Bipedal archosaur locomotion: Bates and Schachner 2011

Bates and Schachner 2011
report on bipedal archosaur locomotion with an emphasis on the basal poposaur, Poposaurus (Fig. 1).

Figure 1. Poposaurus skeleton and skull. Proportions indicate bipedal configuration.

Unfortunately,
Poposaurus and the Poposauridae (Fig. 2) nest just outside the Archosauria in the LRT (Fig. x). The basal croc, Pseudhesperosuchus, and the basal dinosaur, Herrerasaurus (Fig. 3), are valid candidates IF you want to stick to present definitions for the Archosauria.

Figure 2. Poposauridae revised for 2014. Here they are derived from Turfanosuchus at the base of the Archosauria, just before crocs split from dinos.

We need a new name
for the unnamed clade Poposauria + Archosauria: Huperarchosauria (“more than Archosauria”). By just changing the title of Bates and Schachner 2011 to “Disparity and convergence in bipedal huperarchosaur locomotion,” the use of Poposaurus (Fig. 1) as an example becomes valid.

Figure 3. The origin of dinosaurs in the LRT to scale. Gray arrows show the direction of evolution. This image includes Decuriasuchus, Turfanosuchus, Gracilisuchus, Lewisuchus, Pseudhesperosuchus, Trialestes, Herrerasaurus, Tawa and Eoraptor.  Note the phylogenetic miniaturization at the origin of Archosauria (Crocs + Dinos).

From the introduction:
“The clade Archosauria contains a staggering level of morphological, functional and ecological diversity that includes living birds and crocodilians, in addition to an array of enigmatic extinct forms such as dinosaurs and pterosaurs.”

Not pterosaurs. Those have nested apart from archosaurs for the last 20 years (Peters, 2000–2011). Over and over taxon exclusion prevents Bates and Schachner 2011 from understanding the phylogenetic context of their subject matter. For a more complete understanding of archosaur interrelations see the large reptile tree (LRT, 1734+ taxa; subset Fig. x).

Pterosaur and related fenestrasaur bipedalism, based on sprawling lepidosaur hind limbs, was not part of the Bates and Schachner study. Rather they concentrated on the erect hind limb bones and hypothetical muscles in Poposaurus and similar dinosaurs and birds.

Figure x. Subset of the LRT focusing on Archosauriformes. Clade colors match figure 2 overlay.

Add taxa to discover 
all the clade members within the Archosauria (= birds + crocs, their last common ancestor and all descendants). In the LRT Archosauria includes crocs + dinosaurs and nothing more. Poposaurs are the proximal outgroup. Pterosaur nest elsewhere, within Lepidosauria, far from these archosauriform taxa.

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Addendum:
There were 10x more views of the recent post on bat origins than the next most popular blogpost this week. I hope these ‘bat’ blogposts help us all understand the transition of arboreal mammals to flapping flight.

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References
Bates KT and Schachner ER 2011. Disparity and convergence in bipedal archosaur locomotion. Journal of The Royal Society Interface 9(71):1339–1353.
Farlow JO, Schachner ER, Sarrazin JC, Klein H and Currie PJ 2014.Pedal Proportions of Poposaurus gracilis: Convergence and Divergence in the Feet of Archosaurs. The Anatomical Record. DOI 10.1002/ar.22863
Gauthier JA, Nesbitt SJ, Schachner ER, Bever GS and Joyce WG 2011. The bipedal stem crocodilian Poposaurus gracilis: inferring function in fossils and innovation in archosaur locomotion. Bulletin of the Peabody Museum of Natural History 52:107-126.
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods. Ichnos 7:11-41.
Peters D 2000b. A reexamination of four prolacertiforms with implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293–336.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. – Historical Biology 15: 277–301.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29: 1327-1330.
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification
Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605

 

https://pterosaurheresies.wordpress.com/2011/07/16/what-exactly-is-a-pterosaur-part-3-of-3/

Pisanosaurus: dinosaur or silesaurid?

A new paper by Agnolin and Rozadilla 2017
includes new photographs of the holotype that shed new light on Pisanosaurus (Casamiquela 1967, Bonaparte 1976; Late Triassic). This taxon was previously known in the literature chiefly (not exclusively) from drawings. The large reptile tree (LRT, 1043 taxa) nested Pisanosaurus with Haya as a basal ornithischian, confirming prior assessments. Now Agnolin and Rozadilla provide evidence for a Silesaurus affinity among the Poposauridae. Echoing others, they report, “the poor preservation of the specimen is the largest difficulty to overcome when interpreting its morphology. Its phylogenetic position within ornithischians is problematic.”

