Kiwi ancestors

Worthy et al. 2013 reported:
“Until now, kiwi (Apteryx owenii, Apterygidae, Shaw 1813; Fig. 1) have had no pre-Quaternary fossil record to inform on the timing of their arrival in New Zealand or on their inter-ratite relationships.” They described two fossils (femur and quadrate) from the Early Miocene (Fig. 1; 19–16mya) which they named Proapteryx. “The new fossils indicate a markedly smaller and possibly volant bird, supporting a possible overwater dispersal origin to New Zealand of kiwi independent of moa. If the common ancestor of this early Miocene apterygid species and extant kiwi was similarly small and volant, then the phyletic dwarfing hypothesis to explain relatively small body size of kiwi compared with other ratites is incorrect.

Figure 2. Jurapteryx, Pseudocrypturus, Apteryx and Proapteryx to scale.

Figure 2. Jurapteryx, Pseudocrypturus, Apteryx and Proapteryx to scale. In lateral view it is difficult to see the width of the ventral pelvic elements. They are not as wide as the egg diameter. Note the lack of a pygostyle in all three taxa.

By contrast
the large reptile tree (LRT, 1213 taxa) nest the kiwi with Pseudocrypturus (Houde 1988; Early Eocene) apart from other ratites, as the basalmost birds with living representatives.

Apteryx owenii (Shaw 1813) The extant flightless kiwi has an elongate naris that extends to the tip of its beak. Maybe two teeth are there. Here it nests with Pseudocrypturus, but flightless traits push it toward Struthio, by convergence. in the pre-cladistic era, Calder (1978, 1984) considered the kiwi a phylogenetic dwarf derived from the larger moa, but that was invalidated by Worthy et al. 2013 and the large reptile tree.

Proapteryx micromeros (Worthy et. al. 2013) was a slender, tiny Miocene (18 mya) ancestor likely capable of flight.

Pseudocrypturus cercanaxius (Houde 1988; Early Eocene) was originally considered a northern hemisphere ancestor to ratites (like the ostrich, Struthio). Today these primitive flightless birds are chiefly restricted to the southern hemisphere. It could be that early birds did start in the South and had migrated to the North during the Paleocene (66-56 mya).

Since ratites are basal to extant birds, and Pseudocrypturus is basal to ratites (paleognaths), Pseudocrypturus is also quite similar to the ancestor of all extant birds despite its late appearance in the early Eocene. Perhaps something very much like it was one of the few survivors of the K-T extinction event.

It’s notable that Pseudocrypturus has long legs. Early ducks, like Presbyornis, and basal raptors, like Sagittarius, also had long legs. Evidence is building that this is the primitive condition for the clade of living birds arising from the K-T extinction event.

Worthy et al. nest Apteryx
within the order Casuariiformes, which includes cassowaries, emirs, and kiwi, but only in the absence of Pseudocrypturus.

The kiwi egg vs ventral pelvis issue
In most tetrapods, including humans, the egg/baby passes through the cloaca/vagina which passes through the two ischia. That was also likely the case with Archaeopteryx, because this is also the case with Gallus the chicken. In extant birds the ischia posterior tips no longer touch, but are widely separated. Going several steps further, in the kiwi the enormous egg is held in front of the pubis (Fig. 1), which is also in front of the ischia.

The following video of a kiwi laying an egg
shows the cloaca a substantial distance below the swirl that marks its tail. kiwi egg video click to play pretty much located at the tip of the long axis of the egg in figure 1 (maybe a little higher/closer to the tail).

Figure 2. Kiwi laying an egg. Click to play.

Figure 2. Kiwi laying an egg. Click to play.

In the LRT
Pseudocrypturus and Apteryx (Fig. 1) nest together and apart from the ratites. Pseudocrypturus is basal to all living birds. It probably first appeared in the Early Cretaceous. It was found in the Paleocene.

