 8). Odd-looking tetrapods often have uncommon niches and atypical behaviors.

Tanystropheus underwater among tall crinoids and small squids.

Figure 8. Tanystropheus in a vertical strike elevating the neck and raising its blood pressure in order to keep circulation around its brain and another system to keep blood from pooling in its hind limb and tail.

References
Li C, Rieppel O and LaBarbera MC 2004. A Triassic aquatic protorosaur with an extremely long neck. Science 305:1931.
Liu, J. et al. 2017. Live birth in an archosauromorph reptile. Nature Communications 8, 14445 doi: 10.1038/ncomms14445
Peters D, Demes B and Krause DW 2005. Suction feeding in Triassic Protorosaur? Science, 308: 1112-1113.

 

What is Triopticus? (It’s not what they think it is…)

Updated July 13, 2017 with new bone identifications for Triopticus. These further cement the sisterhood to Tanytrachelos. 

It’s been a long time
since an interesting ‘reptile’ showed up in the literature. Especially an ‘enigma’ like this one.

A recent paper by Stocker et al. 2016
reports on a domed and expanded Late Triassic cranium that they identify as an archosaur, but it’s unlike that of any other archosaur. Triopticus primus was named for its three eyes, with a big one on top (Fig. 1). The authors compared the domed appearance of the cranium in Triopticus with a Cretaceous dome-headed ornithischian dinosaur, Stegoceras.  They also discussed convergence in general and provided a CT scan brain endocast of Triopticus.

Unfortunately 
the authors employed a prior cladogram (Fig. 2) by Nesbit et al. 2015, expanded from Pritchard et al. 2015. that was shown to not recover sister taxa that looked alike and did not provide a gradual accumulation of derived traits at several nodes. In their cladogram Triopticus nested without resolution among basal archosauriforms like Proterosuchus, which looks nothing like it. By contrast, the LRT was able nest and fully resolve Triopticus elsewhere.

Figure 1. In round 1 I added characters shown here to the LRT in two passes. One recovered a sisterhood with mesosaurs. The other nested with Tanytrachelos, among the tanystropheid tritosaur lepidosaurs. Both shown here for comparison.

Figure 1. In round 1 I added characters shown here to the LRT in two passes. One recovered a sisterhood with mesosaurs. The other nested with Tanytrachelos, among the tanystropheid tritosaur lepidosaurs. Both shown here for comparison. Triopticus would be 2x the size of the giant Tanyrachelos from New Mexico.

From the Stocker et al. abstract:  “Exemplifying this extreme morphological convergence, we present here a new dome-headed taxon from the assemblage, which further illustrates the extraordinary range of morphological disparity present early in the Late Triassic.” That ‘extraordinary range’ should be — and will be — chopped down substantially with the right sister taxa.

A few problems with the archosauriform hypothesis include:

  1. No other archosauriforms, until you get to pachycelphalosaurs in the Cretaceous, expand the cranium deleting the upper temporal fenestra.
  2. The entire rostrum and mandible is absent, so no naris, antorbital fenestra or teeth are known, even in part.
  3. They dubiously identified an antorbital fenestra and fossa at the edge of the fossil.
  4. …and they were not aware that Tanytrachelos and kin, including pterosaurs within the – Lepidosauria -, also have an antorbital fenestra, but without a fossa.
  5. A large pineal opening is present, but never present at such a size in archosauriforms.
  6. The extreme angle of the rostrum coupled with the large orbit are traits not found in basal archosauriforms that typically have a long boxy rostrum.
Figure 2. Stocker et al. 2106 cladogram nesting Triopticus uncertainly within a set of unresolved basal archosauriforms. The LRT completely resolves that node.

Figure 2. Stocker et al. 2106 cladogram nesting Triopticus uncertainly within a set of unresolved basal archosauriforms and far from the Tanystropheidae. The LRT completely resolves all nodes. Note how this cladogram mixes Lepidosauromorpha with Archosauromorpha and separates the protorosaurus, Protorosaurus and Prolacerta.

This is a perfect problem
for the large reptile tree (LRT) which now provides then 820, now 1036 opportunities for Triopticus to nest in. With that large number of taxa, unfortunately I had to split the matrix in two, even for a simple Heuristic Search. By contrast, the Stocker et al. matrix included 30 taxa and 247 characters.

