Padian 2017 examines pterosaur ankles with taxon and paper exclusion

I’ve had a long history with Dr. Kevin Padian,
one of the smartest paleontologists out there. He made important suggestions to my first book, GIANTS and early in his career made a name for himself by reporting on the bird-like traits of the Jurassic pterosaur, Dimorphodon. 

Unfortunately
Dr. Padian has a blind spot. He holds to the invalidated hypothesis that pterosaurs are related to dinosaurs, despite the complete lack of a series of archosaur taxa demonstrating a gradual accumulation of pterosaur traits. He still believes in the clade ‘Ornithodira.’

Ornithodira
Wikipedia reports, “Gauthier…coined and defined a slightly more restrictive node-based clade, Ornithodira, containing the last common ancestor of the dinosaurs and the pterosaurs and all of its descendants. Paul Sereno in 1991 gave a different definition of Ornithodira, one in which Scleromochlus was explicitly added.”

In the large reptile tree (LRT, 1094 taxa) the last common ancestor of dinosaurs and pterosaurs is the Devonian tetrapod, Tulerpeton at the base of the Lepidosaurormorpha – Archosauromorpha split.

Padian 2017
once again links pterosaurs with dinosaurs as he reviews with old illustrations the ankle bone ‘homologies’ of pterosaurs and archosaurs. Unfortunately he ignores Peters (2000a, b) who reidentified certain tarsals based on homologies with Cosesaurus and other fenestrasaurs (see below).

Figure 1. Peteinosaurus and Dimorphodon BMNH4212 pedes. Four tarsals are present on both.

From the Padian abstract:
“The ankle bone assembly of pterosaurs has received little attention, even though it is critical for understanding the functional morphology of the leg and the foot and has far-reaching implications for interpretations of stance and gait in ornithodirans in general, as well as for any role the leg may have had in the flight of pterosaurs. Of particular importance are the distal tarsal bones, which are seldom preserved clearly.”

Padian found only two large (medial and lateral) tarsals in Dimorphodon, but most early pterosaurs have four tarsals (Fig. 1), as some of his figures show.  In Dimorphodon and Pteranodon the distal and proximal tarsals appear to fuse to one another creating two large side-by-side tarsals with a concave surface for articulation with the tibia/fibula. In all other pterosaurs the proximal tarsals are the astragalus and calcaneum. The ‘distal tarsals’ are actually distal tarsal 4 + the centrale sometimes accompanied by a tiny distal tarsal 3 (Peters 2000a) based on homologies with several tritosaur lepidosaurs, like Macrocnemus.

“Their concave proximal facets articulate with the medial and lateral condyles (comprising the astragalus and, at least basally, the calcaneum) of the tibiotarsus.”

The proximal tarsals are not part of the tibia in pterosaurs. Pterosaurs do not fuse the tibia and tarsus to form a tibiotarsus (Peters 2000a).

“Distally, they articulate with metatarsals II–IV, and the relatively large metatarsal V articulates on the distolateral side of the lateral distal tarsal.”

Not quite. That’s distal tarsal and the calcaneum articulate with metatarsal 5. That is exactly what happens, as Padian shows, in the archosauriforms Euparkeria, Crocodylus and Lagerpeton. That is exactly what also happens in the tritosaurs Huehuecuetzpalli, Macrocnemus, Langobardisaurus, Cosesaurus and Sharovipteryx (Peters 2000a and ReptileEvolution.com).

“The homology of these bones in pterosaurs can be established with reference to other early-branching ornithodirans, and the morphology of the bones implies similar functional roles and ranges of motion.”

Convergence here with tritosaur lepidosaurs. Worth looking at.

“The medial distal tarsal is likely the fusion of distal tarsals 2 C 3, and the lateral distal tarsal is distal tarsal 4, a pattern reflected in ontogeny.”

No and yes. In tritosaurs distal tarsals 1–3 are tiny vestiges. Distal tarsal 3 is retained in many long-tailed pterosaurs. Distal tarsal 4 remains large. The proximal and distal elements fuse in Pteranodon. The medial centrale is Padian’s medial distal tarsal (Peters 2000a).

