Megachirella: Not at the origin of squamates. Lacertulus is older.

We looked at Lacertulus and the origin of the Squamata in the Late Permian
here in October 2011.

We looked at the splitting of the Tritosauria from the Protosquamata
here in December 2014.

Siimòes et al. 2018
proposed to nest Megachirella watchtleri (Fig. 1) at the origin of squamates in the Middle Triassic, 75 million years earlier than the previously known oldest squamate fossils. They reported, “For the first time, to our knowledge, morphological and molecular data are in agreement regarding early squamate evolution, with geckoes—and not iguanians—as the earliest crown clade squamates. Divergence time estimates using relaxed combined morphological and molecular clocks show that lepidosaurs and most other diapsids originated before the Permian/Triassic extinction event, indicating that the Triassic was a period of radiation, not origin, for several diapsid lineages.”

Figure 1. New µCT scans of Megachirella from Simoes et al. 2018.

Figure 1. New µCT scans of Megachirella from Simoes et al. 2018.

|they did not include relevant taxa. According to the large reptile tree (LRT, 1224 taxa, Megachirella nests at the base of the Rhynchocephalia (= Sphenodontia) along with Pleurosaurus (excluded from the Simoes team study) when many more relevant taxa are included.

Figure 2. Megachirella nests in the middle of this cladogram, that also nests turtles between rib gliders and choristoderes.

Figure 2. Megachirella nests in the middle of this cladogram, that also nests turtles between rib gliders and choristoderes.


Lacertulus is older (Late Permian) and more directly related to squamates.

FIgure 2. Megachirella (Renesto and Posenato 2003) is a sister to the BSRUG diapsid.

FIgure 3. Megachirella (Renesto and Posenato 2003) is a sister to the BSRUG diapsid and reconstructed here.

Nesting turtles with rib gliders
(Coelurosauravus) only hints at major flaws in the Simoes et al. cladogram topology. Nesting Sophineta and Palaegama close to and basal to Megachirella confirms findings made years earlier by the LRT. Marmoretta is also close, but nests within the Rhynchocephalia in the LRT.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa.

Tijubina (which Simoes redescribed in 2012) is also missing from the Simoes et al. 2018 study.

Figure 1. Palaegama is basal to Coelurosauravus ('rib' gliders), Megachirella (rhynchocephalians), Lacertulus (protosquamates) and Tijubina (tritosaurs)

Figure 5. Palaegama is basal to Coelurosauravus (‘rib’ gliders), Megachirella (rhynchocephalians), Lacertulus (protosquamates) and Tijubina (tritosaurs)



Simòes T, and 8 co-authors 2018. The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature 557: 706â709 (2018)


More evidence that black vultures are ugly pigeons

There’s at least one pigeon larger than a vulture.
It’s Goura, the Victorian crowned pigeon (Fig.s 1, 2). Goura is  the same size or larger than Coragyps, the black vulture (Fig. 1) and these two nest together in the large reptile tree (LRT, 1224 taxa). Smaller pigeons, like Columba and Caloenas nest together, next to Goura + Coragyps.

Figure 1. The largest pigeon, Goura, nests with Coragyps the black vulture, not with Columba, the rock pigeon.

Figure 1. The largest pigeon, Goura, nests with Coragyps the black vulture, not with Columba, the rock pigeon.

Goura cristata (Pallas 1764; Stephens 1819; Figs. 1, 2) is the extant Western crowned pigeon. It is restricted to New Guinea. It eats fruits and seeds.

Figure 2. Victorian crowned pigeon (genus: Goura) skeleton. Compare to figure 3.

Figure 2. Victorian crowned pigeon (genus: Goura) skeleton. This taxon nests with the black vulture, Coragyps, in the LRT. Compare to figure 3.