Stocker et al. report,
“We chose this dataset because the following combination of character states in Triopticus are also present in some archosauromorph taxa:

  1. presence of a single occipital condyle;
  2. ossified laterosphenoid;
  3. presence of a metotic strut of the otoccipital;
  4. presence of upper and lower temporal fenestrae;
  5. presence of an antorbital fenestra and fossa formed by the lacrimal.”

They provided no reconstructions of included taxa.

First,
I tested Triopticus against basal tetrapods and the new Lepidosaurmorpha and found that Triopticus nested with the aquatic, long-necked tritosaur Tanytrachelos (Fig. 1), large specimens of which were recently found in New Mexico (Fig. 3). Like Triopticus the rostrum descends at a high angle from a tall cranium in Tanytrachelos, which also shares a large orbit and a large pineal foramen (at present known only from sister taxa). Like related fenestrasaurs and langobardisaurs, Tanytrachelos also had a small antorbital fenestra without a fossa, but that would have been beyond the rim of the broken skull in Triopticus (Fig. 5).

Figure 2. A large incomplete Tanytrachelos from New Mexico compared to the smaller more complete East Coast specimen. Triopticus would be twice as large as the New Mexico specimen.

Figure 3. A large incomplete Tanytrachelos from New Mexico compared to the smaller more complete East Coast specimen. Triopticus would be twice as large as the New Mexico specimen.

Second,
I tested Triopticus with the rest of the matrix, the new Archosauromorpha, and found that Triopticus nested with the mesosaurs (Fig. 4), an aquatic enaliosaur clade close to thalattosaurs and ichthyosaurs, all derived from basal pachypleurosaurs. It did not nest with archosauriforms. While basal mesosaurs have typical diapsid temporal regions, Mesosaurus, like Triopticus, closes up the upper temporal fenestra, then the lateral temporal fenestra with bone expansion.  Mesosaurs also retain a relatively large pineal foramen and have large eyes, but they don’t have a sharply descending preorbital region.

Mesosaurus

Figure 4. Mesosaurus, like Triopticus, has a large pineal foramen and expands the skull bones to obliterate former temporal fenestrae.

Digital Graphic Segregation
was applied to the cranial lump that is Triopticus (Fig. 5) and the skull suture patterns, perhaps overlooked by those with firsthand access due to the expansion of the cranial bones, revealed a Tanytrachelos-like morphology (Fig. 2). I illustrate this interpretation here with the hope that this hypothesis can be either confirmed or falsified. This is a tough assignment.

Figure 5. Triopticus reconstructed along the bauplan of Tanytrachelos.  The upper temporal fenestra is the top half of a divided lateral temporal fenestra. At 72 dpi this is 90 percent of actual size.

Figure 5. Triopticus reconstructed along the bauplan of Tanytrachelos. The upper temporal fenestra is the top half of a divided lateral temporal fenestra. At 72 dpi this is 90 percent of actual size.

Tanystropheids
have been reported from the Hayden Quarry of northern New Mexico (Chinle Formation) far from the west central Texas location of Otis Chalk. Stocker et al. included Tanytrachelos in their study, even though they have not provided a reconstruction of it, so it is difficult to imagine how they interpreted it. Tanystropheids, in general, have widely varying skull shapes. Triopticus appears to have expanded the morphospace just a little, not a lot.

The loss of ventral material in the Triopticus fossil
appears to have occurred at the roof the narial/oral opening.

So what other long-necked animal
expands the cranium like Triopticus? Giraffa, the giraffe. Maybe it will turn out to be a better analogy than short-necked Stegoceras?

References
Nesbitt SJ, Flynn JJ, Pritchard AC, Parrish JM, Ranivoharimanana L and Wyss AR 2015. Postcranial osteology of Azendohsaurus madagaskarensis (?Middle to Upper Triassic, Isalo Group, Madagascar) and its systematic position among stem archosaur reptiles. Bulletin of the American Museum of Natural History 899, 1-125.
Pritchard AC, Turner AH, Nesbitt SJ, Irmis RB and Smith ND 2015. Late Triassic tanystropheid (Reptilia, Archosauromorpha) remains from northern New Mexico (Petrified Forest Member, Chinle Formation): insights into distribution, morphology, and paleoecology of Tanystropheidae. Journal of Vertebrate Paleontology, 10.1080/02724634.02722014.02911186.
Stocker MR, NesbittSJ, Criswell KE, Parker WG, Witmer LM, Rowe TB, Ridgely R  Brown MA 2016. A Dome-Headed Stem Archosaur Exemplifies Convergence among Dinosaurs and Their Distant Relatives. Current Biology (advance online publication)DOI: http://dx.doi.org/10.1016/j.cub.2016.07.066   pdf