So, with the new evidence,
let’s test and nest Pisanosaurus 2017! (There are so few traits that can be scored for Pisanosaurus, that the rest of the discussion might seem like I’m pulling a Larry Martin. That happens sometimes, but I’m trying to report results from the LRT.

Before we start…
with present data, shifting Pisanosaurus to Silesaurus in the LRT adds 24 steps. Moreover, Agnolin and Rozadilla did not mention the proximal relatives of Pisanosaurus in the LRT:  Haya, Daemonosaurus, Chilesaurus, Scelidosaurus and Emausaurus. This may be the key to their novel results: taxon exclusion… once again. 

Some general notes to start with:

1. Silesaurus and other poposaurs have a metatarsus no longer than the longest digit. The same hold true for many basal phytodinosaurs, but Pisanosaurus has a longer metatarsus, like its sister in the LRT, Haya.
2. The photo of the pelvis does little to clarify any issues. It is a broken up mess (Fig. 2) with, what appear to be smaller pelvis bones (greens)  and several sacral bones (blues) stirred up in a conglomeration. Not much matches the published drawings. And my earlier imagination describing a rotated pubis based on simple published drawings did not pan out.
3. The anterior dentary appears to be missing a predentary bone, a trait common to the clade Ornithischia, but something like it also appears in Silesaurus.
4. Pisanoaurus comes from South America, home of most of the other basalmost Triassic phytodinosaurs. Popposaurids, all except Sacisaurus, come from somewhere else on the globe. Haya, the LRT sister to Pisanosaurus, comes from China, but it is Late Cretaceous in age.
5. Agnolin and Rozadilla consider Silesaurus part of a clade “that is currently recognized as the sister group to Dinosauria.” The LRT recovers Crocodylomorpha closer to Dinosauria and Silesaurus nests within the next proximal outgroup, Poposauridae.
6. Agnolin and Rozadilla report, “because Pisanosaurus is a unique and very valuable specimen, it is not currently possible to [CT] scan it.”
7. Authors have not agreed whether the pelvis, represented by fragments of bones and bone impressions in rock. is preserved in medial or lateral view. Agnolin and Rozadilla report, “the sacrum is articulated and preserved in life position with respect to the pelvis.”

Figure 1. The Pisanosaurus pelvis here flipped right to left along with drawings and reconstructions by Agnolín and Rozadilla, plus DGS colors applied to what I can see here. Other than the sacral vertebrate on top, not much is clear, but it seems like the pelvic elements are smaller that published and that several sacral vertebrate are sprinkled in this mass. Perhaps a CT scan would be helpful here. Blue = vertebrae. Green = pelvi elements.

Agnolin and Rozadilla provided an emended diagnosis.
Pisanosaurus is a basal dinosaurifordiagnosable by the following autapomorphies:

1. “central teeth bilobate in occlusal view, showing well-developed mesial and distal grooves;
2. distal end of the tibia anteroposteriorly longer than transversely wide;
3. bilobate astragalus in distal view;
4. ascending process of the astragalus being subquadrangular and robust in lateral view;
5. intense transversal compression of the calcaneum.”

Figure 2. Skull of Haya and restored skull of Pisanosaurus compared. The resemblance of preserved elements is apparent here. In both cases the mandibular fenestra is filled in. The other holes in the Pisanosaurus mandible are artifacts of taphonomy. Pisanosaurus data from Irmis et al. 2007b.

Other factors of interest:

1. The number of tooth positions (15) in Pisanosaurus matches both silesaurids and pertinent ornithischians.
2. “Central teeth are bilobate in occlusal view, and show well-developed mesial and distal grooves, a condition unknown in other herbivorous taxa and a trait that may be an autapomorphy of Pisanosaurus.” Not sure if the teeth in Haya are the same, but they look similar in lateral view (Fig. 2). Neither have denticles. Silesaurid teeth are leaf-shaped.
3. “the teeth do not form a palisade or continuous masticatory surface as advocated by some authors.” As in Haya.
4. “Pisanosaurus is similar to saurischians and basal dinosauriforms in having overlapping proximal metatarsals, differing from the non-overlapping condition in ornithischians.” Except Haya.