Calder WA 1978. The kiwi. Scientific American 239(1):132–142.
Calder WA 1984. Size, function and life history. 448 pp. Cambridge (Harvard U Press).
Houde PW 1986. Ostrich ancestors found in the northern hemisphere suggest new hypothesis of ratite origins. Nature 324:563–565.
Houde PW 1988. Paleognathus birds from the early Tertiary of the northern hemisphere. Publications of the Nuttall Ornithological Club 22. 147 pp.
Shaw 1813. Naturalist’s Miscellany 19:
Worthy, TH. et al. (5 coauthors) 2013. Miocene fossils show that kiwi (Apteryx, Apterygidae) are probably not phyletic dwarves. Paleornithological Research 2013, Proceedings of the 8th International Meeting of the Society of Avian Paleontology and Evolution. Retrieved 16 September 2017.

wiki/Apteryx, Kiwi


Quail hip joints are not good models for pterosaur hip joints

Manafzadeh and Padian 2018 tell us:
“Studies of soft tissue effects on joint mobility in extant animals can help to constrain hypotheses about joint mobility in extinct animals. However, joint mobility must be considered in three dimensions simultaneously, and applications of mobility data to extinct taxa require both a phylogenetically informed reconstruction of articular morphology and justifications for why specific structures’ effects on mobility are inferred to be similar. We manipulated cadaveric hip joints of common quail and recorded biplanar fluoroscopic videos to measure a ‘ligamentous’ range of motion (ROM), which was then compared to an ‘osteological’ ROM on a ROM map. Nearly 95% of the joint poses predicted to be possible at the hip based on osteological manipulation were rendered impossible by ligamentous constraints. Because the hip joint capsule reliably includes a ventral ligamentous thickening in extant diapsids,the hip abduction of extinct ornithodirans with an offset femoral head and thin articular cartilage was probably similarly constrained by ligaments as that of birds. Consequently, in the absence of extraordinary evidence to the contrary, our analysis casts doubt on the ‘batlike’ hip pose traditionally inferred for pterosaurs and basal maniraptorans, and underscores that reconstructions of joint mobility based on manipulations of bones alone can be misleading.”

Figure 6. Images of floating lizards. The small ones, like small pterosaurs, take advantage of surface tension to ride high while spread-eagle on the surface.


Manafzadeh and Padian 2018 are not phylogenetically informed.
They should have used lizards. Pterosaurs are not related to birds. Birds are archosaurs. Pterosaurs are lepidosaurs, which universally (except for legless taxa) assume a bat-like pose in their hind limbs when resting (Figs. 1, 2). Many articulated pterosaur fossils are found in the sprawling posture (Fig. 2) typically used for flying…but Manafzadeh and Padian are talking about quail hips and inferring similarity. That is the basic error here.

The clade ‘Ornitodira’
(= pterosaurs + dinosaurs, their last common ancestor and all descendants, Gauthier 1986) is a junior synonym for ‘Amniota’, which is a junior synonym for ‘Reptilia’ when more taxa are added to phylogenetic analysis, as demonstrated here: This growing online study currently tests 1220 specimen-based taxa throughout the Tetrapoda. So here, as nowhere else, pterosaurs have the opportunity to nest with over 1200 candidate sisters.

Pterosaur outgroups
Macrocnemus, Tanystrospheus, Tanytrachleos, Langobardisaurus, Cosesaurus and Sharovipteryx are pterosaur outgroup taxa (Peters 2000, 2007) with an oblique femoral head and sprawling femora. In Peters (2000) pterosaurs and their outgroups were considered prolacertiforms, but with additional taxa (Peters 2007 and taxa listed above join the lepidosaurs Huehuecuetzpalli and Tijubina in a new clade (Tritosauria) nesting between Rhynchocephalia (= Sphenodontia) and Squamata.

Pterosaur femur samples. A

Figure 1b. Pterosaur femur samples. Above, Pteranodon. Below, Anhanguera. Note the oblique angle of the femoral head. When the axes of the femoral neck and laterally-oriented acetabulum lined up a sprawling configuration was produced.

In pterosaurs the angle of the femoral shaft
in relation to the acetabular bowl is determined by the femoral neck, which is nearly at right angles to the shaft in the clade represented by Dimorphodon and Anurognathus. Padian famously compared erect Dimorphodon to erect birds (Padian 1987) and heartily endorsed the Ornithidira hypothesis without testing other pterosaur ancestor candidates among the Lepidosauria, some of which were not published until after 1987. In many other pterosaurs, like Anhanguera, Pteranodon and Quetzalcoatlus, the shaft and head of the femora are much more oblique (Fig. 1b), at times approaching collinear (Fig. 2). No pterosaur femora are presented in Manafzadeh and Padian 2018, only a quail pelvis and femur.

