Litorosuchus: a macrocnemid, not an archosauriform

Good to be getting back to non-mammalian reptiles again…

Li et al. 2016 describe
a Middle Triassic (240mya) semi-aquatic reptile,  Litorosuchus somnii based on a complete and articulated skeleton (Fig. 1) together with hard and soft dermal tissue (see below). Note the antorbital fenestra without a fossa. That’s a key trait that led the Li team astray.

Unfortunately 
Li et al. nested Litorosuchus with the armored and aquatic Vancleavea (Nesbitt et al 2009, Fig. 2), which they still insist is an aberrant stem archosaur (Nesbitt et al. 2009, Nesbitt 2011) despite lacking an upper temporal or antorbital fenestera, among a long list of other autapomorphies. Five years ago here Vancleavea was nested with Helveticosaurus in the Thalattosauria in the large reptile tree (LRT) where it remains today. Taxon exclusion burned the Nesbitt et al. 2009 study on Vancleavea and the Nesbitt 2011 study on archosaurs. Two of those team members (Nesbitt and Stocker) are also on the Li et al. 2016 team. Sadly for Science, Li et al. 2016 got burned again for the same reason. Pride.

Taxon exclusion
So certain that Litorosuchus was an archosauriform, no tritosaur lepidosaur lepidosauriform lepidosauromorphs were included in the Li et al. study matrix (based on Nesbitt et al. 2009 and additions thereafter) despite their overall resemblance to the new taxon (Fig. 3). In the LRT Litorosuchus nests strongly within Macrocnemus, (Fig. 3) another clade that has members with an antorbital fenestra, a trait that appears at least 4 times within the Reptilia.

Let’s take a look at the Li et al. abstract,
then discuss the macrocnemid affinities of the new taxon.

From the Li et al. abstract:
“Reptiles have a long history of transitioning from terrestrial to semi-aquatic or aquatic environments that stretches back at least 250 million years. Within Archosauria, both living crocodylians and birds have semi-aquatic members. Closer to the root of Archosauria and within the closest relatives of the clade, there is a growing body of evidence that early members of those clades had a semi-aquatic lifestyle [1]. However, the morphological adaptations to a semi-aquatic environment remain equivocal in most cases. Here, we introduce a new Middle Triassic (245–235 Ma) archosauriform, Litorosuchus somnii, gen. et sp. nov., based on a nearly complete skeleton from the Zhuganpo Member (Ladinian [241–235 Ma]) of the Falang Formation, Yunnan, China. Our phylogenetic analyses suggest [2] that Litorosuchus is a stem archosaur closely related to the aberrant Vancleavea just outside of Archosauria. The well-preserved skeleton of L. somnii bears a number of morphological characters consistent with other aquatic-adapted tetrapods including: a dorsally directed external naris, tall neural spines and elongate chevrons in an elongated tail, a short and broad scapula, webbed feet, long cervical vertebrae with long slender ribs, and an elongated rostrum with long and pointed teeth [3]. Together these features represent one of the best-supported cases of a semi-aquatic mode of life for a stem archosaur [4]. Together with Vancleavea campi, the discovery of L. somnii demonstrates a growing body of evidence that there was much more diversity in mode of life outside Archosauria. Furthermore, L. somnii helps interpret other possible character states consistent with a semi-aquatic mode of life for archosauriforms, including archosaurs.” [5]

Notes

1. Why restrict the taxon search to the clade Archosauria (crocs + dinos) a priori when Litorosuchus looks nothing like any of them? Better to add the new taxon to a large gamut analysis and let the scores determine the clade nesting, especially knowing that an antorbital fenestra, if present, is convergent over several unrelated clades.
2. The results only ‘suggest’ [we call this a weasel-word because you can weasel your way out of it] because Litorosuchus bears no resemblance to its ‘by default‘ sister taxa. In the LRT the nesting is sure and secure, supported by a long list of traits used as evidence and readily apparent just by looking (Fig. 3).
3. These are the basic traits of the Macrocnemus clade (Fig. 3). Someone evidently had their blinders on.
4. Stem archosaurs (sensu LRT, crocs + dinos only) were bipedal and terrestrial.  Basal archosauriformes (sensu LRT, like Proterosuchus) were indeed semi-aquatic. Macrocnemids also had a semi-aquatic lifestyle. Perhaps they competed.
5. Jimi Hendrix said it best, “Castles made of sand slip into the sea, eventually.” The only question is, how long with the Nesbitt, Stocker excluded taxon cladogram take to find more solid footing with a larger gamut matrix?

I only had to take a look at the foot of Litorosuchus
to know that the Li team had missed a golden opportunity to discuss macrocnemid affinities. The long pointed skull with the long premaxillary ascending process together with the long neck and short limbs all link Litorosuchus to macrocnemids. A sternum is present, which is typically absent in archosauriforms other than birds. The length and depth of the tail are so far unique within this clade, as are the presence or preservation of the dermal  armor. The antorbital fenestra has its genesis in the macrocnemids with a larger appearance in fenestrasaurs like Cosesaurus (Fig. 6) and its descendants. Litorosuchus is not directly related to Cosesaurus, but both are derived from a sister to the BES SC specimen (Fig. 5) which has a small precursor antorbital fenestra.

If we delete all lepidosauromorpha
from the LRT, then Litorosuchus nests not with Vancleavea, but as the sister to Pamelaria and the WMsN specimen assigned to Protorosaurus.

Diandongosuchus is discussed within the Li et al. text
because it is semi-aquatic, coeval and an archosauriform. However it is unrelated to archosaurs except through Youngina UC1528 and basalmost Proterosuchus specimens. Diandongosuchus is therefore unrelated to Litorosuchus in the LRT except through basalmost reptiles. And yes, Nesbitt and Stocker claimed credit for discovering that Diandongosuchus (Fig. 4) was a basal phytosaur when you heard that here first, four years ago. Either this is proof that they don’t Google or proof that they like to stick to outmoded paradigms while ignoring the greater evidence.

Soft tissue
Litorosuchus clearly had a long series of dorsal plumes similar to those discovered on the small BES SC111 specimen assigned to Macrocnemus (Fig. 5) and its sisters, Cosesaurus, Sharovipteryx (Fig. 6), Longisquama (Fig. 8) and Bergamodactylus (Fig. 9) the basalmost pterosaur. Dr. Pierre Ellenberger discussed the plumes of Cosesaurus, but falsely considered them proto-feathers. Critics could see these plumes, too, if they would take the time to do so.

Earlier we looked at the origin of dorsal plumes all in a row, a trait that goes back to the basal tritosaur, Huehuecuetzpalli.

Let’s see how the nodding journalists
handle Litorosuchus. It’s not on Wikipedia yet.

References
Li C, Wu X-C, Zhao L-J, Nesbitt SJ, Stocker MR, Wang L-T 2016. A new armored archosauriform (Diapsida: Archosauromorpha) from the marine Middle Triassic of China, with implications for the diverse life styles of archosauriforms prior to the diversification of Archosauria. The Science of Nature 103: 95. doi:10.1007/s00114-016-1418-4
Nesbitt SJ 2011. The early evolution of archosaurians: relationships and the origin of major clades. Bull Amer Mus Nat Hist 352:1–292.
Nesbitt SJ, Stocker MR, Small BJ and Downs A 2009. The osteology and relationships of Vancleavea campi (Reptilia: Archosauriformes). Zoological Journal of the Linnean Society 157 (4): 814–864. doi:10.1111/j.1096-3642.2009.00530.x.