“The pterosaur ankle was capable of plantarflexion, but adduction and abduction of the feet were greatly limited.”

True.

“A synoptic survey of available tarsal bones of pterosaurs shows that the morphology of these bones remained relatively unchanged from the most basal pterosaurs to the pteranodontids and the azhdarchoids.”

True.

“Comparisons among a variety of ornithodirans show that the basic functional pattern did not vary importantly, although some ornithodiran subgroups evolved unique schemes of development and sequential ossification.”

True.

Dr. Padian writes:
“Pterosaurs were not thought to be particularly close to dinosaurs, or to any other archosaurs.”

When? That’s not current and traditional.

“Bennett, as noted above, does not accept that pterosaurs are ornithodirans. So it is all the more striking that these authors come to the same conclusion as functional morphologists who accept that pterosaurs are ornithodirans. The consensus of these authors is that pterosaurs, like dinosaurs and other ornithodirans, had a mesotarsal ankle that functioned as a hinge joint. Because the knee was also a hinge joint, as were the metatarso-phalangeal joint and the interphalangeal joints (Padian, 1983b, 1991), and the hip joint effectively allowed only protraction and retraction (see Schaeffer, 1956, and also Padian, 1983b), the gait would have been parasagittal and the stance erect (Padian, 2008). No argument has ever been made to counter these observations.”

No argument can be made to counter these observations. However, they can be expanded. Padian ignores the fact that other clades, like lepidosaurs, are also capable of bipedal locomotion and that some (like those list above) also have a simple hinge ankle joint. He also fails to note that in some pterosaurs the femoral head is at right angles to the shaft, but in others it is almost in line with the shaft, creating a splayed femur, like a lepidosaur, yet, like certain lepidosaurs, still capable of erect bipedal locomotion (Fig. 2).

Padian discusses the splayed femur concept
and agrees with Unwin that it would have provided a clumsy, sprawling gait. This is incorrect as anyone can learn from making museum-quality skeletons that have splayed femora and erect hind limbs. The angles all work out (Fig. 2).

And running bipedal lizards are not clumsy. They are speedy wonders!

Figure 2. Standing Pteranodon with sprawling femora. We’ve known this for 17 years.

Way back in the 1980s,
Kevin Padian and Chris Bennett. in the same conversation. cautioned me to employ phylogenetic analysis in my studies. Given present data in the academic literature (Peters 2000a, b) you have to ask yourself why Padian, like Bennett (2012) restricted his taxon list to just archosauromorphs.

For those who wonder why I don’t publish,
maybe Padian’s paper will offer some insight. I have published several papers on pterosaur relationships, wings and feet. None were cited by Dr. Padian. He is listed in the acknowledgments of Peters 2000a for reading an earlier version of the manuscript. The last time we e-mailed he was angry that I made several of the above observations.

References
Bennett SC 2012. The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined. Historical Biology. iFirst article, 2012, 1–19.
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.
Padian K 2017. Structure and evolution of the ankle bones in pterosaurs and other ornithodirans. Journal of Vertebrate Paleontology.
DOI: 10.1080/02724634.2017.1364651
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods.  Ichnos 7:11-41.
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.

A crow, owl and swift are added to the LRT

Today
several taxa (in yellow-green, listed below, Figs. 1-3) were added to the LRT. Along the way several scoring errors were corrected which shifted the giant sea bird with bony teeth, Pelagornis (in amber, Fig. 4), to nest with Macronectes, the petrel.

Figure 1. The latest additions to the LRT are shown in light green. Pelagornis, which has just moved to nest with petrels, is in amber.

Corvus the crow nests between the the heron, Ardea, and a large clade that includes flying sea birds, storks, ducks, vultures, hummingbirds, dippers, wrens and penguins.

Figure 2. Apus the common swift is actually a close relative of the falcon and owl, not a hummingbird.

