Bipedal crocodylomorph (or giant pterosaur tracks?) from Korea

Kim et al. 2020 describe
sets of 18-24cm narrow-gauge tetradactyl (four-toed) bipedal tracks from the Early Cretaceous (Aptian?) coast of South Korea they name Batrachopus grandis (Figs. 1, 2) a new ichnospecies. The authors attribute the tracks to a large (3m) crocodylomorph. They also note: “Surprisingly, the trackways appear to represent bipedal progression which is atypical of all known smaller batrachopodid trackways.”

You might find their logic train interesting. (See below.)

By the way, such narrow-gauge tracks (Fig. 2) are also atypical for Cretaceous crocs and azhdarchid pterosaurs, like the Late Cretaceous trackmaker of the ichnospecies, Haenamichnus (Figs. 2, 3).

On the other hand, basalmost Triassic crocs were all narrow-gauge bipeds. None of these were large (but that can change), plantigrade (but that can change) or left tracks (that we know of).

Such narrow-gauge tracks were also typical for strictly bipedal pterosaurs, like the coeval (Early Cretaceous) Shenzhoupterus (Figs. 5–7), a taxon overlooked by Kim et al. 2018, 2020.

Figure 1. Batrachopodus grandis tracks from Kim et al. 2020. Note the digits are shorter than the metatarsals and the heel is half the maximum width of the foot, matching both Early Jurassic Protosuchus and coeval (Early Cretaceous) Shenzhoupterus.

Figure 2. Batrachopus tracks (2nd from left) compared to other croc tracks.

Figure 2. Batrachopodus tracks (2nd from left) compared to other croc tracks. Haenamichnus uhangriensis, azhdarchid quadrupedal pterosaur tracks shown at far right, with three fingered manus track, outside and slightly behind the oval (here at this scale) pedal track.

From the Kim et al. abstract:
“This interpretation helps solve previous confusion over interpretation of enigmatic tracks of bipeds from younger (? Albian) Haman formation sites by showing they are not pterosaurian as previously inferred. Rather, they support the strong consensus that pterosaurs were obligate quadrupeds, not bipeds.”

Consensus = current opinion. What you really want  and deserve is evidence! (…and not to overlook evidence that is already out there). Peters 2000, 2011 showed that many pterosaurs were bipedal. Specific beachcombing clades were quadrupedal secondarily.

Figure 2. The large azhdarchid pterosaur, Zhejiangppterus. is shown walking over large pterosaur tracks matched to its feet from Korea (CNUPH.p9. Haenamichnus. (Hwang et al. 2002.)

Figure 3. The large azhdarchid pterosaur, Zhejiangppterus. is shown walking over large pterosaur tracks matched to its feet from Korea (CNUPH.p9. Haenamichnus. (Hwang et al. 2002.)

Unfortunately,
basal crocodylomorph feet are almost entirely absent from the fossil record in tested taxa in the large reptile tree (LRT, 1697+ taxa). The only exception is bipedal and likely digitigrade, Terrestrisuchus (Fig. 4). We don’t get another complete set of toes for testing until quadrupedal and plantigrade Protosuchus (Fig. 4), a not so basal crocodylomorph, that had the slightly more gracile digit 4 common to all extant crocs. Digit 4 does not appear to be any more gracile than the other toes in the new South Korean tracks, but let’s overlook that trifle for the moment.

Figure 2. Same feet, reordered according to the large reptile tree. Only Terrestrisuchus and Protosuchus are croc-like archosaurs here. Poposaurs are basal dinosaurs.

Figure 4. Same feet, reordered according to the large reptile tree. Only Terrestrisuchus and Protosuchus are croc-like archosaurs here.

Perhaps that is why Kim et al. write:
“Lower Jurassic Batrachopus with foot lengths (FL) in the 2–8 cm range, and Cretaceous Crocodylopodus (FL up to ~9.0 cm) (Fig. 2) known only from Korea and Spain registered narrow gauge trackways indicating semi-terrestrial/terrestrial quadrupedal gaits. Both ichnogenera, from ichnofamily Batrachopodidae, have been attributed to Protosuchus-like semi-terrestrial crocodylomorphs.”

… with a wider-gauge quadrupedal track.

On that note: The type species for Batrachopus is much smaller, fleshy, quadrupedal, narrow-gauge, with pedal impressions just behind the much smaller manus impressions.

By the start of the Cretaceous all the earlier bipedal crocodylomorphs were extinct, according to the current fossil record. Shenzhoupterus, from China, was a nearby contemporary of the South Korean trackmaker with nearly identical feet and gait. Did I hear someone say, “Occam’s Razor“? Did someone mention, “taxon exclusion”?

Earlier Kim et al. 2012 described similar tracks
as pterosaurian. Back then they were matched here to a giant Shenzhoupterus (Figs. 5–7), a coeval (Aptian, Early Cretaceous) dsungaripterid relative found in nearby China, with forelimbs less likely to reach the ground. Later a partial skeleton of a giant Late Cretaceous pterosaur from France, Mistralazhdarcho (Vullo et al. 2018), was reidentified here as a giant shenzhoupterid, rather than an azhdarchid. So shenzhoupterids were not restricted in size.

Kim et al report on, “Distinguishing crocodilian from pterosaurian trackways.”
“An unexpected result of the discovery of B. grandis trackway has been to shed light on a the controversial issue of pterosaur locomotion debated since the 1980 and 1990s: were pterosaurs bipedal or quadrupedal?

The answer is some were bipedal. Others were quadrupedal (Peters 2000, 2011, not cited by Kim et al.). It all depends on the clade and their niche.

