Baron 2017: 21 ‘unambiguous’ theropod/ornithischian synapomorphies don’t pan out

Yesterday we looked at Baron et al. 2017, who proposed uniting Ornithischia with Theropoda to the exclusion of Sauropodomorpha + Herrerasaurus and kin (Fig. 1), among several other relationships not recovered by the large reptile tree (LRT, 980 taxa). They did so by excluding dinosaur outgroup taxa recovered by the LRT, like Gracilisuchus and Pseudhesperosuchus, while including inappropriate outgroup taxa, like pterosaurs, Lagerpeton and kin, and poposaurs, like Silesaurus. In paleontology this is known as ‘cherry-picking’ and yesterday’s post showed how cherry-picking outgroup taxa, like the pterosaur Dimorphodon, can lead to having scansoriopterygid basal birds recovered as basal dinosaurs. Baron et al. did this by focusing on, and mis-scoring minute traits, not readily visible from an arm’s length of viewing. See below.

By contrast,
the LRT provides a very long list of candidate outgroup taxa going back to Devonian tetrapods and lets the computer decide the topology of the reptile family tree including the Dinosauria. It thereby minimizes a priori bias and subjective or traditional opinion in taxon selection. The LRT also employs more readily observable traits and few to no minutia. The LRT is fully resolved with high Bootstap scores, in contrast to the Baron et al. trees.

Today we’ll dive deeper into Baron et al. 2017
They start with a false premise by supporting the clade ‘Ornithodira‘, which is a junior synonym for Reptilia, since it includes pterosaurs. In the LRT pterosaurs share a last common ancestor with dinosaurs in the Devonian amniote Tulerpeton, the last (and only) known common ancestor of all reptiles.

Baron et al. report, “A formal hypothesis proposing dinosaur monophyly was proposed in 1974, and consolidated in the 1980s. As a direct result of these and other analyses, Ornithischia and Saurischia came to be regarded as monophyletic sister-taxa: this hypothesis of relationships has been universally accepted ever since.” Not in the LRT, which recovered evidence in 2011 to support a clade Phytodinosauria, uniting Sauropodomorpha with Ornithischia + several basal phytodinosaur genera.

Baron et al. report, “No studies on early dinosaur relationships have included an adequate sample of early ornithischians and the majority of studies have also excluded pivotal taxa from other major dinosaur and dinosauromorph (near dinosaur) lineages.” The LRT did so include more than an adequate sample of all pertinent taxa.

Baron et al. report, “In order to examine the possible effects of these biases on our understanding of dinosaur evolution, we carried out a phylogenetic analysis of basal Dinosauria and Dinosauromorpha and compiled, to our knowledge, the largest and most comprehensive dataset of these taxa to date.” No, the LRT is larger and more comprehensive. It is under the authority of the LRT that mistakes can be revealed in the Baron et al. study.

Baron et al. report,Although this study has drawn upon numerous previous studies, no prior assumptions were made about correlated patterns of character evolution or dinosaur interrelationships.” Not true. Their exclusion of appropriate and inclusion of inappropriate taxa demonstrates their assumptions. By this statement they appear to have fooled themselves as well, based on the taxon list of the the LRT.

Baron et al. report, “We analysed a wide range of dinosaurs and dinosauromorphs, including representatives of all known dinosauromorph clades.” Not true. They did not include dinosaur outgroup taxa recovered by the LRT (Fig. 2).

Figure 1. According to Baron et al. 2017 these taxa are related in this fashion.

Figure 1. According to Baron et al. 2017 these taxa are related in this fashion. The LRT does not recover these relationships.

Here is the ‘meat’ of todays post:
Baron et al. report, “The formation of the clade Ornithoscelida [Ornithischia + Theropoda] is strongly supported by 21 unambiguous synapomorphies including: [comments follow]

  1. an anterior premaxillary foramen located on the inside of the narial fossa [present in basal sauropodomorphs Leyesaurus and Pampadromaeus.]
  2. a sharp longitudinal ridge on the lateral surface of the maxilla [present in basal sauropodomorph Pantydraco.]
  3. a jugal that is excluded from the margin of the antorbital fenestra by the lacrimal–maxilla bone contact (this appears convergently in some ‘massospondylids’) [not excluded in Tawa or Coelophysis.]
  4. an anteroventrally oriented quadrate [seemingly all dinosaurs have this sort of quadrate orientation]
  5. short and deep (length of more than twice the dorsoventral height) par occipital processes [apparently a mistake because the figure 2 caption text lists, “elongate par occipital processes.”]
  6. a post-temporal foramen that is entirely enclosed within the par occipital process [I cannot check this minutia with available data]
  7. a supraoccipital that is taller than it is wide [I cannot check this minutia with available data]
  8. a well-developed ventral recess on the parabasisphenoid [I cannot check this minutia with available data]
  9. a surangular foramen positioned posterolaterally on the surangular [I cannot check this minutia with available data]
  10. an entirely posteriorly oriented retroarticularprocess, which lacks any substantial distal upturn [present in basal sauropodomorph Pantydraco.]
  11. at least one dorsosacral vertebra anterior to the primordial pair [I cannot check this with available data]
  12. neural spines of proximal caudals that occupy less than half the length of the neural arches (which are also present in some sauropodomorphs, but absent in Herrerasauridae, Guaibasaurus, and nearly all sauropodomorphs as or more derived than Plateosaurus [it doesn’t matter about derived taxa, we’re looking only at basal taxa, this is a variable trait not present on Scuttelosaurus, but present on Efraasia]
  13.  scapula blade more than three times the distal width (also found in Guaibasaurus) [also found in Herrerasaurus and Sajjuansaurus]
  14. humeral shaft that has an extensively expanded ventral portion of the proximal end, creating a distinct bowing (convergently acquired in plateosaurids and more derived sauropodomorphs) [sounds like a deltopectoral crest, If so, this is universal among Dinosauria]
  15. absence of a medioventral acetabular flange (which was also lost in plateosaurids and more derived sauropodomorphs) [unable to check this minutia with available data]
  16. a straight femur, without a sigmoidal profile (which was also acquired by more derived sauropodomorphs, but absent in basal forms such as Saturnalia and Pampadromaeus, and is also absent in Herrerasauridae) [also absent in Eoraptor, present in Pantydraco]
  17. a well-developed anterior trochanter that is broad and at least partly separated from the shaft of the femur [absent in Eodromaeus and otherwise difficult to check with available data]
  18. a strongly reduced fibular facet on the astragalus [unable to check this minutia with available data]
  19. a transversely compressed calcaneum with reduced posterior projection and medial process [unable to check this minutia with available data]
  20. a first metatarsal that does not reach the ankle joint, but that is instead attached ventrally to the shaft of metatarsal II [not in Tawa, Scelidosaurus or Haya]
  21. fusion of the distal tarsals to the proximal ends of the metatarsals.[not in Tawa, Scelidosaurus or Haya]

several of these ‘traits’ are minutia. The LRT uses larger traits that one can see and measure from a greater viewing distance or with published figures.