Apus the swift nests between Falco the falcon and Tyto, the barn owl. Earlier Eocypselus was promoted as a swift + hummingbird ancestor. Here Eocypselus is only a hummingbird relative.

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

Pelagornis, the giant sea bird, has a new skull shape (Fig. 4). This came about after the realization that some birds have a very long maxilla. Other birds essentially turn the maxilla into a vestige. We’ll look at that major difference and many others in a future blog post.

FIgure 4. Pelagornis, new reconstruction of skull along with overall reconstruction from Mayr and Rubilar-Rogers

Birds are tough,
but I’m learning their details along the way. Phylogenetic analysis is a great way to dissect a taxon, trait by trait.

References
wiki/Macronectes
wiki/Pelagornis
wiki/Apus
wiki/Tyto

Spoonbill: it really is a duck-stork

Maybe the spoonbill
(Fig. 1) is the closest thing we have today to Presbyornis (Fig. 2).

Figure 1. The roseate spoonbill (genus: Platalea) in vivo. Traditionally spoonbills and storks have been nested together. Here it nests between storks and ducks.

Short one today. Pretty obvious results.
The extant spoonbill (Platalea) with its featherless face (Fig. 1), gives the impression of being very ancient. The large reptile tree (LRT, 1090 taxa) nests it between storks (check out those long legs) and ducks (check out that spoon bill), including long-legged ducks (Fig. 2) like Presbyornis.

Figure 4. Presbyornis is the prehistoric long-legged duck,

The only trouble is
spoonbills typically nest only with storks, far from ducks, which typically nest closer to screamers, like Chauna.

You can see the gradual accumulation of traits
using morphological traits. That doesn’t always happen with DNA.

From the ReptileEvolution page:
Presbyornis pervetus (Wetmore 1926; Olson and Feduccia 1980; earliest Eocene, 62 mya) is one of the first of the clade Anseriformes (ducks, geese and kin). It is known from scattered bones and was originally considered a flamingo relative, due to its long legs. The duck-like skull was found later.

References
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Olson SL and Feduccia A 1980. Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae). Smithsonian Contributions to Zoology 323: 1–24.
Wetmore A 1926. Fossil birds from the Green River deposits of Eastern Utah. Annals of the Carnegie Museum 16: 391-402.

wiki/Platalea
wiki/Anas
wiki/Presbyornis

Like dippers, wrens are not passerines either

Updated January 8, 2018
with the nesting of wrens and dippers with crows and woodpeckers.

FIgure 1. Eurasian wren, Troglodytes troglodytes, skull in three views

On a reader request that I became curious about
the Eurasian wren (Troglodytes troglodytes) has been added to the large reptile tree (LRT, 1090 taxa) because it typically nests with the tiny dipper (Cinclus), which nests with woodpeckers and crows.

Figure 1. The Eurasian wren, Troglodytes troglodytes, nests with the dipper, Cinclus, close to kingfishers and penguins.

To no one’s surprise,
the wren nests with the dipper. And both nest with woodpeckers and crows. 

Troglodytes troglodytes (Vieillot 1809; 12cm long,is the Eurasian wren.  Here the cranium is further expanded.

References
Vieillot LJP 1809. Histoire naturelle des oiseaux de l’Amérique Septentrionale

wiki/Troglodytes_wren

Roman Uchytel. Taking paleoart to the next level.

Figure 1. Roman Uchytel is the arist/naturalist who is bringing prehistoric beasts and birds back to life.

Here’s an artist worth noting.
Roman Uchytel (Fig. 1) says it best himself, “Using only their skeletons, I bring creatures to life that roamed the same routes that take you to and from work hundreds of thousands of years ago.”

His mission:
“Roman Uchytel’s galleries constitute the first resource solely dedicated to the reconstruction of prehistoric animals beyond the dinosaurs. These are not photographs, but rather, artistic recreations from the skeletons of ancient animals that roamed the earth millions of years ago. Many of these fascinating creatures are unfamiliar to the public and remain a mystery even to science.”