The Vienna Pterodactylus.

Figure 2. The Vienna Pterodactylus. Click to animate. Wing membranes in situ (when folded) then animated to extend them. The femora are sprawling because this is a lepidosaur, not an archosaur.


Young scientists:
Examples like Manafzadeh and Padian 2018 should inform you that even though some highly regarded paleontologists have made great discoveries and have stood up against Creationists, even they can put on blinders when it comes to direct attacks on cherished hypotheses. Neither Padian nor his students, nor any other professor nor their students, have ever, or will ever find pterosaur sister taxa among the Archosauriformes, no matter how much they believe that someday, somehow what they pray for and have faith in will happen. It’s been 18 years since the Ornithodira was struck down (Peters 2000) and pterosaurs were shown to nest outside the Archosauriformes. Padian and others simple ignore this trifle, hoping it will someday go away. And it will, unless others offer to take up the cause. Unfortunately, that’s the state of paleontology in 2018.

Everywhere, but here
testing the discoveries of others appears to be on the wane (see video below the references)… but that’s life. Question authority. Test evidence for yourself.

Gauthier JA 1986. Saurischian monophyly and the origin of birds. The Origin of Birds and the Evolution of Flight, K. Padian (ed.), Memoirs of the California Academy of Sciences 8:1–55.
Manafzadeh AR and Padian K 2018. ROM mapping of ligamentous constraints on avian hip mobility: implications for extinct ornithodirans. Proceedings of the Royal Society B Biological Sciences. Published 23 May 2018.DOI: 10.1098/rspb.2018.0727
Padian K 1983. Osteology and functional morphology of Dimorphodon macronyx (Buckland) (Pterosauria: Rhamphorhynchoidea) based on new material in the Yale Peabody Museum, Postilla, 189: 1-44.
Peters D 2000. A redescription of four prolacertiform genera and implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293-336.

John Oliver thinks the following science problem is not funny.
Academic publications are unlikely to publish studies that simply confirm earlier discoveries. And yet… science depends on confirmation and ultimately consensus.

(Click to play video). After the first few minutes the video becomes less relevant):

As Oliver puts it:
There’s no Nobel Prize for fact checking.” Perhaps that is why few other workers are even considering taxa listed in the large reptile tree and large pterosaur tree that were shown to be relevant for more focused studies. And those that do (e.g. Baron and Barrett 2017 in their Chilesaurus study) are being notably taciturn about grabbing headlines for discoveries posted and time-stamped years earlier.

Quotes from this Oliver video:
“So you have all these exploratory studies that are taken as fact, that have never actually been confirmed.” 

“Replication studies are rarely funded. No one wants to do them.”

“Too often, a small study with nuanced tentative findings gets blown out of all proportion when it is presented to us, the lay public.”



The black skimmer (Rynchops niger) enters the LRT

And it nests with
the extinct great auk (Pinguinus) in the large reptile tree (LRT, 1217 taxa). No wonder it seems so different than other living birds! It is.

Figure 1. Black skimmer (genus: Rynchops) in vivo.


These, in turn
are close to the puffin (Fratercula), which also has large keratinous extensions of the rostrum and mandible, and not one, but two mandibular fenestrae.

Figure 2. Skull of the the black skimmer (Rynchops niger) with bones colorized. Note the large keratinous extension extending the mandible.

Figure 2. Skull of the the black skimmer (Rynchops niger) with bones colorized. Note the large keratinous extension extending the mandible. Note the slight differences in the two presented skulls.

Rynchops niger (Linneaus 1758; up to 50 cm in length) is the extant black skimmer, famous for having a longer lower bill than upper. It flies low on still waters to skim for fish near the surface. Close relatives include the auk, Pinguinus and the puffin, Fratercula, all derived from skuas and petrels.

Figure 2. Pinguinus the great auk skull.