According to Baron et al.
other shared features uniting Ornithischia with Theropoda included: [comments again follow]

  1. a diastema between the premaxillary and maxillary tooth rows of at least one tooth crown’s length [not in Eodromaeus, Emausaurus]
  2. an extended contact between the quadratojugal and the squamosal bones [not in a wide variety of ornithischians]
  3. an anterior tympanic recess (convergently acquired in Plateosaurus) [unable to check this minutia with available data]
  4. a fibular crest on the lateral side of the proximal portion of the tibia (described as present in Eoraptor, although we could not confirm its presence, which is also absent in Tawa [unable to check this minutia with available data]
  5. an oblique articular end of the tibia in which the outer malleolus extends further distally than the inner malleolus (although this appears to be absent in Pisanosaurus [unable to check this minutia with available data]
  6. fusion of the sacral neural spines [unable to check this minutia with available data, often hidden by the pelves]
  7. presence of an antitrochanter on the ilium [unable to check this minutia with available data]
  8. reduction of the distal end of the fibula [not in Buriolestes, Tawa, Scelidosaurus]
  9. fusion of the tibia, fibula and proximal tarsals into a tibiotarsus [not in BuriolestesTawaScelidosaurus]
  10. fusion of the metatarsals [not in BuriolestesTawaScelidosaurus]

Apparently Baron et al. were not
thorough enough in these assessments and again depended for the most part, on minute traits rather than large, readily observable ones, Apparently referees were likewise not thorough enough on their vetting of this manuscript. I imagine because it is difficult to do when all the data is not gathered into a single readily reference resource, like The present vetting took only a few hours.

According to Baron et al. 
“20 additional steps would be needed to recover Saurischia as previously defined.” But that’s a false goal according to the LRT results that do not recover a clade Saurischia. And with such bad scoring (see above) this goal turns out to be a misstep, not a step.

Baron et al. report,
“in our hypothesis a fully carnivorous feeding strategy is not recovered as the plesiomorphic condition for Dinosauria and we are forced to interpret some of the anatomical similarities between herrerasaurids and theropods as convergences.” In the LRT, herrerasaurids are basal to all remaining dinosaurs, yet have certain autapomorphies that indicate an older, more plesiomorphic last common ancestor of all dinosaurs is awaiting discovery.

Baron et al. report, 
“Dinosauria is recovered in a polytomy with Silesauridae and the enigmatic Late Triassic British taxon Saltopus elginensis.” In the LRT, both of those outgroups are surrounded by other taxa that separate them from Dinosauria.

Figure 1. The origin of dinosaurs to scale. Gray arrows show the direction of evolution. This image includes Decuriasuchus, Turfanosuchus, Gracilisuchus, Lewisuchus, Pseudhesperosuchus, Herrerasaurus, Tawa and Eoraptor.

Figure 2. The origin of dinosaurs to scale according to the LRT.  Gray arrows show the direction of evolution. This image includes Decuriasuchus, Turfanosuchus, Gracilisuchus, Lewisuchus, Pseudhesperosuchus, Herrerasaurus, Tawa and Eoraptor.

Several years ago
the above (Fig. 2) was published online. It remains the best graphic portrayal of basal Dinosauria and their outgroups to date, based on a much larger number of outgroup taxa than has ever been published before. Unfortunately, the Baron et al. team did not take advantage of this readily available and thoroughly verified hypothesis.

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



New radical dinosaur cladogram: Baron, Norman and Barrett 2017

Baron, Norman and Barrett 2017
have just allied Ornithischia with Theropoda to the exclusion of Sauropodomorpha. That radical hypothesis was not recovered by the large reptile tree (LRT, 980 taxa) nor any other study in the long history of dinosaurs. Despite the large size of their study, it was not large enough. And so taxon exclusion bites another group of well-meaning paleontologists who used traditional small inclusion sets.

From the Baron et al. abstract:
“For 130 years, dinosaurs have been divided into two distinct clades—Ornithischia and Saurischia. Here we present a hypothesis for the phylogenetic relationships of the major dinosaurian groups that challenges the current consensus concerning early dinosaur evolution and highlights problematic aspects of current cladistic definitions. Our study has found a sister-group relationship between Ornithischia and Theropoda (united in the new clade Ornithoscelida), with Sauropodomorpha and Herrerasauridae (as the redefined Saurischia) forming its monophyletic outgroup. This new tree topology requires redefinition and rediagnosis of Dinosauria and the subsidiary dinosaurian clades. In addition, it forces re-evaluations of early dinosaur cladogenesis and character evolution, suggests that hypercarnivory was acquired independently in herrerasaurids and theropods, and offers an explanation for many of the anatomical features previously regarded as notable convergences between theropods and early ornithischians.”

As a reminder, the fully resolved cladogram
at finds Herrerasaurus as a basal dinosaur arising from the Pseudhesperosuchus clade. Tawa (Fig. 1) and Buriolestes lead the way toward Theropoda. Barberenasuchus and Eodromaeus are basal to Phytodinosauria, which includes Sauropodomorpha + Ornithischia. So the Nature piece is totally different due to taxon exclusion and improper taxon inclusion.

Earlier heretical dinosaur origins were presented here with images and complete resolution with high Bootstrap scores at every or virtually every node.

Problems with the Baron et al. report

  1. Lack of resolution: Over dozens of nodes, only 5 bootstrap scores were over 50 (the minimum score that PAUP shows as fully resolved).
  2. Lack of correct proximal outgroup taxa (taxon exclusion) and they chose several wrong outgroup taxa (see below) because they had no large gamut analysis that established the correct outgroup taxon out of a larger gamut of choices
  3. Lack of several basal dinosaur taxa. (again, taxon exclusion, see below)
  4. Improper taxon inclusion: poposaurs, pterosaurs and lagerpetons are not related to dinos or their closest kin
  5. Lacking reconstructions for all pertinent basal/transitinal taxa so we can see their data at a glance, see if a gradual accumulation of traits can be observed and not have to slog through all the scores
Figure 1. Unrelated archosaurs. Silesaurus is a poposaur. Eoraptor is a phytodinosaur (note the big belly). And Tawa is a lean theropod.