Coragyps atratus (LaMout 1853; 56-74 cm in length, 1.5m wingspread; Fig. 3) is the extant black vulture and a sister to Goura (Fig. 2). Both were derived from the more primitive giant petrel Macronectes. There are not very many differences between these two skeletons, perhaps one of the reasons bird workers have given up analyzing bone shapes and proportions and have taken to trusting DNA analyses.

black vultures are New World vultures. They are not related to Old World vultures in the LRT, or in any other analysis. At present this is the only New World vulture in the LRT. Old world vultures, like Torgos, nest with birds of prey.

Figure 3. When vultures drift north and start finding fish attractive they evolve into auks and puffins.

Figure 3. Skeleton of Coragyps, the black vulture. Compare to figure 2.

Beautiful as adults…
not so beautiful as chicks, Goura hatchlings look more like black vultures before they get their silky blue adult plumage. This is neotony at work once again. We’re going to look more and more at neotonous sisters and compare them to short-legged, short rostrum chicks of longer-legged, longer-beaked taxa. This is not an isolated incidence.

FIgure 4. Is it any wonder that the Goura chick is so colorless and ugly, given its relation of Coragyps, the black vulture?

FIgure 4.The Goura chick is so colorless and ugly. This makes sense given its relation of Coragyps, the black vulture. And now we know which came first, the pigeon or the vulture. The big pigeon came first, followed by smaller and smaller taxa. 

Gmelin JF 1789. Caroli a Linné … Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, Cum Characteribus, Differentiis, Synonymis, Locis. Editio Decima Tertia, Aucta, Reformata/ cura Jo. Frid. Gmelin. Volume 1, part 3. Lipsiae: Impensis Georg. Emanuel. Beer.
Gray GR 1840. List of Gen. B:59
LeMaout JEM 1853. Les trois regnes de la nature. Regne animal. Histoire naturalle des oiseaux, suivant la classification de M. Isidore Geoffroy-Saint-Hillaire, avec l’indication de leurs moeurs et de leurs rapports avec les arts, le commerce et l’agriculture. Par Emm. Le Maout. L. Curmer. Paris 425 pp.
Pallas PS 1764. Adumbratiunculae avium variorum praecedenti Elencho inserlarum, sed quae in Systemate Naturae Illustr. Linnaei nondum extant. Pp. 1-7 in Vosmaer 1764.
Vieillot LJP 1809. Histoire naturelle des oiseaux de l’Amérique Septentrionale

wiki/Coragyps atratus

What is a limpkin? (genus: Aramus)

Figure 1. The limpkin (Aramus guarauna) is a basal member of the x family.

Figure 1. The limpkin (Aramus guarauna) is a long-legged, wading basal member of the x family.

Aramus guarauna (Linneaus 1766) is the extant limpkin. It is often considerd transitional between rails and cranes. In the large reptile tree (1121 taxa) the limpkin nests basal to seagulls and hummingbirds, plovers and crowned cranes, common cranes and stilts, terns and loons, kingfishers and jabirus, murres and penguins.

Figure 1. Skeleton of the limp kin (Aramus), traditionally nests within the crane and rail order Gruiformes.

Figure 2. Skeleton of the limpkin (Aramus), traditionally nests within the crane and rail order Gruiformes. In the LRT rails are not closely related, so Gruiformes should no longer include rails.

Extant limpkins eat snails.
Primitive limpkins like Aramournis  probably had a more diverse diet. It is known from a distal tarsus.

Traditional rails
like the corn crake (Crex) and the coot (Fulica) are much more basal birds that give rise to chickens, sparrows and parrots. Adding Rallus, the Virginia rail, to the LRT nests it between Aramus and the rest of the clade, which, phylogenetically makes hummingbirds, terns and penguins variations on the rail theme and Rallus at least a Middle Cretaceous taxon radiation.

Figure 4. Virginia rail alongside the rail clade in the LRT.

Figure 4. Virginia rail alongside the rail clade in the LRT.

Congeneric specimens of Aramus
are found in the Miocene, but more derived penguins are found in the Paleocene, pointing to a mid-Cretaceous radiation of this clade.