Figure 3. Haya in lateral view. Note the dorsal laminae, similar to those in Pisanosaurus.

Agnolin and Rozadilla describe the dorsal vertebrae
as having a strong and complex system of laminae. Haya (Fig. 3).has similar laminae. Poposauridae do not.

Figure 4 Silesaurus as a biped and occasional quadruped. Note the squareish cervicals, unlike the parallelograms in figure 5.

Agnolin and Rozadilla considered the vertebrae
(Fig. 5) very different from the cervical vertebrae described for basal dinosauriforms and ornithischians. But they did not look at Haya, which has similar cervicals 1 and 2 (Fig. 5). They considered the cervicals ‘indistinguishable from Sacisaurus cervicals, but Langer and Ferigolo 2013, did not refer the cervical to Sacisaurus due to its relatively large size. Concluding Agnolin and Rozadilla considered these verts to be on uncertain position.

Figure 5. Pisanosaurus cervical vertebrae in left lateral view (not right as published) matches cervical vertebrae 1 and 2 in Haya – and does not match the simpler vertebrae in Silesaurus (Fig. 4).

Sacrals are preserved as moulds in Pisanosaurus. 
Various authors have interpreted five, to two sacrals. Agnolin and Rozadilla concurred with Irmis et al. 2007, who found no trace of sacral elements, reporting, “some features previously considered to be impressions of sacral ribs are actually cracks in the matrix, and there is insufficient fidelity to determine whether any of the centra are fused to each other.” 

Figure 6. Pisanosaurus right pes with digit 2 ghosted in and digit 4 rotated into in vivo position. PILS added. Nnte the brevity of the toes compared to the metatarsus, a trait shared with Haya.

Is the acetabulum open or closed?
Agnolin and Rozadilla ‘suggest’ it is closed, as in poposaurs. If so the closed portion is buried. With available evidence and phylogenetic bracketing, it was probably open. Haya has an acetabulum with a keyhole shape (Fig. 3).

The tibia, tarsus and metatarsus
in Pisanosaurus the cnemial crest does not peak at the knee, but somewhat lower. Haya is similar. The fibula diameter is 70% that of the tibia, as in Scelidosaurus. The fibula for Haya is unknown. Anolín and Rozadilla identified a calcaneal tuber. That is odd because it is so small that it does not extend as far as the fibula does. in Haya the calcaneum extends slightly beyond the astragalus. The astragalus of Pisanosaurus is longer than wide (when the medial condyle is included), which is distinctly different from Haya and other sister taxa and different from Silesaurus.

Figure 8. Calcaneum of Pisanosaurus. You can imagine why some authors saw a tuber while others did not.

A flawed phylogenetic analysis
Other than excluding several taxa that nest close to Pisanosaurus in the LRT, Agnolin and Rozadilla employed the invalid Nesbitt (2011) database, also suffering greatly from taxon exclusion. It does not nest sauropodomorphs with ornithischians as phytodinosaurs, but nests sauropodomorphs, like Pampadromaeus, with Tawa and other theropods. In their first analysis, 20 trees resulted with Pisanosaurus nested as an unresolved polytomy of several dinos and non-dinos. After excluding wild card taxa, 82 trees resulted with Pisanosaurus within the Silesauridae. Bremer support is low in their analysis, but Bootstrap support is high in the LRT.

Discussion
Agnolín and Rozadilla discuss several traits of Pisanosaurus (typically related to herbivory) and their appearances elsewhere within the Archosauria. They find no epipophyses in the cervicals, but Haya lacks these, as well on the pertinent first two verts. Agnolín and Rozadilla note “The vertebral centra are very elongate and transversely compressed, differing from the short and stout dorsal vertebrae of known ornithischians, including heterodontosaurids.” They do not realize the close relationship of Pisanosaurus to sauropodomorphs like Saturnalia and the basalmost ornithischian, Chilesaurus, both with elongate dorsals. Agnolín and Rozadilla made a “tentative reconstruction” of the pelvis (Fig. 1), but it bear little to no resemblance to the in situ fossil. In every comparison made, Agnolín and Rozadilla delete or ignore Haya and related taxa and thus recover semi-blind results.

Today and in the future
we can’t keep going back to the same short lists of taxa for our inclusion sets. We know of so many more now that need to be included in phylogenetic analyses. The LRT can be your guide.