Figure 3. Pinguinus the great auk skull. This flightless extinct bird nests closest to the black skimmer in the LRT. Note the two mandibular fenestra, The curved and expanded premaxilla that extends no further posteriorly than the naris, the pinched rostrum,  the tiny ascending process of the retroarticular process, the quadratojugal that extends to the antorbital fenestra,

Prum et al. 2015
nested Rynchops with the black-headed gull (Chroicocephalus) using DNA data. They did not test the extinct Pinguinius. In the LRT, the black-headed gull nests with another clade of wading birds, beginning with the limpkin, Aramus.

Figure 1. Chroicocephalus, the black-headed sea gull in vivo and as a skeleton.

Figure 4. Chroicocephalus, the black-headed sea gull in vivo and as a skeleton. This taxon nests with hummingbirds, both derived from stilts and other wading birds like the limpikin (Aramus).






Axial rotation: fingers in pterosaurs, toes in birds

A somewhat recent paper by Botelho et al. 2015
looked at the embryological changes that axially rotate metatarsal 1 to produce a backward-pointing, opposable, perching pedal digit 1 (= hallux).

Hallux rotation phylogenetically
Botelho reports: Mesozoic birds closer than Archaeopteryx to modern birds include early short-tailed forms such as the Confuciusornithidae and the toothed Enantiornithes. They present a Mt1 in which the proximal portion is visibly non-twisted, while the distal end is offset (“bent”) producing a unique “j-shaped” morphology. This morphology is arguably an evolutionary intermediate between the straight Mt1 of dinosaurs and the twisted Mt1 of modern birds, and conceivably allowed greater retroversion of Mt1 than Archaeopteryx.”

“D1 in the avian embryo is initially not retroverted9, and therefore becomes opposable during ontogeny, but no embryological descriptions address the shape of Mt1, and no information is available on the mechanisms of retroversion.”

Figure 1. Pes of the most primitive Archaeopteryx, the Thermopolis specimen.

Figure 1. Pes of the most primitive Solnhofen bird, the Thermopolis specimen. This digit 1 never left the substrate.

In Tyrannosaurus,
(Fig. 2) the entire metatarsal 1 with pedal digit 1 is rotated just aft of medial by convergence. It’s not axially rotated. It’s just attached to the palmar side of the pes. This pedal digit 1 was elevated above the substrate.

Figure 2. The semi-retroverted pedal digit 1 of Tyrannosaurus rex in two views.

Figure 2. The semi-retroverted pedal digit 1 of Tyrannosaurus rex in two views. This digit 1 was elevated above the substrate.

In some birds
like the woodpecker, Melanerpes, and the unrelated roadrunner, Geococcyx, pedal digit 4 is also retroverted. Sorry, I digress.

Further digression
The axial rotation of pedal digit 1 in birds is convergent with the axial rotation of metacarpal 4 in Longisquama (Fig. 3) and pterosaurs. In both taxa the manus was elevated off the substrate and permitted to develop in new ways. Manual digit 4 never leaves an impression in pterosaur manus tracks… because it is folded, like a bird wing, against metacarpal 4. In Longisquama such extreme flexion is not yet possible.

Figure 1. Longisquama left and right manus traced using DGS then reconstructed (below). This is a very large hand for a fenestrasaur and manual digit 4 is oversized, as in pterosaurs.

Figure 3. Longisquama left and right manus traced using DGS then reconstructed (below). This is a very large hand for a fenestrasaur and manual digit 4 is oversized and the metacarpal is axially rotated, as in pterosaurs. Manual digit 5 is useless, but not yet a vestige. A pteroid is present, as in Cosesaurus. The coracoid is elongate as in birds. The sternum, interclavicle and clavicle are assembled into a single bone, the sternal complex, as in pterosaurs.

Note the lack of space between
the radius and ulna in Longisquama. This is what also happens in pterosaurs. It prevents the wrist from pronating or supinating, as in birds and bats… which means, the forelimb is flapping, not pressing against the substrate, nor grasping prey. That means all those images of Longsiquama on all fours are bogus. Now you know.

So now we come full circle
While the toes of birds axially rotate and the wing metacarpal of pterosaurs axially rotates, the forearms of birds, pterosaurs and Longisquama do not axially rotate. No one wants their wing to twist.