Limpkins are derived from Cretaceous sisters to
hamerkops (Scopus) and stone curlews (Burhinus), both long-legged taxa. By the evidence shown in the crown bird subset of the LRT (Fig. 4), long legs, like those shown by Aramus, the limpkin, are basal traits. The retention of hatchling short legs occurred several times by convergence, sometimes during the Cretaceous. See the earlier post on post K-T non-arboreal birds. 

Figure 4. Subset of the LRT focusing on the crown bird clade. Brown taxa are all long-legged. Neotony produces the smaller, shorter-legged, arboreal taxa.

Figure 4. Subset of the LRT focusing on the crown bird clade. Brown taxa are all long-legged. Neotony produces the smaller, shorter-legged, arboreal taxa.

Linneaus C von 1766. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. pp. 1–532. Holmiæ. (Salvius)


Bird, pterosaur, dinosaur simplified chronology

Following the earlier post on non-arboreal post K-T boundary birds…

…this one pretty much speaks for itself.
Here (Fig. 1) is a chronology, very much simplified, of birds, pterosaurs and dinosaurs according to the LRT.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades.

Figure 1. Mesozoic chronology of bird, dinosaur and pterosaur clades based on taxa in the LRT.

If you’re curious about any of the taxa,
in the chronology, simply use Keywords to locate them.

Owlets: aptly named!

Owlets, like Aegotheles,
are like little owls, like Tyto, but they are not carnivores. Owlets are large-eyed nocturnal insectivores that feed on the wing. They are also transitional to swifts, like Apus, large-eyed diurnal insectivores that feed on the wing.

Figure 1. Aegotheles skull and in vivo. This clade is transitional from owls to swifts.

Figure 1. Aegotheles skull and in vivo. This clade is transitional from owls to swifts.

owlets are considered Caprimulgiformes. According to Wikipedia, “Traditionally, they were regarded, on morphological grounds, as being midway between the owls (Strigiformes) and the swifts. Like the owls, they are nocturnal hunters with a highly developed sense of sight, and like the swifts they are excellent flyers with small, weak legs.”

Figure 3. Tyto, the barn owl, nests very close to Apus, the swift.

Figure 2. Tyto, the barn owl, nests very close to Apus, the swift.

However… and I hate to tell you this,
Wikipedia reports, “Based on analysis of DNA sequence data – the families of the Caprimulgiformes to be members of the proposed clade Metaves, which also includes the hoatzin, tropicbirds, sandgrouse, pigeons, kagu, sunbittern, mesites, flamingos, grebes and swifts and hummingbirds (Fain and Houde 2004).

Figure 3. Skull of Apus, the common swift, closer to hawks and owls.

Figure 3. Skull of Apus, the common swift, closer to owlets and owls.

Prum et al. 2015, also using molecules,
nested Aegotheles basal to swifts and hummingbirds within a basal clade of owlet-nightjars derived from screamers (genus: Chauna) and currosaws (genus: Crax), both heavy-bodied ground birds.

The results of these and other bird DNA studies
should have reported that DNA analyses do not produce a tree topology that produces a gradual accumulation of traits, and so fail to echo trait studies. Instead, bird workers put their faith in what they could not see directly. Worse yet, they no longer believed the hard evidence that traits display. What happens to science when we no longer believe hard evidence? You get Fain and Houde 2004 and Prum et al. 2015.

Fain, MG and Houde P 2004. Parallel radiations in the primary clades of birds. Evolution. 58 (11): 2558–2573. doi:10.1554/04-235
Prum et al. (6 co-authors) 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526:569–573. online


Balearica: a unique ‘crane’ with skull bumps

This came with some surprise.
The gray-crowned crane (Balearica regulorum) has beautiful plumage, but under the skin this bird has twin skull bumps on the posterior frontals (Fig. 1).

Figure 1. Balearica regulorum in vivo and two skulls (showing individual variation).

Figure 1. Balearica regulorum in vivo and two skulls (showing individual variation).

Distinct from most cranes,
Balearica has a short rostrum (derived from Charadrius, a neotonous crane with juvenile proportions and size as an adult, based on sister taxa in the large reptile tree, LRT, 1221).

Figure 2. Balearica compared to its sister in the LRT, Charadrius, the plover/kildeer.