Triassic gastric pellet semi-reconstructed, better this time…

A while back
Dalla Vecchia, Wild and Muscio (1989) described a small pellet (MFSN 1891, Fig. 1) of Late Triassic bones from the Dolomia di Forni Formation of Firuli (NE Italy) as a small jumble of pterosaur bones. They tentatively referred it to Preondactylus, the only pterosaur known at the time from that formation. This was an early work for all three paleontologists.

Following the original paper
Earlier I attempted a reconstruction of the elements based on the pterosaur model. I recognized then that it didn’t turn out too well. I was working from the original drawings. Now new data has been published and a new hypothesis has been put forth that makes much more sense.

Figure 1. Several views of the Triassic gastric pellet formerly considered pterosaurian, but now considered langobaridsaurian. Elements from a surface photo, a microCT scan of the opposite side still buried in matrix, and DGS colors. Not all bones have been colored here.

Figure 1. Several views of the Triassic gastric pellet formerly considered pterosaurian, but now considered langobaridsaurian. Elements from a surface photo, a microCT scan of the opposite side still buried in matrix, and DGS colors. Not all bones have been colored here, but employed colors are assembled in figure 2. The long cervical at upper left is 1 cm long. So is the scale bar. The pellet is about 5 cm wide.

Recently 
Holdago et al. (2015) redescribed the pellet in much greater detail using microCT acquisition. They concluded “The best candidate for the pellet is not a pterosaur, but a protorosaurian like Langobardisaurus.  Therefore, the skeletal remains could belong to a still unknown small reptile with procoelous dorsal vertebrae, rather elongate and probably procoelous cervical vertebrae with low neural arch and spine, filiform cervical ribs, at least some dicephalous dorsal ribs, elongated and hollow limb bones, and no osteoderms.”

They did not attempt a reconstruction,
so I do so here (Fig. 2) following the hypothesis that the elements belong to a langobardisaur (contra Holdgago, et al., not a protorosaur but a tritosaur lepidosaur).

Figure 2. The elements of MFSN 1891 assembled to form a langobardisaur in a bipedal pose.

Figure 2. The elements of MFSN 1891 assembled to form a langobardisaur in a bipedal pose. Some langobardisaurs have a very long neck, slender limbs and a short tail. Lots of guesswork here.

Lots of guesswork here. 
Everything is tentative. The toes could be ribs. Lots of slivers and scraps left over. More complete langobardisaurs (Fig. 3) have 8 cervicals, but they are related to tanystropheids, with 13 cervicals. Renesto et al. (2002) considered langobardisaurs as likely facultative bipeds in the manner of the many extant facultative bipedal lizards, all with sprawling hind limbs.

Langobardisaurus tonneloi reconstructed. Note the cosesaur-like pectoral girdle.

Figure 3. Langobardisaurus tonneloi reconstructed. Note the cosesaur-like pectoral girdle.

MFSN 1891 needs to be dissembled
in high resolution, then reassembled like a puzzle. I’d like to help if possible. Here (Fig. 2) is a first draft lo rez example leading to others of greater detail in the future. Worthwhile taking another look at the pes (Fig. 3) which greatly resembles a basal pterosaur pes with that elongate p5.1. It resembles a pterosaur pes because these two taxa are related (Peters 2000).

Figure 6. Click to view full scale on a 72 dpi screen. Tanystrachelos compared to the gastric pellet lepidosaur.

Figure 4. Click to view full scale on a 72 dpi screen. Tanystrachelos compared to the gastric pellet lepidosaur. The large hemal arches on the gastric pellet are the genesis of the paddle-like hemal arches on Tanytrachelos and Tanystropheus.

Compared to the tritosaur Tanytrachelos (Fig. 4)
the gastric pellet reptile has a similar number of cervicals, but longer limbs and longer cervicals. Are we seeing the origin of Tanystropheus (Fig. 5) here? Or a hatchling? The large hemal arches appear to have homologs in Tanytrachelos and Tanystropheus.