Kim et al continue:
“These debates, mainly concerning relatively small pterosaurian tracks, have largely been resolved in favor of quadrupedalism.

Largely? Does that mean Kim et al. recognize exceptions? If so, they were not cited. More importantly, look for any other distinguishing traits in what follows from the Kim et al. text.

Kim et al continue:
“However, some uncertainty remained regarding tracks of purported ‘giant’ pterosaurians that were described as ‘enigmatic’ and inferred to have progressed bipedally (Kim et al. 2012). These trackways from the Lower Cretaceous, Haman Formation, at the Gain-ri tracksite, Korea were named Haenamichnus gainensis and inferred to represent, large, plantigrade pterodactyloid pterosaurs that might have walked bipedally so that the long wings did not become mired in the substrate. It was further inferred they may have been wading in shallow water.”

amples from the Lower Cretaceous, Gain, Korea trackway

Figure 5. Samples from the Lower Cretaceous, Gain, Korea trackway (left) along with original tracings of photos, new color tracings of photos with hypothetical digits added in red, then candidate trackmakers from the monophyletic Shenzhoupterus/Tapejarid clade.

Estimating Gain pterosaur trackmakers from track sizes and matching taxa.

Figure 6. Estimating Gain pterosaur trackmakers from track sizes and matching taxa. Note the Shenzhoupterus manus is a wee bit too short to touch the substrate as in Tupuxuara and many other derived pterosaurs.

Figure 1. Mistralazhdarcho compared to reconstructions of Shenzhoupterus and Nemicolopterus.

Figure 7. Mistralazhdarcho compared to reconstructions of Shenzhoupterus and Nemicolopterus.

Kim et al continue:
“We can now confirm confidently, that these tracks from the Gain-ri tracksite and others from Adu Island: are identical to poorly preserved large Batrachopus trackways. Thus, they should be removed from Haenamichnus and regarded as large poorly preserved batrachopodid tracks. The type specimen then tech- nically becomes Batrachopus gainensis (comb nov.). Thus, H. gainensis becomes a footnote to ichnotaxonomic history, shown to be an extramorphological expressions large of Batrachopus, only recognizable retrospectively after comparison with B. grandis. Therefore ichnologists may retrospectively choose to regard H. gainensis as a nomen dubium, and find little value in the trival name (gainensis). Alternatively they may simply refer to the Haman Formation tracks as Batrachopus cf. grandis.

Taken on its face, this is a rare instance of a paleontologist admitting a mistake. The other option is: both tracks are pterosaurian. So far, as you’ll note, the authors have not pointed to any factors, other than ‘bipedalism’, that would dissuade a pterosaurian trackmaker interpretation. I will admit and you can see (Figs. 4–5) that the pedes of Protosuchus and Shenzoupterus are rather close matches when covered with pads.

Kim et al continue:
“Note that the Gain-ri and Adu island trackways are from the Haman Formation and so these occurrences indicate a widespread distribution in space (three sites) and time (two formations) of this distinctive apparently bipedal morphotype. The pes tracks from the two Haman Formation sites are also larger (27.5–39.0 cm long), but with trackway proportions (step, stride, pace angulation etc.,) quite similar to those from the Jinju Formation.”

“The identification of the Haman Formation trackways as poorly preserved large batrachopodid tracks apparently suggests that the trackmakers habitually progressed bipedally. Alternatively the same speculative arguments for apparent rather than real bipedalism would have to be invoked as was the case with the Jinju material. Moreover, in almost all cases the trackways are very narrow gauge with a narrower straddle than seen in modern crocodylians. It is also of interest that least five subparallel more or less equally spaced trackways were registered on the level 4 surface. This suggests either that the trackmakers may have been gregarious, or that they were following a physically controlled route, such as a shoreline, defined by the paleoenvironment.”

Still no distinguishing traits, other than bipedalism, according to the authors. And note, they never considered the coeval and neighboring pterosaur, Shenzhoupterus, which is also a close match for the new tracks. They chose to invent a croc trackmaker rather than consider a pterosaurian trackmaker, evidently bowing to the consensus (their word, not mine, see above) and to follow Dr. Bennett’s curse and keep their blinders on. I wish they had dived deeper into the literature and evidence instead of following the crowd.


References
Hwang KG, Huh M, Lockley MG, Unwin DM and Wright JL 2002. New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea. Geology Magazine 139(4): 421-435.
Kim, JY et al. 2012. Enigmatic giant pterosaur tracks, and associated ichnofauna from the Cretaceous of Korea: implications for bipedal locomotion of pterosaurs. Ichnos 19, 50–65 (2012).
Kim KS, Lockley MG, Lim JD, Bae SM and Romilio A 2020. Trackway evidence for large bipedal crocodylomorphs from the Cretaceous of Korea. Nature Scientific Reports 10:8680 | https://doi.org/10.1038/s41598-020-66008-7
Lockley, MG et al. 2020. First reports of Crocodylopodus from Asia: implications for the paleoecology of the Lower Cretaceous.Cretaceous Research (2020) (online, March 2020).
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods. Ichnos, 7: 11-41
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
Vullo R, Garcia G, Godefroit P, Cincotta A, and Valentin X 2018.
 Mistralazhdarcho maggii, gen. et sp. nov., a new azhdarchid pterosaur from the Upper Cretaceous of southeastern France. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1502670.

https://pterosaurheresies.wordpress.com/2012/03/25/giant-bipedal-pterosaur-tracks-from-korea/

https://pterosaurheresies.wordpress.com/2018/10/19/mistralazhdarcho-a-new-pterosaur-but-not-an-azhdarchid/

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