Figure 2. Balearica compared to its sister in the LRT, Charadrius, the plover/kildeer.

Balearica regulorum (= Ardea regulorum, Anthropoides regularum, Bennett 1834; extant; 1m tall, 2m wingspan) is the gray crowned crane, and a member of the Gruidae/Gruiformes. In the LRT Balearica is most closely related to the neotenous plovers and kildeers (genus: Charadrius,Fig. 2) and shares with them, a short bill. Twin bumps appear on the posterior frontal. Only four phalanges appear on pedal digit 4, which is as long as pedal digit 3. This trait pops up occasionally, apparently autapomorphic each time.

Using DNA
Prum 2015 nested Balearica with another crane, Grus, the limp kin, Aramus, and the trumpeter, Psophia, which is more closely related to roadrunners and cuckoos in the LRT. Prum 2015 nested Charadrius with Burhinus, close to nestings in the LRT, far from Balearica.

Olson 1985 reports
“From North America there is now a considerable representation of small to medium-sized cranes that are closely related to the modern African crowned cranes of the genus Balearica.” That makes sense with so many plovers and killdeer in North America. I see them all the time on St. Louis parking lots. Never thought they were related the one of the most beautiful birds on the African savanna.

Bennett ET 1834. On two new species of Crowned Cranes [Anthropoidea, Vivil.] from Africa. Zoological Society Proceedings pt. 1, 1833:118–119. Oken, Isis, 1835, col. 549–550.
Olson S 1985. The fossil record of birds pp. 80–218 in Farner DS, King JR and Parkes KC (eds.) Avian Biology 8: chapter 2, Academic Press, Inc.
Prum et al. (6 co-authors) 2015. A comprehensive phylogeny of birds (Aves) using targeted next-generation DNA sequencing. Nature 526:569–573. online


Ashland, Oregon, pterosaur exhibit

I recognized some of the models
in this traveling pterosaur exhibit (Fig. 1). They came from Triebold Paleontology, but originated at David Peters Studio. Unfortunately, I’m not keen on the poses the exhibitors gave these pterosaurs. Ironically they are sprawling, like lizards, but no one, but yours truly has adopted the lepidosaur origin of pterosaurs hypothesis.

FIgure 1. Pterosaur exhibit

Here’s the way
Pterodaustro should be posed (Fig. 2). It’s a wader, built for walking knee deep into still shallow waters to dip that long filter-toothed mandible — yet able, in a moment to leap into the air and take flight. Still the hind limbs are sprawling.

Figure 2. Pterodaustro sculpture

Here’s the way
Jeholopterus should be posed (Fig. 3), digitigrade with the shoulders directly over the toes.  The long finger claws of this vampire pterosaur were ideal for latching onto and in to, dinosaur skin. And this is the correct skull. Triebold, tossed out the correct skull and placed the more popular, but invalid, Bennett anurognathid skull (the one that mistook the mandibles for sclerotic (eyeball) rings.

Model of Jeholopterus, the famous vampire pterosaur.

Figure 2. Model of Jeholopterus, the famous vampire pterosaur.

Here’s the way
Dimorphodon would run around on the ground – digitigrade (proximal phalanges elevated, too), shoulders over the toes, pedal digit 5 retroverted. Nothing clumsy or awkward about this basal pterosaur! It was fast, agile and keeping those large finger claws sharp for clinging to tree trunks.


Figure x. Dimorphodon skeleton.

Figure x. Dimorphodon skeleton. The tail, not found with the rest of the skeleton, making this a chimaera, is too long.

Here’s the text of the online article:
“Long before Tyrannosaurus rex, the world was filled with pterosaurs — bizarre-looking flying reptiles, some as huge as a fighter jet, who ate everything, terrorizing the Mesozoic Age for 160 million years until they, like the dinosaurs, were killed off by a big asteroid.

“That little-known world, our window into which has been vastly expanded by science in the last few decades, has been recreated in a stunning new ScienceWorks Hands-On Museum exhibit that invites you, at one point, to virtually think, feel and fly like a pterosaur by flexing your arms and body.