Tanystropheus and kin going back to Huehuecuetzpalli.

Figure 5. Tanystropheus and kin going back to Huehuecuetzpalli. Two scales here, one yellow, one white.

Then we have Fuyuanssaurus, 
a tiny tanystropheid (Fig. 6) about twice the size of the gastric pellet reptile. Unfortunately we don’t know if it was long-legged or not. Notably the skull elements of Fuyuansaurus, which we looked at earlier here were all quite slender. This is the model we should use for the gastric pellet lizard until data suggests another model.

Figure 2. Click to enlarge. Reconstruction of Fuyuanasaurus. Fraser et al. identified a strange circular object as the pubis, but no sister taxa have a circular pubis. Here it is tentatively ID'd as an egg because a standard pubis is found nearby.

Figure 6. Click to enlarge. Reconstruction of Fuyuanasaurus. Fraser et al. identified a strange circular object as the pubis, but no sister taxa have a circular pubis. Here it is tentatively ID’d as an egg because a standard pubis is found nearby.

References
Dalla Vecchia FM, Wild R and Muscio G 1989. Pterosaur remains in a gastric pellet from Upper Triassic (Norian) of Rio Seazza valley (Udine, Italy). Gortania 10: 121–132.
Holgado B, Dalla Vecchia FM, Fortuny J, Bernardini F and Tuniz C 2015. A Reappraisal of the Purported Gastric Pellet with Pterosaurian Bones from the Upper Triassic of Italy. PLoS ONE 10(11): e0141275. doi:10.1371/journal.pone.0141275
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Renesto S, Dalla Vecchia FM, Peters D. 2002. Morphological evidence for bipedalism in the Late Triassic prolacertiform reptile Langobardisaurus. In: Gudo M, Gutmann M, Scholz J, editors. Concepts of functionalengineering and constructional morphology: biomechanical approaches on fossil and recent organisms. Senckenb Lethaea 82(1): 95–106.

 

News on the Origin of Pterosaurs on YouTube

I just uploaded a pterosaur origins video on YouTube. Click here to view it.

Click to view this "Origin of Pterosaurs" video on YouTube.

Click to view this “Origin of Pterosaurs” video on YouTube. 17 minutes long. 

Tanytrachelos in New Mexico – taxon exclusion problems

Modified June 09, 2015 with the addition of clades named by Peters 2000 overlooked by Pritchard et al. 2015. 

Pritchard et al. (2015)
report on 3D Tanytrachelos (Fig. 1) individual bones from New Mexico (Late Triassic, Chinle Formation). And I think they’re spot on with regard to bone identification.

Tanytrachelos

Figure 1. Tanytrachelos – close to Tanystropheus, but tiny with a distinct skull. The New Mexico material matches both in shape and size this North Carolina and Virginia material from the Late Triassic.

The problem comes from their phylogenetic analysis.
From the Pritchard et al. text: “Our analysis incorporated a range of fossil taxa that have traditionally been allied with Tanystropheus and Macrocnemus.” Unfortunately that tradition is ‘bogus’ based on the larger taxon list of the large reptile tree in which macronemids and tanystropheids are lepidosauromorphs, not archosauromorphs. In the Pritchard et al. cladogram (Fig. 2) note the separation of Prolacerta and Protorosaurus to make room for a “by default clade” of tanystropheids that should nest within the Lepidosauromorpha when more taxa are added. This abbreviated taxon list and “by default clade” actually separates the two prorotorosaurs from each other.

Figure 2. Cladogram from Pritchard et al. nesting tanystropheids between two protorosaurs, which should have nested together.

Figure 2. Cladogram from Pritchard et al. nesting tanystropheids between two protorosaurs, which should have nested together (Fig. 3 and large reptile tree). Lepidosauromorphs are in yellow. Archosaurmorphs are in white.

A subset
of the large reptile tree taxon list (Fig. 3) matched (as closely as possible) to the Pritchard et al taxon list demonstrates the problems with such a short taxon list using these taxa in which archosauromorphs and lepidosauromorphs are shuffled like a deck of cards. And sister taxa do not resemble one another at each color-shift clade. For instance, in figure 2 Macrocnemus does not resemble Mesosuchus and in figure 3 Macrocnemus does not resemble Petrolacosaurus.