Figure 1. Unrelated archosaurs mentioned in this blog. Silesaurus is a poposaur. Eoraptor is a phytodinosaur (note the big belly). And Tawa is a lean theropod.

LRT differences with the Baron et. al results

  1. Carnivorous Staurikosaurus, Herrerasaurus, Chindesaurus and Sanjuansaurus nest at the base of the herbivorous Sauropodomorpha.
  2. Herbivorous Eoraptor nests at the base of the Theropod with Tawa.
  3. Poorly known Saltopus sometimes nests as the last common ancestor of Dinosauria.
  4. Six taxa nest basal to dinosaurs in SupFig1 including the poposaur Silesaurus and kin. Silesaurus has ornithischian and theropod traits and so appears to make an ideal outgroup taxon,  but nests with neither clade when more taxa are included. This is the key problem with the study: pertinent taxon exclusion. 
  5. The lack of Gracilisuchus and other bipedal basal crocs that nest basal to dinos in the LRT certainly skewed results.

In an effort to understand Baron et al. I duplicated their outgroup taxon list
but retained all the LRT dinosaurs to see what would happen. The SupFigs are available free online at

  1. SupFig 1: When Euparkeria is the outgroup and Postosuchus is included: 3 trees result and (theropods Herrerasaurus + Tawa + Buriolestes) + (poposaurs Sacisaurus + Silesaurus) nest as the base of the Phytodinosauria, while bipedal croc Saltopus nests at the base of the Theropoda.
  2. SupFig 2: When the lepidosaur pterosaur Dimorphodon is the outgroup and Euparkeria + Postosuchus are excluded: 12 trees and basal scansoriopterygid birds (come to think of it, they DO look like Dimorphodon!) nest as basal dinosaurs, then the bird cladogram gets reversed such that basal becomes derived, but Phytodinosauria is retained.
  3. SupFig. 3: when Silesaurus is the outgroup: 12 trees and Phytodinosauria is retained in the LRT
  4. SupFig. 4: when no characters were treated as ordered. Neither does the  LRT order any characters, so this test was moot.

Dr. Kevin Padian said, 
“‘original and provocative reassessment of dinosaur origins and relationships”. And because Baron and his colleagues used well-accepted methods, he notes, the results can’t simply be dismissed as a different opinion or as mere speculation. “This will send people back to the drawing board,” he added in an interview.”

“There have been a lot of studies on the phylogenetic relationships, the family tree of the dinosaurs, but they’ve mostly been on individual dinosaurian groups. They haven’t really examined the entire dinosaur tree in such depth. And so this analysis had the advantage of using a different and larger set of critters than most previous trees. They’ve analyzed the characters used by others before and then also adding their own characteristics and getting their selves quite different configurations, radically different in fact.

The LRT has had, for several years, an even larger set of taxa, so large that any bias in selecting an outgroup taxon list has been minimized. Unfortunately, Baron et al. were biased and used traditional outgroup taxa that skewed their results.

Dr. Hans-DieterSues reported,
“For one thing, palaeontologists’ analyses of relations among species are keenly sensitive to which species are considered, as well as which and how many anatomical features are included, he says.”

Many more outgroup taxa would have minimized the inherent bias clearly present in Baron et al. When Silesaurus is your outgroup, herbivores will nest with carnivores. When you start your study with a goal in mind (read and listen to Baron’s comments) that’s never good. When you exclude taxa that have been shown to be pertinent to your study, that’s never good.

That’s what is here for (on the worldwide web). Free. Testable. And with a demonstrable gradual accumulation of traits along with minimal bias due to its large gamut.

I was surprised to see Nature print this
because they have not published relationship hypotheses in favor of  new specimens of note. Co-author Dr. David Norman has published for several decades and has a great reputation.

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

You heard it here first: Lagerpeton is NOT a dinosauromorph

Thanks to Dr. Max Langer
for sending this abstract.

Five years after
Lagerpeton was removed from the Dinosauromorpha online here, Novas and Agnolin 2016 follow suit by nesting Lagerpeton with Tropidosuchus, among the Proterochampsidae.

This is confirmation of methodology
and the value of a large gamut analysis in reducing a priori bias and the influence of paradigm. You don’t have to see the fossil firsthand to make a contribution to paleontology. The Science is in the testing of heretical hypotheses. The Pseudoscience is in the critical blackwashing of results without testing.

Not sure how the critics will take this news.
I know they hate it every time testing reveals something that is not in their paradigm.

From the Novas and Agnolin 2016? abstract
“The relationships of basal ornithodirans [1] constitute a hotly debated topic of vertebrate evolution. Among one of the most important basal ornithodirans is the enigmatic Lagerpeton chanarensis, coming from the Middle Triassic Los Chañares Formation, Talampaya National Park, La Rioja Province, Argentina. With the aim to evaluate the phylogenetic position of this taxon, a detailed comparison with other Triassic archosauriforms was conducted. Surprinsingly, extensive similarities between Lagerpeton and proterochampsids, particularly Tropidosuchus, were found. Because of that, an integrative phylogenetic analysis including Lagerpeton and several archosauriform clades was constructed. The analysis resulted in the recognition of Lagerpeton as a basal archosaurifom nested within Proterochampsidae, and nearly related to the genus

“Lagerpeton resembles proterochampsids in several derived features, including 1) proximal pubis with robust ambiens process; 2) lateral margin of pubis sigmoid in anterior view; 3) acetabulum cup-like and ellipsoidal shaped; 4) transverse processes on caudal vertebrae long and narrow; 5) femoral 4th trochanter proximodistally expanded; 6) caudal surface of distal tibia with a middle tubercle surrounded by two shallow concavities; 7) astragalus with anteromedial corner acute; and 8) metatarsal II tranversely thick. Furthermore, Lagerpeton and Tropidosuchus share elongate and compact metatarsus with metatarsal V reduced and devoid of phalanges, and with articular surface for distal tarsal 4 subparallel to the longitudinal axis of shaft, and metatarsal IV longer than III. In spite of the similarities noted above, Lagerpeton differs from proterochampsids and resembles dinosauriforms in having a well-defined and proximally convex femoral head, and absence of osteoderms.