“Entirely created by the Ashland’s museum staff and volunteers, it was built for under $250,000, a modest amount among museums these days, especially when they saw a San Francisco museum did their pterosaur exhibit for 30 times that, says Steve Utt, co-creator and president of ScienceWorks board of directors.

“Costs for ScienceWorks can be recouped by leasing it out, he says.

“A self-described “Silicon Valley escapee” eight years ago, Utt did all the seemingly magical if not miraculous software and video that plops you right in the middle of the pterosaur’s world, which started 228 million years ago and seems a lot stranger than any science fiction movie.

“Pterosaurs (pronounced “terra-soars”) have replaced the once terrible tyrannosaurus rex, hero of Jurassic Park, as an object of fascination because, says ScienceWorks exhibit director Leo Palombo, “there have been so many discoveries, so much we didn’t know about 10 or 15 years ago, and that’s what you see here — flying reptiles. They are not dinosaurs, not birds. Some had hair, not feathers — so many amazing sizes and shapes.”

“They all used to be called pterodactyls, but that word is outmoded now and applies only to a small subcategory. Displays at ScienceWorks seek to show the immense, newly-discovered range of body types, sizes, combs (those wild shapes on top of their heads), as well as their body architecture, which can only be described as an extremely inventive chapter of evolution.

“Displays explain that pterosaurs in general had long, pointy heads, usually with teeth, could fly up to 70 mph and would gather food by scooping it from water, land or air. They are not like bats, though they have skin-like wings, and these were made possible by the evolution of the fourth finger to hold a wing.

“Many of the exhibits teach you what various species did, how and where they did it — and then you turn around and there’s a video of a familiar beach on the Oregon coast with a couple of pterosaurs soaring in among the breakers, then alighting on our big beach rocks, where they sit and peck and preen. It’s just, simply, hard to believe this ever happened in what’s now Oregon, let alone that we have an accurate, scientific depiction of it.

“Len Eisenberg of ScienceWorks’ science advisory board stands at the most popular interactive pterosaur “ride,” urging participants to arch their heads back and wave their arms, as sensors pick up all these cues. There’s a learning curve and most who try it get chomped by a giant-jawed mososaur (sic) when they crash in the water. You get points for various foods you kill — squid, fish or ammonite. A sign shows the best score of the day, a 20, and you, usually have zero. It takes several times in a long line to get up to the skill of the pterosaur.

“This display and the science around pterosaurs is interesting because we’ve found lots more fossils and footprints in the last decade,” says Eisenberg, “all of which explain how they lived and got food.”

“Another interactive ride shows a seeming x-ray of your flapping human thorax, set beside the ancient creature and giving us a window on how much muscle and thin, fragile, lightweight bone had to be brought into play for it to fly.

“The stunning centerpiece of the new exhibit is the lifesize, 16-foot tall wood model of Quetzalcoatlus, the largest known flying creature of all time, which exhibit technician Rachel Benbrook and others fashioned using Turbo CAD and Adobe Illustrator.

“The public reception to this exhibit has been overwhelmingly positive and,” she says, “many people, seriously, have been blown away. That’s what we want — to inspire and encourage science education to the next level.”
— John Darling is an Ashland freelance writer. Reach him at”


Post K-T event birds were all non-arboreal…

…whether tested using DNA or skeletal traits…

Field et al. 2018
used DNA to produce a cladogram of extant birds to determine that basal taxa were all non-arboreal. Earlier the large reptile tree (LRT, 1121 taxa) came to the same conclusion using trait analysis and fossils (Figs. 1, 2). The only difference is the Field team determined that the crown bird radiation was post-Cretaceous. The LRT recovered a crown bird radiation in the post-Jurassic and post-Cretaceous birds were also non-arboreal (Fig. 1, 2). An earlier radiation explains the Paleocene appearance of very derived fossil penguins and the Early Cretaceous appearance of the fossil chicken, Eogranivora.