Figure 3. Subset of the large reptile tree matched to the Pritchard et al. taxon list. Here Protorosaurus and Prolacerta nest together, but the other clades interweave archosauromorphs and lepidosauromorphs.

Figure 3. Subset of the large reptile tree matched to the Pritchard et al. taxon list. Here Protorosaurus and Prolacerta nest together, but the other clades interweave archosauromorphs (white) and lepidosauromorphs (yellow). If you think this cladogram needs more taxa, you are right. And you can find them at the large reptile tree. And this cladogram proves there is a limit to taxon exclusion past which the tree topology breaks down. But then, we are cherry-picking here from two widely separated clades.

Traditions need to be tested
The Pritchard et al. studied relied on a traditional taxon list that was falsified four years ago here. So why did the referees let this manuscript get published? (Answer: tradition, status quo, established paradigm, plus shunning, marginalizing, ignoring the larger gamut study).

The sternum issue
Tanystropheids, like all tritosaurs and most squamates (but not Adirosaurus through snakes because the forelimbs are shrinking), have a sternum not found in protorosaurs and other archosauromorphs. I know I just pulled a “Larry Martin” by noting one and only one trait…

So,
below is the list of all the other traits from the large reptile tree that unambiguously separate tanystropheids (T) from protorosaurs (P). There are 30. Many of these traits extend to other tritosaurs (a subset of the Lepidosauria) and are not found in other archosauromorphs or vice versa.

  1. ventral naris: T = chiefly mx; P = chiefly pmx
  2. dorsal nasal shape: T = pmx invasion; P = narrows toward naris
  3. pmx orientation: T = horizontal; P = down
  4. naris placement: T = displaced or elongated; P = snout tip
  5. posterolateral pmx: T = absent; P = narrower than naris
  6. frontal/parietal suture: T = straight and > than nasal suture; P = not
  7. frontal shape: T = wider posteriorly; P = not
  8. frontal posterior process: T = absent; P = present
  9. postparietals: T = absent; P= present
  10. tabulars: T = absent; P = present
  11. friontal/nasal suture: T = anteriorly oriented; P = zigzag
  12. quadratojugal presence: T = jugal ramus only; P = quadrate ramus only
  13. squamosal/quadratojugal indent: T = no qj ascending process; P = semicircle
  14. parietal and frontal fusion: T = both fused; P = no fusion
  15. pterygoid lateral edge: T = ectopterygoid continues margin laterally; P = sharp angle
  16. pterygoid shape: T = narrow; P = broad triangular
  17. procumbent pmx teeth: T = present; P = absent
  18. posterior mandible shape: T = deeper anteriorly; P = mid rise
  19. caudal transverse processes: T = absent beyond 8th caudal; P = present beyond
  20. short lumbar ribs: T = present; P = not short
  21. second sacral rib: T = not bifurcate; P = bifurcate
  22. chevron shape: T = parallel to centra; P = descends, distal wider
  23. anterior caudal spines: T = shorter than centra; P = taller than centra
  24. sternum: T = present; P = absent
  25. scapula shape: T = not robust; P = robust
  26. pubic apron: T = not present; P = present and wide
  27. tarsus: T = not fenestrated; P = fenestrated
  28. calcaneal tuber: T = no tuber: P = lateral tuber
  29. metacarpal 5: T = straighter or twisted: P = hooked
  30. pedal 3.1 longer than p2.1: T = present; P = not

This reference probably snuck under the radar
Pritchard et al. noted several unnamed clades that were actually named in Peters 2000, some 15 years ago. Further work with the large reptile tree has shown that these clades are all lepidosaurian, not archosaurian or protorosaurian.

Clades named by Peters 2000

Tapinoplatia
Macrocnemus + Characiopoda

Characiopoda
Tanystropheidae + Langobardisaurus + Fenestrasauria

Fenestrasauria
Cosesaurus + Sharovipteryx + Longisquama + Pterosauria

References
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Pritchard AC, Turner AH, Nesbitt SJ, Irmis RB and Smith ND 2015. Late Triassic tanystropheids (Reptilia, Archosauromorpha) from northern New Mexico (Petrified Forest Member, Chinle Formation) and the biogeography, functional morphology, and evolution of Tanystropheidae. Journal of Vertebrate Paleontology 25(2):e911186 (20pp).