Botelho JF, Smith-Paredes D, Soto-Acuña S, Mpodozis J, Palma V and Vargas AO 2015. Skeletal plasticity in response to embryonic muscular activity underlies the development and evolution of the perching digit of birds. Article in Reports · May 2015 DOI: 10.1038/srep09840

A 1986 tribute to every paleontologist’s favorite cartoonist

Here are 14 minutes of a 1986 interview with Gary Larson
whose Far Side cartoons decorated the doors of every professional paleontologist back in the day. Click on the image to view in YouTube.

The Far Side series ended
on January 1, 1995 with Larson’s retirement.

Figure 1. Gary Larson, recent photo.

Figure 1. Gary Larson, recent photo.

Larson has asked people
not to use Far Side cartoons on the internet, writing a widely distributed letter in which he explains the “emotional cost” to him of people displaying his cartoons on their websites and asks them to stop doing so. If you want to see his work, it’s on his website below.



Bergamodactylus (basal pterosaur) back ‘under the microscope’

This all started with Kellner 2015
who proposed 6 states of pterosaur ontogeny based on skeletal fusion of discrete elements. This hypothesis was tested in phylogenetic analysis and shown to be invalid. Pterosaurs don’t fuse bones during ontogeny. Fusion appears in phylogenic patterns. Oblivious to this fact, Dalla Vecchia 2018 dismissed Kellner’s hypothesis by writing, “Kellner’s six ontogenetic stages are an oversimplification mixing ontogenetic features of different taxa that probably had distinct growth patterns. Finding commonality across all pterosaurs is impossible, because there is much variation in pterosaur ontogeny and the available sample is highly restricted.” 

Neither Kellner nor Dalla Vecchia recognize
the lepidosaurian affinities of pterosaurs, and do not realize that as lepidosaurs pterosaurs mature differently than archosaurs. Some lepidosaurs continue growing after fusing elements (Maisano 2002). Others never fuse elements. Fusion of elements in pterosaurs is phylogenetic, not ontogenetic. Pterosaurs mature isometrically, not allometrically as proven by every full-term embryo and every known juvenile among a wide variety of pterosaur specimens. Plus, all of the small purported Solnhofen juveniles phylogenetically nest as key transitional taxa linking larger long-tail primitive pterosaurs to larger short-tail derived pterosaurs (Peters 2007). That’s how those clades survived the extinction events that doomed their fellow, larger, longer-tailed kin.

Kellner 2015 also
distinguished a small pterosaur MPUM 6009 from the holotype of Eudimorphodon and from Carniadactylus (MFSN 1797, Dalla Vecchia 2009; Fig. 1) and gave MPUM 6009 the name Bergamodactylus (Fig. 1) after Peters 2007 had done the same (without renaming MPUM 6009), in phylogenetic analysis. Neither Kellner nor Dalla Vecchia performed a phylogenetic analysis, but preferred to describe similar bones. That rarely works out well.

Figure 1. Bergamodactylus compared to Carniadactylus. These two nest apart from one another in the LRT.

Figure 1. Bergamodactylus (MPUM 6009) compared to Carniadactylus (MFSN 1797). These two nest apart from one another in the LRT. Contra Dalla Vecchia 2018, these two share relatively few traits in common. The feet, cervicals, sternal complex coracoids and legs are different.

Dalla Vecchia 2018 concludes, 
“The anatomical differences between MPUM 6009 and MFSN 1797 are too small to support the erection a new genus for MPUM 6009.” That is incorrect (Fig. 1). Several taxa nest between these two taxa in the large pterosaur tree (LPT, 232 taxa). Their feet alone (Fig. 1) were shown to be very different in Peters (2011).


Figure 2. Bergamodactylus compared to Cosesaurus. Hypothetical hatchling also shown.