References
Agnolín FL and Rozadilla S 2017. Phylogenetic reassessment of Pisanosaurus mertii Casamiquela, 1967, a basal dinosauriform from the Late Triassic of Argentina. Journal of Systematic Palaeontology. http://dx.doi.org/10.1080/14772019.2017.1352623
Ferigolo and Langer 2006. A Late Triassic dinosauriform from south Brazil and the origin of the ornithischian predentary bone. Historical Biology, 2006; 1–11, iFirst article
Nesbitt SJ 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History, 352, 1–292.

wiki/Sacisaurus
wiki/Pisanosaurus
wiki/Haya

 

Effigia palate and occiput

Updated March 13, 2015 with a new palate figure based on the photo, not the original drawing in Effigia and Shuvosaurus was modified in turn.

We looked at the poposaur Effigia earlier here and here. Having never attempted a reconstruction of the palate I do so here.

Figure 1. Effigia. is an odd derived poposaur with tiny hands and no teeth. In competition with dinosaurs, poplars did not fare as well. The dentary and predentary have been modified here from prior attempts to more closely match the mandible of Shuvosaurus (Fig. 3).

Effigia okeeffeae (Nesbitt and Norell 2006, Nesbitt 2007) Carnian, Late Triassic, ~210 mya, ~ 2 m in length, was originally considered an early theropod dinosaur by Colbert, who collected the specimen in the late 1940s but never removed it from its jacket. A recent reassessment by Nesbitt and Norell (2006) and Nesbitt (2007) nested Effigia among the poposauridis. It is an odd bipedal poposaur and perhaps the most derived member of a clade composed almost entirely of odd derived members. The reconstruction of the skull has been controversial. Perhaps only a direct tracing and shifting of the elements can solve this puzzle. All the pieces in the disarticulated fossil will come together precisely if they are correctly reassembled.

The palate 
It is possible that the palatine (Fig. 2). was misidentified originally as the right ectopterygoid. If so, then the palate resembles that of known sister taxa, like Shuvosaurus (Fig. 3)..

Figure 2. Effigia palate in situ (left) and reconstructed by reassembling colored elements (at right). Click to enlarge.

Due to the long premaxilla
and the short maxilla the Effigia palate shifts most of the palatal elements into a smaller space. Even so all maintain their original and typical connections to the other skull elements.

Figure 3. Shuvosaurus, a sister to Effigia, has a similar palate in this reconstruction, but it was not reconstructed like this originally.

You really can’t talk about
the palate of Effigia without comparing it to its sister, Shuvosaurus (Fig. 3). Here the main triangular part of the pterygoid must be imagined, but the quadrate processes are present and quite robust. The palatines frame the internal nares posterior to the palatal processes of the maxilla and premaxilla.

Figure 4, the occiput of Effigia colorized here to segregate elements. That’s the central supraoccipital in pink flanked by two opisthotics in lavender, all displace dorsally. Originally they were framed by the squamosals in gold. Quadrates in red and basisphenoid in purple.

Effigia occiput
The above image (Fig. 4, Nesbitt 2007) is a CT scan of the Effigia occiput colorized to aid identification of the elements. The occiput is so inclined it is almost continuous with the palate. Originally the supraoccipital + opisthotics were identified as the two parietals with no median element recognized. Neither the supraocipital or the opisthotic were identified otherwise.

Effigia References
Nesbitt SJ and Norell MA 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B 273:1045–1048. online
Nesbitt S 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, 302: 84 pp. online pdf
AMNH Effigia webpage
wiki/Effigia

Shuvosaurus References
Alcober O, Parrish JM. 1997. A new poposaurid from the upper Triassic of Argentina. Journal of Vertebrate Paleontology 17:548–556
Brusatte SL, Benton MJ, Desojo JB and Langer MC 2010. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida), Journal of Systematic Palaeontology, 8:1, 3-47.
Chatterjee S 1991. An unusual toothless archosaur from the Triassic of Texas: the world’s oldest ostrich dinosaur? Abstract, Journal of Vertebrate Paleontology, 8(3): 11A.
Chatterjee S 1993. Shuvosaurus, a new theropod: an unusual theropod dinosaur from the Triassic of Texas. National Geographic Research and Exploration 9 (3): 274–285.
Rauhut OWM 1997. On the cranial anatomy of Shuvosaurus inexpectatus (Dinosauria: Theropoda). In: Sachs, S., Rauhut, O. W. M. & Weigert, A. (eds) 1. Treffen der deutschsprachigen Palaeoherpetologen, Düsseldorf, 21.-23.02.1997; Extended Abstracts. Terra Nostra 7/97, pp. 17-21.
Long R and Murry P 1995. Late Triassic (Carnian-Norian) Tetrapods from the Southwestern United States. New Mexico Museum of Natural History and Science Bulletin 4, Pp. 153-163.

wiki/Shuvosaurus

 

Not Arizonasaurus, but Postosuchus, made the giant Isochirotherium tracks

A recent paper by Diedrich (2015) purported to match the Arizonasaurus to giant Isochirotherium tracks from the Middle Triassic of Germany (Fig. 1).

The problem is,
no manus or pes are known for Arizonasaurus. Furthermore, all related taxa in the large reptile tree have digit 3 the longest, and all digits are elongate. The giant Isochirotherium tracks indicate that both digits 2 and 3 are the longest, and they are short. So matching candidates have to be found elsewhere, not close to Arizonoasaurus (although the size and time are right!).

Among the 504 taxa in the large reptile tree that are possible candidates with digits 2 and 3 the longest are Erythrosuchus (Fig. 1), Shansisuchus, Lotosaurus and the Postosuchus alisonae (Peyer 2008, Fig. 1). It turns out that only the latter is the best match when scaled up to the size of P. kirkpatrchicki (Chatterjee 1985, Fig. 1).

Figure 1. Giant Isochirotherium tracks matched to Postosuchus alisonae scaled up to the size of P. kirkpatrcki. Click to enlarge. This taxon was not considered originally because it is Late Triassic and the tracks are Middle Triassic.

Postosuchus was not mentioned in the text
because Diedrich (recent email) knew Postosuchus was Late Triassic, not Middle Triassic. He did not accept the idea that between the origin, radiation and extinction of Postosuchus there might have been a Middle Triassic relative.

Diedrich also saw the small manus tracks and assumed they were produced by a large poposaurid. Unfortunately, Arizonasaurus does not nest with poposaurids either. And poposaurids, other than Lotosaurus, do not match the track morphology.

It would have been helpful,
I suppose, to do what I did and make a list of possible candidates from a large list, AND THEN delete the possible candidates one by one as bad matches. Other than that phylogenetic bracketing mismatch, Diedrich does good work with excellent graphics. It took a leap of faith, I suppose to match tracks to a taxon for which no manus or foot is known.

C. Diedrich writes:
“Watch my ARTE docu – there you see Arizonasaurus (Ticinosuchus and Macrocnemus) walking in my point of view combining trak/sleketal records”:http://www.youtube.com/watch?v=b9GcVmb6OtE

References
Chatterjee S 1985. Postosuchus, a new Thecodontian reptile from the Triassic of Texas and the origin of Tyrannosaurs. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 309 (1139): 395–460. doi:10.1098/rstb.1985.0092.
Diedrich C 2015. Isochirotherium trackways, their possible trackmakers (?Arizonasaurus): intercontinental giant archosaur migrations in the Middle Triassic tsunami-influenced carbonate intertidal mud flats of the European Germanic Basin  Carbonates and Evaporites  DOI 10.1007/s13146-014-0228-z
Novak SE 2004. A new specimen of Postosuchus from the Late Triassic Coelophysis Quarry, siltstone member, Chinle Formation, Ghost Ranch, New Mexico. M.S. thesis, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina.
Peyer K Carter, JG, Sues H-D, Novak SE, and Olsen PE 2008. A new Suchian Archosaur from the Upper Triassic of North Carolina. Journal of Vertebrate Paleontology 28 (2): 363–381. doi:10.1671/0272-4634(2008)28[363:ANSAFT]2.0.CO;2.

Did Effigia have a postfrontal?

Applying color to the dorsal skull of Effigia appears to show unfused postfrontals (Fig. 1), a trait seen in archosaurs that are not crocs and not dinos (in other words: virtually all archosaurs). Very few basal crocs, like Gracilisuchus, retain an unfused postfrontal. Nesbitt (2007) did not identify a postfrontal but did identify an asymmetrical posterior nasal suture, which may not have been accurate.

Figure 1. Dorsal view of Effigia skull from Nesbitt 2007. At left postfrontals are identified. Bits of the broken nasals are also in that cranial area.

Earlier the poposaurs, including Effigia, shifted their position on the large reptile tree to nest with Turfanosuchus, a basal poposaur, which also has a postfrontal. A postfrontal has been reported on Lotosaurus (Parrish 1993), but I haven’t seen a good dorsal view of the skull. Poposaur skulls, and poposaur cranial regions are quite rare. This is one of the few examples.

References
Nesbitt SJ and Norell MA 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B 273:1045–1048. online
Nesbitt S 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, 302: 84 pp. online pdf
Parrish JM 1993. Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly. Journal of Vertebrate Paleontology 13:287–308
Zhang F-K 1975. A new thecodont Lotosaurus, from Middle Triassic of Hunan. Vertebrata PalAsiatica 13:144-147.

AMNH Effigia webpage
wiki/Effigia

The manus of Effigia revisited

The manus of Effigia is tiny.

Figure 1. Effigia. Note the tiny hands.

Really tiny.
Even so, it is one of the few manus examples from a wide range of post-Proterosuchus and pre-Herrerasaurus taxa. The Effigia manus (Figs. 1, 2) includes only 3 metacarpals and 2 phalanges, plus two carpals. Another metacarpal, one that looked like mc1 of Coelophysis and measuring 12×5.5 mm was described by Nesbitt (2007).

Not much to go on, but PILs and sister taxa (even distantly related) can help (Fig. 1) make the restoration — in this case two restorations: one if you switch the two longest metacarpals, one if you don’t. I think you should, to match another poposaurid manus, that of Poposaurus (Fig.1), which we looked at earlier here so metacarpals 1-3 align. In either case, tiny digit 5 appears to be missing, so the known digits are 2-4, not 3-5.

Figure 1. The manus of Effigia reconstructed. Metacarpal 1 was described with measurements. Here two possible hands can be restored, but only when one switches the longest metacarpals. That’s the one I think will prevail.

From Nesbitt (2007):
“Portions of the right manus were recovered somewhat articulated with both the radius and the ulna. Metacarpals III, IV, and V were found articulated together and a potential metacarpal I and the carpals were found together nearby. Unfortunately, the identity of the two carpals cannot be determined with certainty. The proposed metacarpal I proximal portion is broken, shifted ventrally, and appressed against the shaft. Metacarpal I is estimated to have been 12 mm long and 5.5 mm wide. It has a dorsoventrally compressed shaft and the distal end is asymmetrical; only the medial side bears a rounded articular surface; the lateral side tapers to a sharp edge. The morphology is strongly reminiscent of metacarpal I of Coelophysis and other theropods. Metacarpal II was not preserved.

“A much more robust phalanx is also present and may belong with either metacarpal III or metacarpal I. This phalanx indicates that at least one of the digits was not as reduced as much as digit I.”

If you shift the two longest metacarpals and add appropriate phalanges that fit the few established PILs, then you get a very poposaur-ish manus. If you don’t, you get an odd sort of manus, very autapomorphic. The carpals were matched to those of aetosaurs, Lotosaurus and Herrerasaurus, the closest known sisters with known carpals. If tiny distal carpals were present, they were not recovered.

On a side note,
the manus of Lotosaurus is looking ‘funky’ when compared to other popoosaurs. If anyone has good data on that piece of anatomy, please send it.

References
Nesbitt SJ and Norell MA 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B 273:1045–1048. online
Nesbitt S 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, 302: 84 pp. online pdf
wiki/Effigia

Is Nyasasaurus the oldest dinosaur?

Nesbitt et al. (2014) report, “Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister–taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs.”

Nesbitt et al. (2012) did not provide a reconstruction. The specimen is only known from a few Middle Triassic (Anisian) vertebrae and a proximal humerus. Here (Fig. 1) is a reconstruction that seems to fit pretty well based on the holotype and referred specimens of Nyasasaurus placed onto the bauplan of a large and slightly larger (for the cervicals) specimen of Turfanosuchus, a basal archosaur at the base of the Poposauridae (now nesting basal to archosaurs in the large reptile tree). This was, what you might call, an early ‘false start’ mimicking the actual rise of the Dinosauria.

Figure 1. Click to enlarge. Nyasasaurus bones placed on an enlargement of Turfanosuchus, a big. middle Triassic basal archosaur, not a dinosaur. Dinos and crocs all started out as tiny bipeds, also derived from Turfanosuchus, but smaller.