“Previous authoritative papers by J. Bonaparte and P. Sereno and A. Arcucci have illustrated the hindlimbs of Lagerpeton as vertically positioned. However, manual articulation of the femoral head within the deep and elliptical-shaped acetabulum avoids a parasagittal position of the entire hindlimb. On the contrary, the most unforced position of the hindlimbs in Lagerpeton depicts its femora in a sprawling posture. In this position, distal femoral condyles orientate almost ventrally to articulate with tibia and fibula. In lateral view, the articulated hindlimb is oblique, being subhorizontally oriented. In these respects, Lagerpeton resembles basal archosauriforms, including proterochampsids, but differs from the parasagittal posture of ornithodirans.

“The exclusion of Lagerpeton from the dinosaur lineage results in the removal of the clade Dinosauromorpha, [2, 3] which was originally conceived to encompass Lagerpeton plus Dinosauriformes. The recognition of Lagerpeton as a derived proterochampsian widens the morphological radiation that this clade of basal archosauriforms manifested during mid-Triassic times.”


  1. Ornithodira is still an invalid clade encompassing pterosaurs and dinosaurs and their kin, which, in the LRT, encompasses all Reptilia.
  2. We may want to resurrect a clade name for the archosaur outgroups to the Dinosauria listed here and discussed and illustrated here and here.
  3. Dinosauromorpha was also removed here five years ago.

Novas FE and Agnolin FL 2016 Lagerpeton chanarensis Romer (Archosauriformes): A derived proterochampsian from the middle Triassic of NW Argentina. Simposio. From Eventos del Mesozoico temprano en la evolución de los dinosaurios”. Programa VCLAPV. Conferencia plenaria: Hidrodinámica y modo de vida de los primeros vertebrados. Héctor Botella (Universitat de València, España) 2016

Early evolution of archosaurs – SVP abstract 2016

Nesbitt et al. 2016 discuss the early evolution of bird-line archosaurs.

From the Nesbitt et al. abstract:
“Bird-line archosaurs (= Avemetatarsalia, the clade containing dinosaurs, pterosaurs, and their kin) (1) had their origin in the Triassic Period. However, that origin is poorly documented (2) as fossils from their early evolutionary history are extremely rare and  consist mostly of postcrania. (3) Here, we report the discovery of a new reptile (femoral length = 17 cm) (4) from the lower portion of the Middle Triassic Lifua Member (Manda beds) of the Ruhuhu Basin, southwestern Tanzania. Material referred to the new taxon includes a partial skeleton of a single individual including cervical, trunk, and caudal vertebrae, pectoral, pelvic, forelimb, and hind limb material (= ‘Teleocrater’ of A. Charig), and parts (skull elements, vertebrae, pectoral, pelvic, and limb elements) of a minimum of three individuals collected from a bonebed discovered in 2015 very close to Charig’s original partial skeleton. Character states of the limbs, vertebrae, and ilium indicate a close relationship with early dinosauromorphs including: elongated cervical vertebrae, an ilium with a slightly concave ischial peduncle and clear anterior crest, a weakly developed anterior trochanter of the femur, an anteriorly compressed fibula with long strap-like iliofibularis crest, and absence of osteoderms (5). Many character states suggest that the new reptile taxon falls outside of the pterosaur-dinosaur clade (= Ornithodira). (6) However, the distributions of some of these character states at the base of Archosauria are unclear and some character states of the new taxon suggest a more basal relationship outside Archosauria (e.g., absence of two medial tubera of the proximal femur). No matter the position within or outside Archosauria, the new Lifua taxon shares seemingly unique character states with the poorly known Dongusuchus from the Middle Triassic of Russia (known from femora) and Yarasuchus from the Middle Triassic of India (known from partial skeletons), rather than with other archosauriforms. As a result, these forms appear to represent a globally distributed clade of early diverging avemetatarsalians. (7) The larger body size of the Manda form and its potential phylogenetic position outside of pterosaurs and dinosauromorphs indicates that there was a size decrease at the origin of Ornithodira. (8) This new taxon, and other new discoveries from the Middle to Late Triassic, are elucidating the sequence of character acquisitions in Avemetatarsalia and fill a crucial gap in the evolutionary history that led to the flourishing of dinosaurs later in the Mesozoic”. (9)


  1. Avemetatarsalia is a parphyletic clade or a junior synonym of the clade Reptilia because pterosaurs do not nest with dinosaurs and archosaurs in the LRT. Evidently taxon exclusion bias keeps Nesbitt et al. from considering fenestrasaur, tritosaur lepidosaurs as pterosaur ancestors. Only crocs and dinos comprise the Archosauria in the LRT. Like Dr. Naish, this clade of paleontologists are not exploring all the possibilities offered by a large gamut analysis, but opting to stay with traditional untested taxon lists.
  2. In the LRT the origins of all reptilian clades are well-documented going all the way back to basal tetrapods.
  3. Bogus claim, probably based on the mistaken assumption that headless Lagerpeton is basal to dinos. The origin of dinos is documented here.
  4. Similar in size to Lewisuchus, a pro-dinosaur.
  5. Some ‘dinosauromorph traits’ are in contention based on the inclusion set and recovered taxa basal to dinos.
  6. The ‘Ornithodira‘ is an invalid clade, or at the least, a junior synonym for ‘Reptilia’ in the LRT.
  7. Yarasuchus nests with a variety of genera in a clade between Rauisuchidae and Archosauria in the LRT.
  8. Indeed there was a size decrease at the origin of pterosaurs and at the origin of dinosaurs, but they had separate origins.
  9. If related to Yarasuchus, the new taxon is indeed distantly related to the origin of archosaurs, which included Gracilisuchus and the PVL specimen attributed to Gracilisuchus, both derived from sisters to the much larger Decuriasuchus and Turfanosuchus. The purported ‘gap’ reported by Nesbitt et al. appears to be because they are looking at the wrong outgroup taxa due to taxon exclusion in their phylogenetic analysis.
Nesbitt SJ, Butler RJ, Barrett PM, Stocker MR, Sidor CA, Angielczyk KD, Ezcurra MD and Smith RM 2016. The early evolution of bird-line archosaurs: a possible new clade of globally distributed avemetatarsalians just outside the dinosaur-pterosaur split. Abstract from the 2016 meeting of the Society of Vertebrate Paleontology.