From the Dalla Vecchia 2018 abstract
“Six stages (OS1-6) were identified by Kellner (2015) to establish the ontogeny of a given pterosaur fossil. These were used to support the erection of several new Triassic taxa including Bergamodactylus wildi, which is based on a single specimen (MPUM 6009) from the Norian of Lombardy, Italy. However, those ontogenetic stages are not valid because different pterosaur taxa had different tempos of skeletal development. Purported diagnostic characters of Bergamodactylus wildi are not autapomorphic or were incorrectly identified. Although minor differences do exist between MPUM 6009 and the holotype of Carniadactylus rosenfeldi, these do not warrant generic differentiation. Thus, MPUM 6009 is here retained within the taxon Carniadactylus rosenfeldi as proposed by Dalla Vecchia (2009a).” \

Dalla Vecchia is basing his opinion on comparing a few cherry-picked traits, possibly convergent, rather than running both taxa and a long list of other pterosaurs through phylogenetic analysis, to see where unbiased software nests both taxa among the others.

Plus, as mentioned above, both authors are working from an antiquated set or rules that no longer apply now that pterosaurs have been tested and validated as lepidosaurs.

Figure 2. Bergamodactylus skull colorized with DGS and reconstructed.

Figure 3. Bergamodactylus skull colorized with DGS and reconstructed. Palatal and occipital bones shown here were missed by Dalla Vecchia 2018 and prior workers who did not use DGS.

Phylogenetic analysis
employing a large gamut of taxa, like the large reptile tree (LRT, 1215 taxa), invalidates traditional arguments that pterosaurs arose without obvious precedent among the archosauriforms, which most pterosaur workers, including both Kellner and Dalla Vecchia, still cling to, despite no evidence of support. Pterosaurs arose from fenestrasaur tritosaur lepidosaurs (Fig. 7).

Figure 4. The skull of Bergamodactylus traced by Kellner 2015, Dalla Vecchia 2018 and by me using DGS.

Figure 4. The skull of Bergamodactylus traced by Kellner 2015, Dalla Vecchia 2018 and by me using DGS. See figure 2 for a reconstruction of the DGS tracing.  Prior authors missed all the palatal and occipital bones along with several others.

The metacarpus of Bergamodactylus
has a few disarticulated elements. When replaced to their in vivo positions the axial rotation of metacarpal 4 (convergent with the axial rotation of pedal digit 1 in birds) enables the wing finger to fold in the plane of the hand, not against the palmar surface. Manual digit 5, a vestige, goes along for the ride, rotating the dorsal surface of the hand (Fig. 5).

Figure 5. Metacarpus of Bergamodactylus (MPUM 6009) in situ and reconstructed.

Figure 5. Metacarpus of Bergamodactylus (MPUM 6009) in situ and reconstructed. Apparently the pteroid splintered apart, overlooked by those with direct access to the specimen. The distal carpals are not co-ossified, as they are in later pterosaurs. The laterally longer fingers, up to digit 4, is a tritosaur trait. Note ungual 1 lies on top of the posterior face of metacarpal 4. That was overlooked by those who had direct access to the specimen, which supports the utility of DGS.


Bergamodactylus, as the most basal pterosaur,
is itself a transitional taxon bridging non-volant fenestrasaurs with all other pterosaurs. And the wing (Fig. 6) was about the last thing to evolve.

Figure 6. Click to enlarge. The origin of the pterosaur wing and the migration of the pteroid and preaxial carpal. A. Sphenodon. B. Huehuecuetzpalli. C. Cosesaurus. D. Sharovipteryx. E. Longisquama. F-H. The Milan specimen MPUM 6009, a basal pterosaur.

Bergamodactylus to scale
with Cosesaurus and Longisquama (Fig. 7), demonstrate the variety within the Fenestrasauria. Pterosaurs arose more or less directly from a sister to Cosesaurus (based on overall proportions), but note that both Sharovipteryx and Longisquama have more pterosaurian traits than Cosesaurus does. This pattern is convergent with that of birds, of which several clades of Solnhofen bird descendants arose of similar yet distinct structure.

Figure 8. Taxa at the genesis of pterosaurs: Cosesaurus, Longisquama and Bergamodactylus.

Figure 7. Taxa at the genesis of pterosaurs: Cosesaurus, Longisquama and Bergamodactylus.

See rollover images
of Bergamodactylus in situ here. You’ll see how DGS is able to pull out post-cranial details overlooked by others in the chaos and confusion of layers of bones and impressions in MPUM 6009. Cranial details are best seen in figure 3 above, which is based on higher resolution images.

