Laquintasaura: verrrry basal ceratopsian from the Early Jurassic

Figure 2. Phytodinosauria with a focus on Stegosauria (yellow green).

Figure 1. Subset of the LRT focusing on the Phytodinosauria. Here Laqunitasaura nests at the base of the Ceratopsia.

I still hold to the hypothesis|
that a phylogenetic analysis that is able to lump and separate taxa is better than one that cannot do this. In the large reptile tree (LRT, 989 taxa), Laquintasaura venezuelae (Barrett et al. 2014; Early Jurassic, 200mya ~1m in overall length; Fig. 2) nests at the base of the ceratopsia (outside of Hexinlusaurus and Yinlong) and not far from the base of the Ornithopoda (outside of Changchunsaurus). It is very plesiomorphic and very early even for an ornithischian, let alone a ceratopsian.

Figure 1. Laquintasaura and tooth from Barrett et al. 2014. The early and plesiomorphic ornithischian has a naris shifted dorsally and other traits that nest it between the base of the onithopoda (Changchunsaurus) and the base of the ceratopidae (Hexinlusaurus).

Figure 2. Laquintasaura and tooth from Barrett et al. 2014. The early and plesiomorphic ornithischian has a naris shifted dorsally and other traits that nest it between the base of the onithopoda (Changchunsaurus) and the base of the ceratopidae (Hexinlusaurus). Compare to premaxillary teeth in figure 3.

Barrett et al. were not so sure where Laquintasaura nested
as they reported, “A strict consensus of these 2160 MPTs places Laquintasaura in an unresolved polytomy with the major ornithischian clades Heterodontosauridae, Neornithischia and Thyreophora along with other early ornithischian taxa, such as Lesothosaurus.”

The Barrett et al. diagnosis reports:
“Laquintasaura can be differentiated from other early ornithischians by the following autapomorphic combination  of dental characters: cheek tooth crowns have isosceles-shaped outlines, which are apicobasally elongate, taper apically, are mesiodistally widest immediately apical to the root/crown junction, possess coarse marginal denticles extending for the full lengths of the crown margins, and possess prominent apicobasally extending striations on their labial and lingual surfaces. Postcranial autapomorphies include: sharply inflected dorsal margin of ischium dorsal to the obturator process; femoral fibula epicondyle medially inset in posterior or ventral views; and astragalus with a deep, broad, ‘U’-shaped notch in anterior surface.”

I had no access to the fossil(s).
And I had to trust the drawing produced by Barrett et al. (Fig. 1) for my data. Contra the Barrett et all. analysis, there was no loss of resolution with Laquintasaura in the LRT.

Figure 2. The skull of Yinlong a basal certatopsian.

Figure 3 The skull of Yinlong a basal certatopsian. Those premaxillary teeth are quite similar to those figure in Barrett et al. for Laquintasaura. Note the dorsal naris, horizontal ventral premaxilla.

References
Barrett PM, Butler RJ, Mundil R, Scheyer TM, Irmis RB, Sánchez-Villagra MR. 2014. A palaeoequatorial ornithischian and new constraints on early dinosaur diversification. Proceedings of the Royal Society B 281:20141147. http://dx.doi.org/10.1098/rspb.2014.1147

One of the largest Pterodaustro specimens had stomach stones

aka: Gastroliths.
And that’s unique for pterosaurs of all sorts. So, what’s the story here?

Figure 1. The V263 specimen compared to other Pterodaustro specimens to scale.

Figure 1. The MIC V263 specimen compared to other Pterodaustro specimens to scale. Its one of the largest and therefore, most elderly.

One of the largest Pterodaustro specimens
MIC V263 (Figs. 1-5), was reported (Codorniú, Chiappe and Cid 2013) to have stomach stones (gastroliths). That made news because that represented the first time gastroliths have been observed in 300 Pterodaustro specimens and thousands of pterosaurs of all sorts.