Trialestes rises again!!

Lecuona et al. 2016
redescribe in greater detail Trialestes (Reig 1963; Figs. 1, 2), more than 50 years after its original discovery and publication. Glad to see this! Data used to nest Trialestes in the LRT as the proximal outgroup for the Dinosauria consisted of a few old drawings (Fig. 1), nothing more. The new data do not move the nesting much.

Figure 1. Tracings from old drawings are the data used to create this reconstruction of Trialestes, which nested it basal to the Dinosauria.

Figure 1. Tracings from old drawings are the data used to create this reconstruction of Trialestes, which nested it basal to the Dinosauria. New data from Lecuona et al. 2016 greatly reduce the guesswork here. 

From the Lecuona et al. abstract:
“Here, we describe in detail all the material assignable to the species and test its phylogenetic relationships using a comprehensive data matrix focused on early archosaurs. We support the referral of PVL 3889 to Trialestes and reject the presence of a mesotarsal ankle joint in this specimen. We recovered Trialestes within Crocodylomorpha, closer to Crocodyliformes than Pseudhesperosuchus, Hesperosuchus, Dromicosuchus and Sphenosuchus. Therefore, Trialestes represents the most completely known of the earliest non-crocodyliform crocodylomorph taxa known to date.”

They report, “In contrast, Reig’s third taxon, ‘Triassolestes’ romeri has received relatively little attention. In his original publication, Reig (1963) interpreted this taxon as a theropod dinosaur of the family Podokesauridae, a group now considered roughly equivalent to Coelophysoidea (Holtz 1994). Because of this interpretation, Reig considered only part of PVL 2561 as the holotype of the dinosaur ‘Triassolestes’, including an incomplete cranium and mandible, four cervical vertebrae, and 16 caudal vertebrae. Other postcranial remains associated with PVL 2561 were excluded from this genus, including a scapula, humerus, ulna, radius, carpus and proximal metacarpus, which were interpreted by Reig (1963, p. 15) as crocodilian elements because of the presence of an elongated radiale and ulnare.”

FIgure 2. Assembly of the many Trialestes parts featured in Lecuona et al. 2016.  This is an odd combination of robust cervicals and gracile limbs and girdles.

FIgure 2. Assembly of the many Trialestes parts featured in Lecuona et al. 2016. This is an odd combination of robust cervicals and gracile limbs and girdles.

Those proximal wrist elements
(radiale and ulnare) are never elongated in dinosaurs, but they are elongated in dinosaur ancestors like Trialestes. The name ‘Triassolestes’ was preoccupied by an Australian Triassic dragonfly, hence the change we know today.

Lecuona et al. continue: “Clark (in Benton & Clark 1988, fig. 8.6) hypothesized Trialestes as the sister taxon of Crocodylomorpha, but expressed some caution given the structure of the ankle and the poor knowledge of the specimens.”

“Novas (1989) recognized that the referred specimen PVL 2559 contained elements belonging to two individuals of different sizes (Reig 1963, fig. 4b part.) and assigned both of them to Herrerasauridae indet. (Novas 1989, 1993); presently, they can only be assigned to an indeterminate saurischian dinosaur.”

“Clark et al. (2000) questioned the referral of PVL 3889 to Trialestes romeri, suggesting that this specimen was more likely assignable to a basal dinosaur. Nevertheless, these authors could not reject the alternative explanation that PVL 2561 and PVL 3889 belong to one taxon with a combination of crocodylomorph and dinosaurian character states.”

“In addition, Trialestes has not been included in a quantitative phylogenetic analysis so far and, thus its affinities remain untested using modern methodologies.”

Well, the large reptile tree did that several years ago. But let’s keep an open mind moving forward.

Under materials and methods, Lecuona et al report,
“Herein we study the two specimens assigned to Trialestes romeri, the holotype PVL 2561 and PVL 3889; we exclude PVL 2559 because we agree with previous authors that it represents an indeterminate saurischian.”

Lecuona et al. based their analysis on a cladogram originated by Nesbitt (2011) which had several problems listed here. They also included data from Butler et al. (2014) and added Carnufex, a related taxon.

The Trialestes fossils have a pristine beauty. The authors did not create a reconstruction, so I attempt one here (Fig. 2).

A little background data
Trialestes romeri (Bonaparte 1982)= Triassolestes (Reig, 1963/Tillyard, 1918) Carnian, Late Triassic ~235 mya is known from scattered parts. Clark, Sues and Berman (2000) redescribed the known parts and admitted the possibility that this taxon combined dinosaurian and crocodylomorph characters. As it nests here, Trialestes was derived from a sister to Carnufex. This clade phylogenetically preceded Herrerasaurus and the Dinosauria. We looked at this heretical relationship several years ago here.

Distinct from Pseudhesperosuchus,
the skull of Trialestes had a larger antorbital fenestra and a deeper rostrum. The mandibular fenestrae (yes, there are two!) were smaller.

The vertebral centra had excavated lateral surfaces, for bird-like air sacs. The radius was longer than the humerus, a character otherwise known only in dinosaurs. The radiale was smaller than the ulnare, matching the radius and ulna. The fingers were tiny.

The pelvis was semi-perforated, as in basal dinosaurs, with a well-developed supraacetabular crest. The dorsal pelvis was straight, as in Gracilisuchus. The femoral head was not inturned, suggesting a variable posture, promoted by that really long forearm. The ankle joint had a crocodile normal configuration and a functionally pentadactyl pes. Most crocs lose pedal digit 5, but not those basal to dinos, like PVL4597.

Bonaparte JF 1982. Classification of the Thecodontia. Geobios Mem. Spec. 6, 99-112
Clark JM, Sues H-D and Berman DS 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.
deFranca MAG, Bittencourt JdS and Langer MC 2013. Reavaliação taxonomica de Barberenasuchus brasiliensis (Archosauriformes), Ladiniado do Rio Grande do Sul (Zona-Assembleia de Dinodontosaurus). Palaenotogia em Destaque Edição Especial Octubro 2013: 230.|
Irmis RB, Nesbitt SJ and Sues H-D 2013. Early Crocodylomorpha. Pp. 275–302 in Nesbitt, Desojo and Irmis (eds). Anatomy, phylogeny and palaeobiology of early archosaurs and their kin. The Geological Society of London. doi:10.1144/SP379.24.
Kischlat EE 2000. Tecodôncios: a aurora dos arcossáurios no Triássico. Pp. 273–316 in Holz and De Ros (eds.). Paleontologia do Rio Grande do Sul. Porto Alegre: CIGO/UFRGS.
Lecuona A, Ezcurra MD and Irmis RB 2016. Revision of the early crocodylomorph Trialestes romeri (Archosauria, Suchia) from the lower Upper Triassic Ischigualasto Formation of Argentina: one of the oldest-known crocodylomorphs. Papers in Palaeontology (advance online publication). DOI: 10.1002/spp2.1056
Nesbitt S 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352: 292 pp.