Figure 2. Basal bird phylogeny based on the LRT (morphology)

Figure 1. Basal bird phylogeny based on the LRT (morphology.

Field et al. also recovered flamingos with grebes, chickens with ducks, and many other physical trait mismatches, like those in Prum et al. 2015. Such mismatches are ignored by DNA workers.

Figure 1. Click to enlarge. Duck origins recovered by the LRT. Duck descendants were long-legged walkers and later waders.

Figure 2. Click to enlarge. Duck origins recovered by the LRT. Duck descendants were long-legged walkers and later waders.

From the Field et al. 2018 abstract:
“We suggest that ecological filtering due to the temporary loss of significant plant cover across the K-Pg boundary selected against any flying dinosaurs (Avialae) committed to arboreal ecologies, resulting in a predominantly non-arboreal post-extinction neornithine avifauna composed of total-clade Palaeognathae, Galloanserae, and terrestrial total-clade Neoaves that rapidly diversified into the broad range of avian ecologies familiar today. The explanation proposed here provides a unifying hypothesis for the K-Pg-associated mass extinction of arboreal stem birds, as well as for the post-K-Pg radiation of arboreal crown birds.”

the loss of an arboreal habitat due to world-wide fires does not explain the disappearance of the Cretaceous toothed sea birds, Ichthyornis and Hesperornis. Other explanations must be invoked.

Figure 1. More taxa, updated tree, new clade names.

Figure 3. More fossil taxa, updated tree, new clade names. Note the derived position of the penguin Aptenodytes, with with fossil penguins in the Paleocene.

The mechanism for crown birds invading trees
appears to have been neotony, with chick-sized adults with short legs and short necks arising from larger, long-legged, long-necked predecessors (e.g. Passer the sparrow and Opisthocomus, the hoatzin, arising  from Gallus, the chicken). Notably the hatchlings of arboreal taxa are typically not precocial, hatching out a more helpless stage in their ontogeny and growing to fledgling size rapidly.

Field et al. correctly report,
“…virtually the entirety of the avian crown-group fossil record is restricted to sediments of Cenozoic age, and the earliest well-supported crown bird fossil is scarcely older than the end-Cretaceous, at approximately 67 Ma.” True. This is one of the unresolved mysteries of paleontology, only now starting to crack with discoveries like Eogranivora, the early Cretaceous chicken, and the nesting of Cretaceous toothed birds between paleognaths and neognaths (Fig. 3), something the Field analysis was not able to recover.

Figure 1. Megapodius is the extant bird nesting at the base of all neognathae (all living birds except ratites).

Figure 4. Megapodius is the extant bird nesting at the base of all neognathae (all living birds except ratites). Post K-T event birds look more or less like this one. One might ask, if all the adults were killed, would the precocious hatchlings, hidden beneath thick mounds, form the first generation of K-T event survivors?

One might ask,

if all adult birds worldwide were killed on exposure to oven-like temperatures and subsequent lack of trees, would the buried, precocious hatchlings of mound-builders, like Megapodius (Fig. 4), create the first generation of K-T event bird survivors? If so, perhaps the tinamou ancestors of modern tinamous and ratites were likewise mound builders. Currently tinamous and ratites are not mound-builders.

Basal members of all bird clades in the LRT
appear to have survived the K-T event, based on the Paleocene presence of fossil penguins, like Waimanu (Fig. 5). Overlooked by Field et al., basal members of all the major crown bird clades in the LRT (Fig. 3) are all non-arboreal, long-legged, wading taxa (Fig. 2), that do not nest in trees.

Figure 1. The world at the K-T boundary, 65 mya and the distribution of Paleocene birds.

Figure 5. The world at the K-T boundary, 65 mya and the distribution of Paleocene birds.

Field  DJ et al. (7 co-authors) 2018. Early Evolution of Modern Birds Structured by Global Forest Collapse at the End-Cretaceous Mass Extinction. Current Biology (advance online publication) DOI:

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.

Figure 1a. 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 ‘Ornithodira’
(= 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.
Peters D 2007. The origin and radiation of the Pterosauria. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27

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.”