Unfortunately,
Codorniu, Chiappe and Cid followed tradition when they aligned pterosaurs with archosaurs, like dinos (including birds) and crocs. Those taxa also employ gastroliths for grinding devices. According to Codorniú, Chiappe and Cid, other uses include as a personal mineral supply, maintaining a microbial flora, elimination of parasites and hunger appeasement. Shelled crustaceans may have formed a large part of the Pterodauastro diet and stones could have come in handy on crushing their ‘shells’ according to the authors.

FIgure 2. Pterodaustro specimen MIC V263 in situ and as originally traced.

FIgure 2. Pterodaustro specimen MIC V263 in situ and as originally traced.

The authors also noticed
an odd thickening of the anterior dentary teeth and the relatively large size of the MIC V 263 specimen (Fig. 1) and suggested their use as devices for acquiring stones.

The wingspan of this big Pterodaustro is estimated at 3.6 meters.

Figure 1. Pterodaustro elements from specimen MIC V263.

Figure 3. Pterodaustro elements from specimen MIC V263.

Unfortunately,
the authors overlooked a wingtip ungual (Fig. 4), or so it seems… The confirming wingtip ungula is off the matrix block. But they weren’t looking for it…

Figure 2. One wing ungual was preserved in this specimen of Pterodaustro. The other is missing off the edge of the matrix.

Figure 4. One wing ungual was preserved in this specimen of Pterodaustro. The other is missing off the edge of the matrix.

The authors overlooked a distal phalanges on the lateral toe (Fig. 5). It is hard to see. And they were not looking for it. Note the double pulley joint between p2.1 and p2.2. That’s where the big bend comes in basal pterosaurs.

Figure 5. Pterodaustro MIC V263 pes in situ and with pedal digit 2 reconstructed from overlooked bones.

Figure 5. Pterodaustro MIC V263 pes in situ and with pedal digit 2 reconstructed from overlooked bones.

The authors overlooked a manual digit 5, the vestigial near the carpus (Fig. 6) displaced to the disarticulated carpus during taphonomy. Again, easy to overlook. And they were not looking for it…

Figure 6. Carpus of the Pterodaustro specimen MIC V263 withe elements colorized. Manual digit 5 elements are in blue on the pink ulnare.

Figure 6. Carpus of the Pterodaustro specimen MIC V263 withe elements colorized. Manual digit 5 elements are in blue on the pink ulnare. Not sure where carpal 5 is.

The authors
labeled the unguals correctly (Fig. 7), but some of the phalanges escaped them. Note the manual unguals are not highly curved, like those of Dimorphodon and Jeholopterus. And for good reason. Pterodaustro is a quadrupedal beachcomber with the smallest fingers of all pterosaurs. It’s not a tree clinger. And for the same reason, pterosaurs with long curved manual claws are not quadrupeds. Paleontologists traditionally attempt to say all pterosaurs are quadrupeds, rather than taking each genus or clade individually. Beachcombers made most of the quadrupedal tracks. It’s also interesting to note that Pterodaustro fingers bend sideways at the knuckle, in the plane of the palm, probably in addition to flexing toward the palm. It’s easier for lizards to do this, btw. Not archosaurs. That’s how you get pterosaur manual tracks with digit 3 oriented posteriorly, different from all other tetrapods.

Figure 7. Pterodaustro MIC V 263 fingers reconstructed and restored.

Figure 7. Pterodaustro MIC V 263 fingers reconstructed and restored. Pterodaustro is unusual in having metacarpals 1 > 2 > 3. Note the flat tipped manual unguals. Not good for climbing trees, like those of many other pterosaurs.

So the question is: why did this specimen have stones inside—
when other pterosaurs do not? Since MIC V263 is larger, it is probably older, closer to death by old age. Was it supplementing an internal grinding structure that had begun to fail? Was this some sort of self-medication for a stomach ailment? It’s not standard operating procedure for pterosaurs to have stomach stones. So alternate explanations will have to do for now. Let’s not assume or pretend that all pterosaurs had gastroliths. They don’t.

Figure 8. Elements of the MIC V263 specimen applied to the smaller PPVL 3860 specimen scaled to the length of the metacarpals. At this scale the large Pterodaustro had a shorter wing and shorter fingers with smaller unguals.

Figure 8. Elements of the MIC V263 specimen applied to the smaller PVL 3860 specimen scaled to the length of the metacarpals. At this scale the large Pterodaustro had a shorter wing and shorter fingers with smaller unguals.

Compared to the largely complete and articulated Pterodaustro specimen,
PVL 3860, there are subtle differences in proportion (Fig 8) to the larger MIC V263 specimen. If metacarpals are the same length, then the wing is shorter in the larger specimen. This follows a morphological pattern in which no two tested pterosaurs are identical. Still looking for a pair of twins.

References
Codorniú L, Chiappe LM and Cid FD 2013. First occurrence of stomach stones in pterosaurs. Journal of Vertebrate Paleontology 33:647-654.

Zhongjianosaurus: a tiny dromaeosaurid? No.

Wikipedia reports,
“Zhongjianosaurus yang (Xu and Qin 2017, Yixian Fm. ~60 cm in ln length; ) is a genus of dromaeosaurid belonging to the Microraptoria.”

Unfortunately
the large reptile tree (LRT) nested Zhongjianosaurus with the scansoriopterygid bird, Mei long (Fig. 1). Neither does Microraptor nest with dromaeosaurids. It nests closer to Ornitholestes. Increasing the taxon list will resolve this issue for other workers as it did here.

Figure 1. Zhongjianosaurus compared to Mei long, a scansoriopterygid bird.

Figure 1. Zhongjianosaurus compared to Mei long, a scansoriopterygid bird. Both have relatively short forelimbs vs. long hind limbs among other traits.

Xu and Qin report,
The distal carpal is represented by the compound ‘semilunate’ carpal, formed by the addition of distal carpal 4 on its ventrolateral corner, and this morphology also is present in the troodontid Mei long (Xu et al., 2014a).”

Well, 
Mei long is indeed a troodontid, but so are all birds. Better to label it a scansoriopterygid bird to avoid confusion.

When you read the PDF, bear in mind
that the authors do not label the manual digits 1–3, but 2–4 as they pay homage to Limusaurus with what I call digit 0.

Perhaps if the pelvis or skull was preserved
in Zhongjianosaurus it would nest elsewhere. At present shifting Zhongjianosaurus to Microraptor adds 6 steps. Shifting Zhongjianosaurus to Velociraptor adds 9 steps. With the given data set and character list, this is how it all shakes out at present. And, you have to admit, it’s a pretty good match!

References
Xu X; Qin Z-C 2017. A new tiny dromaeosaurid dinosaur from the Lower Cretaceous Jehol Group of western Liaoning and niche differentiation among the Jehol dromaeosaurids” (PDF). Vertebrata PalAsiatica. In press.

wiki/Zhongjianosaurus

The forgotten clade: the REAL proximal ancestors to Dinosauria

Ignored by Baron et al. 2017, and everybody else
the Junggarsuchus clade (including Pseudhesperosuchus, Carnufex and Trialestes in order of increasing quadrupedality, Figs. 1–4) nests as the proximal ancestors to Herrerasaurus (Fig. 1) and the rest of the Dinosauria (Fig. 5) in the large reptile tree (LRT). That cladogram tests a wider gamut of taxa in greater detail than any other reptile cladogram ever published, attempting to not overlook anything. The Junggarsuchia is a sister clade to the Crocodylomorpha with both arising from a taxon near Lewisuchus (Fig. 1). Traditional paleontology (see Wikipedia) nests this largely ignored clade with the sphenosuchian crocodylomorphs (Fig. 4)… and for two good reasons!

Figure 1. Members of the Junggarsuchus clade were derived from a sister to the basal crocodylomorph, Lewisuchus and produced one line that includes Pseudhesperosuchus and Trialestes. The other line produced dinosaurs. These taxa are shown to scale. Note the evolution from a bipedal configuration to a quadrupedal stance.

Figure 1. Members of the Junggarsuchus clade were derived from a sister to the basal crocodylomorph, Lewisuchus and produced one line that includes Pseudhesperosuchus and Trialestes. The other line produced dinosaurs. These taxa are shown to scale. Note the evolution from a bipedal configuration to a quadrupedal stance.

One: Paleontologists never seem to include Dinosauria
in their smaller gamut croc analyses because they’re looking at crocs!~. So once again, taxon exclusion is holding some workers back from seeing ‘the big picture’. ReptileEvolution.com and the blog you are currently reading is all about examining ‘the big picture.’

Figure 2. Skulls of the Junggarsuchus clade not to scale. Herrerasaurus is the basalmost dinosaur.

Figure 2. Skulls of the Junggarsuchus clade not to scale. Herrerasaurus is the basalmost dinosaur, closely related to Junggarsuchus.

Two: Junggarsuchians ALSO have elongate proximal wrist bones
Elongate proximal carpals are found in both sphenosuchian crocs and derived members of the Junggarsuchus clade. Paleontolgists wrongly assumed such odd wrist bones were homologous. It’s an easy mistake to make. However, the LRT makes clear that intervening taxa, including Junggarsuchus, do not have elongate wrist bones.

Among taxa that preserve the manus,
(Fig. 3) it is Junggarsuchus that nests closest to Herrerasaurus and the Dinosauria.

Figure 3. Hands of Lewisuchus, Herrerasaurus, Junggarsuchus, Pseudhesperosuchus and Trialestes. The proximal carpals (radiale and ulnare) were elongate by convergence with a line of crocodylomorphs. This has confused paleontologists and mentally removed them from possible ancestry to the Dinosauria. Note the very short proximal carpals in Junggarsuchus.

Figure 3. Hands of Lewisuchus, Herrerasaurus, Junggarsuchus, Pseudhesperosuchus and Trialestes. The proximal carpals (radiale and ulnare) were elongate by convergence with a line of crocodylomorphs. This has confused paleontologists and mentally removed them from possible ancestry to the Dinosauria. Note the very short proximal carpals in Junggarsuchus.

Like the basal members of the Crocodylomorpha
the Junggarsuchus clade (the Prodinosauria here) transition from bipedal basal members to quadrupedal derived members, with the longest forelimbs belonging to the most derived member, Trialestes (Fig. 3). Distinct from the others and contra the original interpretation, I think Trialestes may have had a larger ulnare than radiale, to match its larger ulna.

Figure 4. Crocodylomorph manus and carpus samples including Terrestrisuchus, Erpetosuchus, Hesperosuchus and Dibothrosuchus along with Scleromochlus documenting the elongate radiale and ulnare on derived taxa. Ticinosuchus is the closest example of an ancestral/plesiomorphic manus in the LRT.

Figure 4. Crocodylomorph manus and carpus samples including Terrestrisuchus, Erpetosuchus, Hesperosuchus and Dibothrosuchus along with Scleromochlus documenting the elongate radiale and ulnare on derived taxa. Ticinosuchus is the closest example of an ancestral/plesiomorphic manus in the LRT.

Let’s not forget
PVL 4597 (Fig. 6) which was mistakenly considered a specimen of Gracilisuchus by (Lecuona and Desojo 2011), but under phylogenetic analysis in the LRT, still nests as the proximal outgroup to Herrerasaurus. It is tiny specimen, supporting the hypothesis of phylogenetic miniaturization at clade origin. And it retains a small proximally oriented calcaneal tuber, as found in other Junggarsuchians.

Figure 1. Subset of the LRT focusing on the Archosauria (Crocodylomorpha + Dinosauria and kin). Gray areas document specimens with elongate proximal carpals (radiale and ulnare).

Figure 5. Subset of the LRT focusing on the Archosauria (Crocodylomorpha + Dinosauria and kin). Gray areas document specimens with elongate proximal carpals (radiale and ulnare).

We looked at
phylogenetic miniaturization at the origin of several pterosaur clades. Well, it happens here too, at the base of the Dinosauria (Fig. 1) with PVL 4597 (Fig. 6), easily overlooked, easily mistaken for something else.