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.

Riff D et al. 2012. Crocodilomorfos: a maior diversidade de répteis fósseis do Brasil. TERRÆ 9: 12-40, 2012.

‘Origin of Dinosaurs’ video. 


The Origin of Dinosaurs x2 (2010) revisited

Several years ago
the top vertebrate paleontologists in the world (Brusatte et al. 2010) reported on the origin of dinosaurs. Coincidentally Langer et al. (2010) wrote a similar report.  It is now 6 years later. Let’s see how well those report have held up as they compare to the current data (2016) in the large reptile tree.

From the Brusatte et al. introduction
“During the past 25 years, numerous new fossils, reinterpretations of long-forgotten specimens, and numerical analyses have significantly revised our understanding of this major macroevolutionary event, which is one of the most profound and important evolutionary radiations in the history of life.”

What has stood the test of time:

  1. Dinosaurs are archosaurs: birds+crocs and last common ancestor
  2. Archosaurs are diapsid reptiles = Eudibamis, Petrolacosaurus and all their descendants.
  3. Dinosaurs are: “Triceratops horridus, Passer domesticus, and all descendants of their most recent common ancestor.” Or alternatively: ““the least inclusive clade containing Megalosaurus and Iguanodon.” Thus dinosaurs are monophyletic.
  4. The suite of traits common to dinosaurs include: 1) upright and fully erect posture [shared with basal crocs and dinosauromorphs]; 2) an enlarged deltopectoral crest on the humerus [shared with Trialestes]; 3) a “specialized” hand; 4) a perforated acetabulum (hip socket) [which may close]; 5) a well-developed fourth trochanter on the femur; 6) a lesser trochanter on the femur; 7) and a simple hinge ankle joint with proximal tarsals fixed immovably to the tibia and fibula [shared with basal crocs and dinosauromorphs].
  5. Dinosaurs likely originated during the Middle Triassic. They are diverse at the earliest Late Triassic.
  6. Herrerasaurus and Eoraptor are some of the most complete specimens of any early dinosaur.
  7. Langer: Herrerasaurs are basal to the Ornithischia-Saurischia dichotomy, but the actual dichotomy is Theropoda/Phytodinosauria
  8. Langer: The oldest dinosaurs include Herrerasaurus, Eoraptor, Staurikosaurus, Saturnalia and Panphagia all from the Carnian (early Late Triassic). These are also among the most primitive dinosaurs. Missing from this list is Barberenasuchus, also Carnian, not commonly considered a dinosaur, but nests as a sister to Eodromaeus.

What has not stood the test of time:

  1. Archosaurs (crocs + dinos alone) no longer include pterosaurs
  2. Diapsids no longer include lizards, snakes, rhynchocephalians (including rhynchosaurs and trilophosaurs) and pterosaurs. Those have a diapsid skull by convergence.
  3. Arizonasaurus is no longer an archosaur since crocs and birds had a more recent common ancestor, a sister to Gracilisuchus.
  4. The clades Crurotarsi (= Pseudosuchia) and Avemetatarsalia (= Ornithodira, Ornithosuchia) are now junior synonyms for older nomenclature based on their inclusion sets (Archosauriformes and Reptilia respectively).
  5. Pterosaurs no longer nest with archosaurs, but with lepidosaurs, in a new clade known as the Tritosauria nesting between basal rhynchocephalians and basal protosquamates.
  6. Lagerpeton is not a dinosauromorph, but a sister to Tropidosuchus.
  7. Marausuchus is does not nest outside the Dinosauria, but as a basal theropod.
  8. Sacisaurus, Silesaurus and Asilisaurus are not the immediate sisters of dinosaurs. Rather they now nest with poposaurs, the proximal outgroup to the Archosauria (crocs + dinos only).
  9. Overlooked by Brusatte et al., Lewisuchus, Zupaysaurus, Pseudhesperosuchus, Trialestes, and their kin are the now the immediate sisters of dinosaur, the true dinosauromorphs.
  10. Some manner of feathers now diagnose the Dinosauria, which primitively had naked (not scaly) skin, like a plucked chicken.
  11. Herrerasaurus and Eoraptor are no longer incerta sedis but the most basal dinosaur and one of the basal phytodinosaurs respectively.
  12. Zupaysaurus no longer nests as a theropod, but a dinosauromorph
  13. Berberosaurus no longer nests as a theropod, but as the basalmost phytodinosaur
  14. Ornithischia no longer branch off first from Saurischia, but are derived from basal phytodinosaurs. Sauropodomorpha are sisters to basal Ornithisichia with Daemonosaurus and Chilesaurus at the base.
  15. Langer: Eusaurischia (Sauropodomorpha + Theropoda) is a junior synonym for Dinosauria
  16. Langer: Silsauridae (all taxa closer to Silesaurus than to Marasuchus + Heterodontosaurus) is a junior synonym for Poposauria, if kept monophyletic.
  17. Langer: the basal-most dinosaurs were not probably omnivorous,
  18. Langer: herrerasaurs were not theropods
  19. Langer: there is no Onithischia-Saurischia dichotomy. Saurischia is a junior synonym  for Dinosauria.
  20. Langer: Agnophitys is a dinosaur sister to Marasuchus.
  21. Langer: Putative dinosaur Saltopus is a basal archosaur close to Gracilisuchus.
Figure 1. Click to enlarge. Subset of the large reptile tree focusing on the Archosauria (crocs + dinos). Sharp-eyed observers will find minor changes here.

Figure 1. Click to enlarge. Subset of the large reptile tree focusing on the Archosauria (crocs + dinos). Sharp-eyed observers will find minor changes here.

Langer et al. (2010) mentioned Staurikosaurus (Colbert 1970) as the first consensual early dinosaur to be collected. Here it nests as a basal theropod, basal to a clade of theropods that is often overlooked that includes Marasuchus, Procompsognathus and Segisaurus. Yes, Staurikosaurus has but two sacral vertebrae. So do other clade members.