Dalla Vecchia FM 2009. Anatomy and systematics of the pterosaur Carniadactylus gen. n. rosenfeldi (Dalla Vecchia, 1995). Riv. It. Paleontol. Strat., 115: 159-186.
Dalla Vecchia FM 2018. Comments on Triassic pterosaurs with a commentary on the “ontogenetic stages” of Kellner (2015) and the validity of Bergamodactylus wildi.  Rivista Italiana di Paleontologia e Stratigrafia 124(2): 317-341. DOI:
Kellner AWA. 2015. Comments on Triassic pterosaurs with discussion about ontogeny and description of new taxa. Anais Acad. Brasil. Ciênc., 87(2): 669-689.
Maisano JA 2002. Terminal fusions of skeletal elements as indicators of maturity in squamates. Journal of Vertebrate Paleontology 22:268-275.
Peters D. 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification. Ichnos 18(2):114-141.

The last common ancestor of all dinosaurs in the LRT: ?Buriolestes

Müller et al. 2018
describe a new dinosaur skeleton they attribute to Buriolestes shultzi (Cabreria et al. 2016, ULBRA-PVT280, Figs. 2, 3). In the large reptile tree (LRT, 2015 taxa; subset Fig. 1) the holotype now nests at the base of the Phytodinosauria. The referred specimen is different enough to nest between the herrerasaurs and all other dinosaurs. This, of course, removes herrerasaurs from the definition of the Dinosauria (Passer + Triceratops, their last common ancestor (= CAPPA/UFSM 0035) and all descendants).

Figure 1. Subset of the LRT including the new specimen of Buriolestes (CAPPA/UFSM 0035) nesting at the base of all dinosaurs.



Buriolestes schultzi (Cabreria et al. 2016; Late Triassic, Carnian; 230mya) was originally and later (Müller et al. 2018) considered a carnivorous sauropodomorph, but here two specimens nest as the basalmost dinosaur (CAPPA/UFSM 0035) and the basalmost phytodinosaur (ULBRA-PVT280).

Figure 2. The two skulls attributed to Buriolestes (holotype on the right). The one on the left nests as the basalmost dinosaur, basal to theropods and phytodinosaurs.

Figure 2. The two skulls attributed to Buriolestes (holotype on the right). The one on the left nests as the basalmost dinosaur, basal to theropods and phytodinosaurs. It should have a distinct name.

All cladograms agree that Buriolestes
is a very basal dinosaur. Taxon exclusion changes the tree topology of competing cladograms. The broad autapomorphic ‘eyebrow’ of the CAPPA specimen indicates it is a derived trait in this Late Triassic representative of an earlier genesis.

Figure 3. Herrerasaurus, Buriolestes and Tawa to scale.

Figure 3. Herrerasaurus, Buriolestes and Tawa to scale.

The Müller et al. cladogram
combined both specimens attributed to Buriolestes (never a good idea, but it happens all the time). The Müller et al. cladogram excluded a long list of basal bipedal crocodylomorpha, but did include Lewisuchus. It excluded the archosaur outgroups PVL 4597Turfanosuchus and Decuriasuchus. The Müller et al. cladogram nested Ornithischia basal to Saurischia (= Herrerasauridae + Agnophitys, Eodromaeus, Daemonosaurus + Theropoda + Sauropodomorpha) with Buriolestes nesting between Eoraptor and Panphagia. The CAPPA specimen of Buriolestes is also a sister to the basalmost theropod, Tawa (Fig. 3)… and not far from the other basal archosaur, Junggarsuchus (Fig. 4).

Figure 8. The CAPPA specimen of Buriolestes compared to the more primitive Junggarsuchus, basal to the other branch of archosaurs, the crocs.

Figure 4. The CAPPA specimen of Buriolestes compared to Junggarsuchus, basal to the other branch of archosaurs, the crocs.

Cabreira SF et al. (13 co-authors) 2016. A unique Late Triassic dinosauromorph assemblage reveals dinosaur ancestral anatomy and diet. Current Biology (2016),
Müller RT et al. (5 co-authors 2018. Early evolution of sauropodomorphs: anatomy and phylogenetic relationships of a remarkably well-preserved dinosaur from the Upper Triassic of southern Brazil. Zoological Journal of the Linnean Society, zly009 (advance online publication) doi: