Testing for bipedalism in archosaurs (and pterosaurs)

Grinham, VanBuren and Norman 2019
looked at the origin of bipedalism in the archosaur and pre-archosaur ancestors of birds.

They report, “We test whether facultative bipedality is a transitionary state of locomotor mode evolution in the most recent early archosaur phylogenies using maximum-likelihood ancestral state reconstructions for the first time. Across a total of seven independent transitions from quadrupedality to a state of obligate bipedality, we find that facultative bipedality exists as an intermediary mode only once, despite being acquired a total of 14 times. We also report more independent acquisitions of obligate bipedality in archosaurs than previously hypothesized, suggesting that locomotor mode is more evolutionarily fluid than expected and more readily experimented with in these reptiles.”

The authors used the cladograms of Ezcurra 2016 and Nesbitt 2011,
both of which are riddled with inappropriate taxon inclusion and exclusion problems as reported earlier here and here. Therefore comparisons regarding the number of times obligate bipedality in archosaurs occurred is useless lacking a consensus phylogenetic contaxt. In the large reptile tree (LRT, 1542 taxa) bipedality occurs only once in archosaurs. It just precedes the origin of the archosaurs (crocs + dinos only). Ezcurra, Nesbitt and Grinham et al. include a long list of inappropriate taxa in their inclusion set according to the LRT that skews results (e.g. the lepidosauromorphs: Jesairisosaurus, Macrocnemus, Mesosuchus, Gephyrosaurus, Planocephalosaurus, Eudimorphodon, Dimorphodon).

Grinham, VanBuren and Norman 2019
follow Nesbitt 2011 who listed the pterosaurs Eudimorphodon and Dimorphodon as archosauriforms. Grinham et al. 2017 considered both to be quadrupeds without explanation. The only pterosaur paper cited by Grinham et al. is Padian 2008. Peters 2007 recovered pterosaurs with lepidosaurs like Huehuecuetzpalli, later validated, expanded and published online in LRT. Peters 2000, 2011 reported on bipedal pterosaur tracks and restricted most cited pterosaur ichnites to flat-footed beach-combing pterosaur clades. Use keyword “bipedal pterosaur tracks” in the SEARCH box to see prior samples of digitigrade and bipedal tracks reported by this blogpost along with their citations.

Padian 2008 reported
“Peters (2000) also reached the conclusion that pterosaurs were not ornithodirans, and found instead that they were nested within what is traditionally considered the Prolacertiformes. It remains to be seen whether other workers can duplicate this result, but a recent analysis by Hone and Benton (2007) failed to find support for Peters’ analyses. For the present, because five different analyses have found that pterosaurs are ornithodirans, and the systematic community seems to have largely accepted this, the present paper will proceed with this provisional conclusion, without discounting other possible solutions.”

We looked at the bogus results
of Hone and Benton 2007 earlier here. They dropped taxa proposed as pterosaur ancestors by Peters 2000 because their inclusion would have tilted their supertree toward the topology recovered by Peters 2000, who tested four previously published cladograms by adding novel taxa to them. One year earlier than Peters 2000, co-author Benton 1999 had proposed Scleromochlus as a pterosaur sister/ancestor, which Peters 2000 invalidated. Evidently professor Benton did not appreciate that and succeeded, at least in Padian’s eyes, to dismiss Peters 2000 as an unacceptable and suppressible minority view.

Note that none
of Padian’s “five different analyses” used novel taxa proposed by Peters 2000. Padian’s report, “The systematic community seems to have largely accepted this,” demonstrates that Padian and his community were adverse to testing the novel taxa of Peters 2000 on their own terms, preferring the cozy comfort of tradition and orthodoxy — and they did this after Peters 2000 invalidated earlier efforts simply by adding a few taxa. Very easy to do. Even today it remains impossible to explain the origin of pterosaurs as archosaurs in a phylogenetic context because they are not archosaurs. In the world of academics, taxon exclusion remains a useful tool. We should all fight against this practice.

Later Padian 2008 reports, 
“Alternatively, if we consider that pterosaurs evolved from quadrupedal basal archosauromorphs such as Prolacertiformes (Peters, 2000), a rather different model of limb evolution must be proposed. In prolacertiforms the humerus is longer than the forearm and the femur is longer than the tibia; the glenoacetabular length is also long, as in most terrestrial quadrupeds. To attain the proportions seen in basal pterosaurs, the relative lengths of humerus and forearm and of femur and tibia would have to have been reversed, and the vertebral column would have had to shorten considerably (or the limb segments increase). These changes are independent of the extensive reorganization of the joints for erect posture and parasagittal gait, for which there is no evidence so far in prolacertiforms.”

Note: Padian 2008 chose to ignore the limb proportions
of Longisquama (Figs. 1, 2) another taxon proposed by Peters 2000 with a humerus shorter than the forearm, as in pterosaurs. He also ignored Sharovipteryx, another taxon proposed by Peters 2000, with a femur shorter than the tibia. In the world of academics, taxon exclusion remains a useful tool. We should all fight against this.

Padian 2008 also chose to ignore the evidence for bipedalism
in Cosesaurus (Fig. 2) matching facutatively bipedal Rotodactylus tracks (Peters 2000) and Sharovipteryx (Fig. 2), an obligate biped based on proportions. Both have the short torso relative to the limb length sought for and purposefully overlooked by Padian 2008 (see above quotation). In the world of academics, taxon exclusion remains a useful tool. We should all fight against this.

Students of paleontology:
I’m sorry, this is just the way it is.

Getting back to bipedalism in archosaurs,
the LRT, subset Fig. 4) documents the patterns and possibilities of bipedal locomotion in taxa preceding dinosaurs. The topology here employs more taxa, pushes pterosaurs over to lepidosaurs (Peters 2007) and nests only Crocodylomorpha + Dinosauria within the Archosauria. Poposauria is the proximal outgroup. This is where bipedalism in archosaurs first appeared. Other bipedal taxa achieved this ability by convergence. Secondary quadrupedalism occurred several times in archosaurs, and by convergence in certain derived pterosaurs (e.g. ctenochasmatids and azhdarchids), as evidenced by their backward pointing manual digit 3 in ichnites.

As documented here and elsewhere
It does not matter if certain hypotheses are peer-reviewed and published or not.
Academic authors can choose to omit pertinent taxa and papers knowing that ‘friendly’ academic referees and editors will likewise choose to overlook such omissions. Apparently all academics seek and work to maintain the orthodox line, no matter how invalid it may be.

That’s why this blogpost and ReptileEvolution.com came into being.
We’re talking about hard science. Ignoring and omitting hard evidence cannot be tolerated or coddled. I ask only that academic workers rise to the professionalism they seek to inspire in their own students. History will put this all into perspective. Professional legacies may end up in shame unless they take action soon. Just test the taxa. 

References
Benton MJ 1999. Scleromochlus taylori and the origin of the pterosaurs. Philosophical Transactions of the Royal Society London, Series B 354 1423-1446. Online pdf
Ezcurra MD 2016 The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4, e1778. (doi:10.7717/peerj.1778)
Grinham LR, VanBuren CS and Norman DB 2019. Testing for a facultative locomotor mode in the acquisition of archosaur bipedality. R. Soc. open sci. 6: 190569. http://dx.doi.org/10.1098/rsos.190569
Hone DWE and Benton MJ 2007. An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles. Journal of Systematic Palaeontology 5:465–469.
Hone DWE and Benton MJ 2008. Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.
Nesbitt SJ 2011. The early evolution ofArchosaurs: relationships and the origin of major clades. Bull. Am. Museum Nat. Hist. 352, 1–292. (doi:10.1206/352.1)
Padian K 2008. Were pterosaur ancestors bipedal or quadrupedal? Morphometric,
functional, and phylogenetic considerations. Zitteliana R. B Abhandlungen der Bayer.
Staatssammlung fur Palaontologie und Geol. 28B, 21–28.
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods.  Ichnos 7:11-41.
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 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. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27
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