Langer et al. (2010) also mentioned Guaibasaurus (Bonaparte et al., 1999) who reported, “The mesotarsal condition and the outline of the distal section of tibia indicate the saurischian nature of this new form, but the almost unreduced medial wall of the acetabular portion of ilium shows an unrecorded primitive condition within the cited group. Several features suggesting affinities with both the Prosauropoda and Theropoda, imply that Guaibasaurus candelariensis may belong to the ancestral group for both of them.” The large reptile tree nests Guaibasaurus as a basal theropod and as the sister to Marasuchus + Procompsognathus, not far from Staurikosaurus. 

The Novas (1992) dinosaur definition
According to Langer et al., Novas (1992b) provided the first phylogenetic definition of Dinosauria as ‘‘the common ancestor of Herrerasauridae and Saurischia + Ornithischia, and all of its descendants’’. The addition of herrerasaurs does not change the current tree (Fig. 1). Padian & May (1993) explicitly restricted the use of Dinosauria to the clade composed of Saurischia and Ornithischia, exclusive of ‘‘Herrerasaurus and its allies’’. But Novas has priority. Moreover, the last common ancestor of Saurischia and Ornithischia is currently a herrerasaur. The diagnosis of the Dinosauria has seen some changes over the years. Many are traits that are not covered by the large reptile tree. Please check out the references below for lists and histories of those lists.

What does the large reptile tree diagnose dinosaurs?
The following suite of traits are found in basal dinosaurs and not their proximal outgroups, Trialestes, the Pseudhesperosuchus clade. However many of these traits are found elsewhere on the tree. And many traits are lost in more derived dinos.

  1. Naris opening lateral
  2. Parietal skull table weakly constructed
  3. Mandible tip straight (neither upturned nor down)
  4. Interclavicle poorly ossified or absent
  5. Coracoid shape disc-like, even if fused (elongate or strap shape in outgroup)
  6. Radiale and ulnare not elongated (as in outgroup)
  7. Manus with long penultimate phalanxes and raptorial claws
  8. Femoral head interned and sub rectangular (reversed in the Marasuchus clade).
  9. Longest metatarsal: 3
  10. Proximal metatarsals: 1 and 5 reduced

has long been touted as a key dinosaurian trait, but dinosaurs evolved from basal bipedal crocodylomorphs, like Gracilisuchus and Scleromochlus. Interesting that Scleromochlus has been often associated with unrelated pterosaurs. Pterosaur removal sets things a little straighter in the retelling of the dinosaur ancestry story. Scleromochlus has not often been touted as a dinosaur ancestor, but by virtue of its false association with pterosaurs in various cladograms, it has always been there.

The long coracoids and proximal carpals of basal bipedal crocs
have set them apart from consideration as possible dino ancestors. But if you just let the software do its job, then you’ll recover nestings that indicate the elongate coracoids and proximal carpals became reduced to shorter, more primitive conditions in basal dinos.

Traits found in dinosaurs exclusive of Herrerasaurus:

  1. Feathers (not on the matrix, but worth mentioning)
  2. Skull shorter than cervicals
  3. Cranium convex
  4. Naris opening
  5. Maxilla ventral margin straight
  6. Jugal qj process straight
  7. Quadrate curls posterodorsally
  8. Jaw joint aligned with ventral maxilla
  9. Canine maxillary teeth not present
  10. Nine or more cervical vertebrae
  11. Some caudal vertebrae 3x longer than tall
  12. Tibia not shorter than femur
  13. Metatarsus not shorter than half the tibia
  14. Phalanges on metatarsal 5: 0 (reversed in higher clades)

Then if wanted to
you could simply list all the traits of Herrerasaurus, the basalmost dinosaur, knowing full well that Herrerasaurus itself is derived from the first, as yet undiscovered, dinosaurs.

Bonaparte JF, Ferigolo J and Ribeiro M 1999. A new early Late Triassic saurischian dinosaur from Rio Grande do Sol state, Brazil” (PDF). Proceedings of the Second Gondwanan Dinosaur Symposium, National Science Museum Monographs 15: 89–109.
Brusatte SL, Nesbitt SJ, Irmis RB, Butler RG, Benton MJ and Norell MA 2010.
The origin and early radiation of dinosaurs. Earth-Science Reviews 101 (2010) 68–10.
Colbert EH 1970. A Saurischian dinosaur from the Triassic of Brazil. American Museum Novitates 2405; 1-39
Langer MC. Ezcurra MD, BittencourtJS, Novas FE 2010. The origin and early evolution of dinosaurs. Biological Review 85, 55–110.

Basal dinosaurs: what was their competitive advantage?

A Paul Sereno YouTube video (2011) on the discovery of Eodromaeus talks about the search for the competitive advantage that basal dinosaurs had that enabled them to eclipse their Triassic competitors and survive into the Jurassic and rise to prominence throughout the rest of the Mesozoic.

that search has not yet produced any distinct  and widely agreed upon answers. Dr. Sereno suggests it might have been, “a lucky break” for dinosaurs.

It’s not an upright/bipedal configuration
Basal crocs,  some poposaurssome rauisuchians, Lagerpeton + Tropidosuchus, and maybe Arizonasaurus were all bipedal, but they did not survive into the Jurassic. Pterosaurs were derived from Triassic bipeds, but that’s another story.

It might be feathers
Feathers used to define birds. Now they define dinosaurs with both small theropods and small phytodinosaurs having them. Big derived dinosaurs had scales, but in birds scales on the feet are former feathers. Phylogenetic bracketing indicates that certain types of feathers go back to basalmost dinosaurs, like Herrerasaurus and/or its ancestors among the protodinosaurs, like Lewisuchus and Trialestes. No evidence of soft tissue is known for these taxa.

Our only clue to basal croc widespread dermal protection
comes from a patch of tiny scales on the back of Scleromochlus. Scales like these don’t ultimately spread across the body in scaly crocs, like the extant Caiman. Osteoderms form along the spine in rauisuchians and are retained in certain basal crocs like Erpetosuchus, and Gracilisuchus. These do spread across the body in Simosuchus and extant crocs.

Evidently spinal osteoderms are not found
in protodinosaurs like Pseudhesperosuchus and kin. We don’t know if they were scaly or naked.

later derived armored dinosaurs, like Scelidosaurus and Scutellosaurus independently redevelop protective osteoderms.