Nesbitt et al. ran their analysis and found Nyasasaurus to nest between 1) Lewisuchus and dinosaurs; 2) basal to Ornithischia; or 3) as the sister to Dilophosaurus, a Jurassic dinosaur. I didn’t run an analysis as my characters would not resolve relationships based on so few parts.

The large reptile tree finds two small specimens, Trialestes and an unnamed specimen incorrectly referred by Lecuona and Desojo (2011) to Gracilisuchus (PVL 1259) at the base of the Dinosauria. The latter is as old as Nyasasaurus, fulfilling chronological predictions.

Nesbitt et al. note a ventrally elongate deltopectoral crest on the humerus, but that assumes a short humerus. The crest is not so elongate if the humerus is a little longer.

Nesbitt et al. note three sacral vertebrae, but basal dinos don’t have three sacrals, only two. Turfanosuchus also has only two, but look at the size difference! Poposaurus, a sister taxon, has five sacrals. So Nyasasaurus is something else. Nesbitt et al. note hyposphene–hypantrum intervertebral articulations in the pre sacral vertebrae. Sorry, not much about that in Turfanosuchus data. In Turfanosuchus, as in Nyasaurus, the cervical vertebrae are laterally concave.

References
Lecuona A and Desojo, JB 2011. Hind limb osteology of Gracilisuchus stipanicicorum (Archosauria: Pseudosuchia). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 102 (2): 105–128.
Nesbitt SJ, Barrett PM, Werning S, Sidor CA and Charig AJ (posthumously) 2012. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biology Letters 9: 20120949.

 

 

Poposaurs – Topology Shift

Yes, I was wrong. And it’s time to man-up.
Earlier, based on available data, the large reptile tree nested poposaurids with phytodinosaurs and attributed the appearance and growth of the calcaneal tuber in certain poposaurs to convergence with the Crocodylomorpha. So the earlier data recovered poposaurs as dinosaurs with an odd ankle, not crocs with a “massive convergence” with dinosaurs, which is still the widespread hypothesis (see Nesbitt 2011 and others).

That seemed to make sense — except some poposaurs, like Lotosaurus and Asilisaurus (Fig. 1), appeared a little too early in the Triassic. They seemed to be anachronistic, and that can be a red flag.

So going back to the phylogenetic analysis,
I reexamined certain specimens, discovered a few items not originally presented (I trusted original tracings instead of making my own from in situ photos) and I found several bad scores. The newly recovered tree finds poposaurids derived from Turfanosuchus, a taxon that earlier stood alone at the base of the Archosauria (basal to crocs and dinos). Now things seem to make more sense, phylogenetically, chronologically (Fig.1) and morphologically. See if you agree…

Figure 1. Poposauridae revised for 2014. Here they are derived from Turfanosuchus at the base of the Archosauria, just before crocs split from dinos. Among these, only Silesaurus and Asilisaurus lost the calcaneal tuber.

Figure 2. The Euarchosauriformes featuring a new nesting for the Poposauridae.

Now the odd thing is: 
Poposaurs appear to provide a sort of preview to what would eventually evolve in the Dinosauria itself, likely filling similar niches in earlier strata.

Turfanosuchus and Poposaurus (Fig. 1) were convergent with theropods. The larger Asilisaurus was convergent with sauropodomorphs. The remainder were convergent with various ornithischians, even down to the toothless predentary they shared by convergence. Lotosaurus was a stegosaur mimic. Shuvosaurus was a Dryosaurus mimic. Silesaurus was a Camptosaurus mimic, down to losing the calcaneal tuber. Sacisaurus was a little Agilisaurus mimic. Effigia was still the oddball with those vestigial hands and back-sloped braincase.

So poposaurids are not dinosaurs. They are also not basal to rauisuchidae (contra Nesbitt 2011), but were derived from basal rauisuchia like Decuriasuchus and Vjushkovia. They are the most basal archosaurs. Basal poposaurs were the last common ancestors of crocs and birds. From their basalmost taxon, a sister to little Turfanosuchus, both tiny basal bipedal crocs and tiny bipedal basal dinos evolved.

Size
Poposaurs, in the form of Nyasasaurus, Asilisaurus and Lotosaurus (Fig. 1), were the first archosaurs to evolve substantial size in the Middle Triassic. Crocs and dinos remained small until the late Triassic (mid-Triassic for the basal Herrerasaurus) when they had their great radiation and poposaurs began to fade. This is an unrecognized faunal turnover.