One should not ‘choose’ outgroup taxa
based on paradigm, tradition, guessing, convenience or opinion. Rather outgroup taxa should ‘choose themselves’ based on rigorous testing of a large gamut of outgroup candidates in phylogenetic analysis. To minimize selection bias, the LRT provides 858 outgroup taxa the opportunity to nest close to dinosaurs.

Figure 6. The closest known taxa to the Dinosauria, PVL 4597, is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

Figure 6. The closest known taxa to the Dinosauria, PVL 4597, is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

 

References
Baron MG, Norman DB, Barrett PM 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543:501–506.
Bonaparte JF 1969. 
Dos nuevos “faunas” de reptiles triásicos de Argentina. Gondwana Stratigraphy. Paris: UNESCO. pp. 283–306.
Butler RJ. et al. 2014. New clade of enigmatic early archosaurs yields insights into early pseudosuchian phylogeny and the biogeography of the archosaur radiation. BMC Evol. Biol. 14, 128.
Clark JM et al. 2000. A new specimen of Hesperosuchus agilis from the Upper Triassic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology 20 (4): 683–704.
doi:10.1671/0272-4634(2000)020[0683:ANSOHA]2.0.CO;2.
Clark JM, Xu X, Forster CA and Wang Y 2004. A Middle Jurassic ‘sphenosuchian’ from China and the origin of the crocodilian skull. Nature 430:1021-1024.
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 2011. The early evolution of archosaurs: relationship and the origin ofmajor clades. Bull. Amer. Mus. Nat. Hist. 352, 1–292.
Novas FE 1994. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto
Reig OA 1963. La presencia de dinosaurios saurisquios en los “Estratos de Ischigualasto” (Mesotriásico Superior) de las provincias de San Juan y La Rioja (República Argentina). Ameghiniana 3: 3-20.
Sereno PC and Novas FE 1993. The skull and neck of the basal theropod Herrerasaurusischigualastensis. Journal of Vertebrate Paleontology 13: 451-476. doi:10.1080/02724634.1994.10011525.
Zanno LE, Drymala S, Nesbitt SJ and Schneider VP 2015. Early Crocodylomorph increases top tier predator diversity during rise of dinosaurs. Scientific Reports 5:9276 DOI: 10.1038/srep09276.

wiki/Pseudhesperosuchus
wiki/Junggarsuchus
wiki/Carnufex
wiki/Herrerasaurus
wiki/Sanjuansaurus

 

What traits separate phytodinosaurs from theropods?

Yesterday we looked at the origin of dinosaurs in the context of and contra the recent Baron et al. 2017 paper. Today we’ll look at the basal split between basal phytodinosaurs, like Eodromaeus (Figs. 1, 2), with the closely related basal theropods, like Tawa (Fig. 1).

Figure 1. The theropod Tawa compared to the closely related phytodinosaur, Eodromaeus.

Figure 1. The theropod Tawa compared to the closely related smaller phytodinosaur, Eodromaeus.

Placed side-by-side to scale
Tawa and Eodromaeus are similar overall, though the plant-eaters were initially smaller. The details (below) demonstrate the initial steps toward herbivory that characterize the Phytodinosauria, distinct from the Theropoda and basal Dinosauria from which they evolved (contra Baron et al. 2017).

Figure 1. Eodromaeus reconstructed. We will look at this taxon in more detail tomorrow.

Figure 1. Eodromaeus reconstructed. We will look at this taxon in more detail tomorrow.

How do basal phytodinosaurs differ from the basal theropods?
Here’s the LRT list:

  1. lateral rostral shape: convex and smoothly curved (also in ancestral Herrerasaurus and Gracilisuchus);
  2. premaxilla/maxilla angle 25–45º;
  3. naris shape in lateral view almost round (not longer than tall or taller than long);
  4. postfrontal has no contact with the upper temporal fenestra;
  5. opisthotic oriented laterally without posttemporal fenestrae;
  6. palatal teeth (only on basalmost taxa);
  7. maxillary tooth depth ≤ 2x width in lateral view;
  8. last maxillary tooth at mid orbit (also in Herrerasaurus);
  9. olecranon process present (convergent in Buriolestes);
  10. metacarpals 2 and 3 align with m1.1 (except Eodromaeus);
  11. acetabulum laterally oriented (no ventral deflection, as in basal theropods);
  12. femoral head with neck and offset (appears later in theropods);
  13. penultimate manual phalanges not the longest in each series;
  14. loss of pubic boot (likely plesiomorphic because outgroups to Herrerasaurus do not have a pubic boot).