Secondary sexual traits
don’t usually provide the keys to survival in terms of weather, viruses and other external factors. So feathers as secondary sexual traits probably do not give dinos the competitive advantage vs. non-conspecific competitors, like basal crocs, rauisuchians, etc.

Feathers as insulation
A dense coat of feathers insulates the naked skin of dinosaurs (including extant birds) from the elements, including air temperature, ultraviolet light, and damaging obstacles in the environment, including, at times, predators.

Figure 1. Kulindadromaeus, a sister to Heterodontosaurus with proto-feathers. Images from and traced from Godefroit et al. 2014. Since theropods and heterodontosaurs both had something like feathers, if they were the same kind of feathers, their last common ancestor had feathers. That last common ancestor was a herrerasaur or its proximal predecessor. Note the Godefroit et al. skull does not match their description but has a standard maxilla ascending process. See color overlays for correct ed interpretation.

Figure 1. Kulindadromeus, a sister to Heterodontosaurus with proto-feathers. Images from and traced from Godefroit et al. 2014. Since theropods and heterodontosaurs both had something like feathers, if they were the same kind of feathers, their last common ancestor had feathers. That last common ancestor was a herrerasaur or its proximal predecessor. Note the Godefroit et al. skull does not match their description but has a standard maxilla ascending process. See color examples for correct ed interpretation. Click to enlarge.

Pycnofibers as insulation
By convergence, that ‘other story,’ the fenestrasaurs (including pterosaurs) likewise had a coat of insulating fibers (pycnofibers). And pterosaurs, like dinosaurs, survived into the Jurassic and beyond.

Hair/Fur as insulation
By convergence, hairy/furry mammals survived into the Jurassic and beyond.

Does this tell us something?
It’s just a clue, but a 3x winner is a pretty good clue IMHO.

On the flip side
uninsulated lepidosaurs, turtles, crocs, choristoderes and enaliosaurs also survived into the Jurassic and beyond. Am I forgetting any other large clades? Some of these were aquatic. Others had a protective carapace. Still others could have found refuge in leaf litter.  Was there something in the air, like radiation, that killed uninsulated mid-sized terrestrial reptiles leaving smaller and aquatic taxa alone? Or was it just a ‘lucky break’? More on this topic follows:

On the same note, a recent paper 
by Benton, Forth and Langer (2014) notes that dinosaurs arose in the Middle Triassic (see PVL 4597, Herrerasaurus and Lewisuchus). They mention Nyasaurus and Asilisaurusbut those are poposaurs, not as close relatives to dinosaurs as are the crocs in the large reptile tree.

Benton, Forth and Langer report, “Tracking the forebears of crocodiles and birds back in time points to a common ancestor in the Early Triassic, and close relatives in the latest Permian, represented by Archosaurus from Russia.” That’s not true according to the large reptile tree which puts the last common ancestor near the Gracilsuchus/PVL 4597 split in the Middle Triassic.

More untested paradigms from Benton, Forth and Langer
who also report, “Within Archosauria, the bird line, Avemetatarsalia (Box 1), includes two subclades, Pterosauria (the flying reptiles) and Dinosauromorpha.”  Pterosaurs are lepidosaurs in the large reptile tree, which tests all these relationships.

Then they dig themselves deeper
“All avemetatarsalians have elongate hindlimbs (suggesting bipedal posture), elongate tibiae (suggesting adaptations to fast running), and three or four slender, elon- gate metatarsals in a tightly bound bundle, so these animals all stood high on their tip-toes (digitigrade posture).” So do basal crocs, which they seem to be forgetting.

“Dinosauromorphs had all these characters, as well as further elongation of the metatarsals and reduction of the fifth toe to a short single element.” So do basal crocs, which they seem to be forgetting.

“Dinosauriformes added to these specializations of the hindlimb some further modification of the pelvis and femur for speedy and efficient movement on two legs.” So do basal crocs, which they seem to be forgetting. But I think they are talking about Lagerpeton, which achieved bipedal configuration by convergence in the large reptile tree.

“Among these, the astragalus, the main ankle bone, sends a thin plate of bone up the front of the tibia, so linking the ankle firmly to the shin as a single functional unit. Many of these characters were once seen as exclusive to Dinosauria, but they are now known to exist in larger clades.” Not so fast… the astragalus sends up a thin plate of bone up the -back- of the tibia in Lagerpeton.

“Dinosaurs are characterized by some skull features, an elongate deltopectoral crest on the humerus (a major muscle attachment of the forearm), and an expanded articulation for the tibia on the astragalus.” None of these appear to be key to their survival. The large reptile tree finds another suite of traits for dinosaurs, none of which are exclusive and few of which are retained in all derived forms.

Getting back to competitive advantages
Benton, Forth and Langer (2014) note: “What were the characters that enabled dinosaurs, and indeed archosaurs more widely, to profit from these ecological crises? Two key attributes are their exceptional growth rates and efficient respiration systems. Recent work on dinosaurs shows… rates of growth in line with modern mammals rather than modern reptiles. Dinosaurs almost certainly possessed the unidirectional respiratory system of birds, and apparently crocodiles, which is more efficient than the tidal system in mammals, and this might have characterized all archosaurs.” In the large reptile tree “all archosaurs” includes only crocs and dinos, but high growth rates and efficient respiration systems are both competitive advantages both for quickly producing lots of little dinosaurs and running away from predators without tiring.

To their credit,
Benton, Forth and Langer (2014) note: “Dinosaurian thermoregulation is somewhat speculative, but the majority of evidence now supports a high metabolic rate, especially in the small- and medium-sized feathered dinosaurs. Considering that these include members of both the saurischian [60] and ornithischian [61] branches, phyloge- netic bracketing implies that the first dinosaurs might have had a high thermal inertia [62], given the insulation provided by the coverage of filamentous integumentary structures (Figure 2b), as well as fast growth and avian-like breathing.”

And so we come full circle,
in complete agreement. It’s not upright posture. It’s not great size. It’s not a bipedal configuration. It just might be feathery insulation (on top of upright posture, bipedal configuration and rapid growth) and everything else that comes with that suite of traits.

bigger, more derived, more robust dinos did not need feathers, and so those shrank to become dino scales, distinct from lepidosaur, turtle and croc scales.

Benton MJ, Forth J and Langer MC 2014. Models for the Rise of the Dinosaurs. Current Biology 24, R87–R95, online pdf