Discovering and correcting errors is what scientists do. 
And I was happy that these new insights appeared.

What took so long?
Inattention to red flags. We should all look more closely at problems. They lead to new insights.

M.M. I hope this helps the cause. And yes, I have made and will make changes to earlier posts on this subject.

References
Brusatte SL, Benton MJ, Desojo JB and Langer MC 2010. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida), Journal of Systematic Palaeontology, 8:1, 3-47.
Irmis RB, Nesbitt SJ, Padian K, Smith ND, Turner AH, Woody D and Downs A 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317 (5836): 358–361. doi:10.1126/science.1143325. PMID 17641198.
Nesbitt SJ 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352: 292 pp.
Nesbitt SJ, Irmis RB, Parker WG, Smith ND, Turner AH and Rowe T 2009. Hindlimb osteology and distribution of basal dinosauromorphs from the Late Triassic of North America. Journal of Vertebrate Paleontology 29 (2): 498–516. doi:10.1671/039.029.0218

The manus of Poposaurus revised — again

 

Revised April 23, 2014 based on further study. 

Figure 1. Poposaurus manus as originally restored and with digit 1 switched to 5. Note the resemblance to dinosaur and basal croc hands in this basal archosaur.

The manus of basal archosaurs is very rare.
What few clues we have indicate that metatarsals 1-3 aligned distally and digit 5 is a vestige. Revising the manus of Poposaurus to that pattern is demonstrated here (Fig. 1).

References
Gauthier JA, Nesbitt SJ, Schachner ER, Bever GS and Joyce WG 2011. The bipedal stem crocodilian Poposaurus gracilis: inferring function in fossils and innovation in archosaur locomotion. Bulletin of the Peabody Museum of Natural History 52:107-126.
Mehl MG 1915. Poposaurus gracilis, a new reptile from the Triassic of Wyoming. Journal of Geology 23:516–522.

wiki/Poposaurus

A slightly new take on the new Poposaurus skull

More thoughts on the new Poposaurus skull (Fig. 1) that we looked at earlier (since modified). Slightly raising the broken maxilla ascending process puts a new spin on the possibilities for the skull shape and brings it into line with multi-documented sister taxa like Lotosaurus and Shuvosaurus.

Figure 4. The restored skull of the PEFO specimen referred to Poposaurus based on the Nesbitt identification of the anterior as a maxilla. The blue articular is not part of the PEFO specimen, but is described as a Poposaurus articular by Parker and Nesbitt (2013) scaled to fit. Their scale bars indicate it was 4x larger, which may be a typo. As is, the elements are part of a longer, more robust skull than any other poposaurid. See the revised skull reconstruction, figure 1B.

The teeth
The teeth are indeed sharp. Parker and Nesbitt 2013 described them as belonging to a hyper-carnivore. But in the large reptile tree Poposaurus nests with beaked herbivores. In the fossil the teeth really don’t descend very much beyond the jawline. They are deeply rooted and triangular, not long and recurved.

Other indicators
Earlier we noted the less than trenchant claws on the forelimb, more appropriate for an herbivore than a carnivore. The cervicals are quite robust, able to handle a taller skull. The new reconstruction, with a possible raised maxilla ascending process, opens the possibility of a taller skull with a larger orbit, also more like an herbivore and more like its sisters.

Figure 1. Revised skull reconstruction for the PEFO specimen. Here the anterior is considered a premaxilla. Those teeth are shaped like triangles, but they are very deeply rooted and exposed very little, which casts doubts on its hypercarnivory.

At this point it’s just best to explore possibilities, some indicated by phylogenetic bracketing. That skull remains quite incomplete.

References
Gauthier JA, Nesbitt SJ, Schachner ER, Bever GS and Joyce WG 2011. The bipedal stem crocodilian Poposaurus gracilis: inferring function in fossils and innovation in archosaur locomotion. Bulletin of the Peabody Museum of Natural History 52:107-126.
Mehl MG 1915. Poposaurus gracilis, a new reptile from the Triassic of Wyoming. Journal of Geology 23:516–522.
Parker WG and Nesbitt 2013. Cranial remains of Poposaurus gracilis (Pseudosuchia: Poposauroidea) from the Upper Triassic, the distribution of the taxon, and its implications for poposauroid evolution. Geological Society, London, Special Publications 379: 22 pp.

wiki/Poposaurus