Summary and significance
Compared to the closely related theropod Tawa, the overall similar phytodinosaur Eodromaeus had a taller rounder rostrum, shorter teeth, a higher coronoid process, a longer dorsal region with more robust dorsal vertebrae, reduced gastralia, a more robust pectoral girdle and forelimb with shorter, less raptorial fingers, a deeper pubis and ischium with more robust hind limbs. The shorter teeth and larger belly together with the more robust limbs and back are traits seen in a wide variety of herbivores, even if only transitional at this early stage.

References
Baron MG, Norman DB, Barrett PM 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543:501–506.

Basal dinosaurs: what makes them dinosaurs?

Earlier we looked at Baron et al. 2017, who heretically allied ornithischians with theropods to create a new clade ‘Ornithoscelida.’ After a week of study and several tests, I came away with the realization that the Baron et al. tree topology did not depend on taxon exclusion, based on comparable results from subsets of the large reptile tree (LRT). Reducing the list of LRT taxa to match the Baron et al. taxon list did not change the dinosaur tree topology. Rather scoring inaccuracies were the problem, and that goes back to the original matrix presented by Nesbitt 2011, which Baron et al. continued using. You might remember, that’s the study that nests pterosaurs with parasuchians with no intervening taxa, among many other odd nestings. We’ll detail those inaccuracies later this week.

One of the reasons
Baron et al. allied theropods with ornithisichians is because they nested two phytodinosaurs, Eodromaeus (Fig. 1) and Eoraptor, as basalmost theropods. So when they thought they were comparing the ‘theropod’ Eoraptor with the ornithischian, Heterodontosaurus (their figure 2), they were actually comparing two phytodinosaurs. So, of course they shared a long list of characters. To really test their assertion, Baron et al. should have tested Heterodontosaurus against a real basal theropod, like Tawa (Fig. 2). (Tomorrow we’ll compare Tawa with Eodromaeus.)

Figure 1. Eodromaeus reconstructed. We will look at this taxon in more detail tomorrow.

Figure 1. Eodromaeus reconstructed. We will look at this taxon in more detail tomorrow.

Along the way, 
I was able to better understand what was going on at the  base of the Dinosauria and at the basal Theropod/Phytodinosaur split. For instance, as noted earlier, Herrerasaurus (Fig. 3) has a smaller number of cervicals than other dinos do, matching the number found in ancestral basal crocodylomorpha, like Lewisuchus, not silesaurids, like Silesaurus.

Figure 2. Tawa, a basal theropod. Note how closely it resembles the phytodinosaur, Eodromaeus. That's because they are closely related as basalmost dinosaurs.

Figure 2. Tawa, a basal theropod. Note how closely it resembles the phytodinosaur, Eodromaeus. That’s because they are closely related as basalmost dinosaurs.

How do basal dinosaurs differ from the basal croc ancestors?
Here’s the latest list:

  1. skull shorter than cervical series (not Herrerasaurus):
  2. skull width less than 1.2x height;
  3. posterolateral premaxilla narrower than naris (not Herrerasaurus);
  4. quadrate curls posterodorsally (not Herrerasaurus);
  5. dentary tip straight or descends (does not rise);
  6. deep canine maxillary teeth absent (not Herrerasaurus);
  7. nine or more cervicals (not Herrerasaurus);
  8. cervical ribs slender, parallel centra (Herrerasaurus unknown);
  9. 3–4 sacral ribs (not Herrerasaurus, which has 2);
  10. mid-caudal centra 3x longer than tall where known (not Herrerasaurus);
  11. clavicles poorly ossified or absent (reappear in later taxa);
  12. interclavicle poorly ossified or absent;
  13. manual unguals long with penultimate phalanges longer than proximal phalanges;
  14. tibia not shorter than femur (not Herrerasaurus);
  15. fourth trochanter sharp (also in the proximal dinosaur outgroup taxon PVL 4597);
  16. tibia not < 2x ilium length (not Herrerasaurus);
  17. advanced and simple metatarsal joint without a calcaneal tuber (also in Lewisuchus, convergent in Silesaurus);
  18. compact metatarsus;
  19. longest metatarsal: #3;
  20. metatarsal #1 < 75% of metatarsal #3 length;
  21. metatarsals 2–4 ≥ tibia (not Herrerasaurus);
  22. proximal metatarsals 1 and/or 5 reduced in diameter.

Summary and significance
Arising from the morphology of basal bipedal crocodylomorphs, many of these basal dinosaur traits document a longer, leaner bipedal morphology with an increasingly robust fulcrum (pelvis and sacrum), along with longer distal hind limb elements with more reliance on the middle digits for faster locomotion. By contrast the proximal outgroup clade, Junggarsuchus and kin, documents a return to quadrupedalism terminating with Trialestes and its long forelimbs. The increasingly elongate radiale and ulnare of the Junggarsuchus clade turn out to be convergent, rather than homologous, with those of other bipedal crocodylomorphs, like Terrestrisuchus and Hesperosuchus, a clade that also transitioned toward a quadrupedal configuration, as seen in extant crocodylomorphs. We’ll take a closer look at this clade and this transition later this week.

Figure 1. Lewisuchus cervical/dorsal transition at top photo and the same for Herrerasaurus drawings, including a foreword shift of the pectoral girdle in a 2-frame GIF movie. The cervical ribs are imagined.

Figure 1. Lewisuchus cervical/dorsal transition at top photo and the same for Herrerasaurus drawings, including a foreword shift of the pectoral girdle in a 2-frame GIF movie. The cervical ribs are imagined.

References
Baron MG, Norman DB, Barrett PM 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543:501–506.

 

Tulerpeton pes options

Earlier the pedal elements of the amphibian-like reptile Tulerpeton were moved around to produce a reasonable reconstruction. Today I offer a few more options (Fig. 1) including one with six toes. All appear to be reasonable.

Figure 1. Tulerpeton pes reconstruction options using published images of the in situ fossil.

Figure 1. Tulerpeton pes reconstruction options using published images of the in situ fossil.

None of these reconstructions
changes the nesting of Tulerpeton as the basalmost Reptile (=Amniote). Such long toes with so many phalanges in these patterns of relative length are not found in basal tetrapods.

Figure 1. Tulerpeton parts from Lebedev and Coates 1995 here colorized and newly reconstructed. Manus and pes enlarged in figure 2.

Figure 2 Tulerpeton parts from Lebedev and Coates 1995 here colorized and newly reconstructed. Manus and pes enlarged in figure 2.

A little backstory
Tulerpeton curtum (Lebedev 1984, Fammenian, Latest Devonian, 365 mya) was described as, “one of the first true tetrapods to have arisen.” Here it nests as the basalmost reptile, pushing Gephyostegus bohemicus back to the pre-amniotes. Very little other than the limbs are known. In life it would have been similar to and the size of Gephyrostegus, Urumqia and EldeceeonTulerpeton lived in shallow marine waters.

References
Coates MI and Ruta M 2001 2002. Fins to limbs: What the fossils say. Evolution & Development 4(5): 390–401.
Lebedev OA 1984. The first find of a Devonian tetrapod in USSR. Doklady Akad. Navk. SSSR. 278: 1407–1413.
Lebedev OA and Clack JA 1993. Upper Devonian tetrapods from Andreyeva, Tula Region, Russia. Paleontology36: 721-734.
Lebedev OA and Coates MI 1995. postcranial skeleton of the Devonian tetrapod Tulerpeton curtum Lebedev. Zoological Journal of the Linnean Society. 114 (3): 307–348.

wiki/Tulerpeton