Figure 2. Homepage for Roman Uchytel’s images. Click to visit.

Check out his website
and you will be filled with wonder: https://prehistoric-fauna.com

 

Revisiting the pes of Pectodens

Earlier we looked at Pectodens, a long-necked tritosaur that nests at the base of the tanystropheids + langobardisaurs and the fenestrasaurs, which ultimately gave rise to pterosaurs.

Metacarpal 5 is the problem (Fig. 1).

Figure 1. The right pes of Pectodens in situ (left), sans the matrix (right), and rearranged to match sister taxa (center). The question is: is the rearrangement valid?

 

Which bone is metacarpal 5?
Is it the long bone similar to metacarpal 4? That would make sense with most taxa, except Pectodens nests with other long-necked taxa, like Langobardisaurus and Tanystropheus. In those taxa metacarpal 5 is short and pedal 5.1 is metapodial (= very long).

Did taphonomy change things?
Or do we trust phylogenetic bracketing?

One more thing…
If the long bone is the metacarpal, then the phalangeal count matches sister taxa (4 phalanges). If the short bone is the metacarpal, then there is one extra phalanx. Did the preparator add a bone? Or did this taxon have an extra bone?

And take a look
at the width of the tibia + fibula. It’s the right width if the short bone is metacarpal 5. The width is not quite wide enough if the long bone is metacarpal 5.

Sometimes
you have to make a decision in paleontology. Sometimes you have to point your finger at a preparator’s mistake. Sometimes you make the mistake when you use your brain OR when you accept the data as presented.

What to do… what to do…

Figure 2. Pectodens reconstructed using the original tracings of the in situ fossil in Li et al. 2017.

Here’s what I wrote a while back
at ReptileEvolution.com:

Pectodens zhenyuensis (Li et al. 2017; IVPP V18578; Anisian, Middle Triassic; 38cm in length) was originally considered to be a diapsid and a possible protorosaur. Here Pectodens nests between Macrocnemus and Langobardisaurus. Originally the interclavicle, sternum and quadratojugal were overlooked. Note the large orbit, the long metarsal 5 and the perforated pubis. The elongate caudal transverse processes anchor powerful leg muscles.

Figure 3. Pectodens skull traced using DGS techniques and reassembled.

With the short metacarpal 5
Pectodens cleanly nests with fewer autapomorphies at the base of the Langonbardisaurus/Tanystropheus clade.

References
Li C, Fraser NC, Rieppel O, Zhao L-J and Wang L-T 2017. A new diapsid from the Middle Triassic of southern China. Journal of Paleontology.7 pp. doi: 10.1017/jpa.2017.12

What is a flamingo in the cladogram of birds?

In the present subset
of the large reptile tree (LRT, 1090 taxa, Fig. 3) flamingoes (genus: Phoenicopterus, Fig. 1) nest with the seriema (genus: Cariama, Fig. 2). That makes the flamingo a sort of bird of prey, now concentrating on tiny plankton (algae + invertebrates). That’s why they are so distinct.

Wikipedia lists several studies
that nest flamingoes with ducks, spoonbills and/or doves.

Prum et al. 2015 used DNA
to nest flamingoes with grebes. These submersible sharp-rostrum, hind limb swimmers have not yet been added to the LRT, but grebes look more like loons, similar to Gavia. Until that analysis,  here’s a loon skeleton online. So when grebes are added, I’ll let you know how that works out.

The closest tested relative
of Phoenicopterus in the Prum et al. tree is Uria, the murre, which we looked at yesterday, Here, Uria nests between dippers and penguins, far from Phoenicopterus (Fig. 3).

So, apparently there is no consensus
out there regarding flamingo relatives. Are the flamingo-like traits of Cariama convergent or homologous? The answer has to come from comparative anatomy. DNA fails too often to deliver sisters who actually look like one could evolved from the other or from a common ancestor sharing a long list of traits.

Figure 1. Phoenicopterus, the flamingo, currently and provisionally nests with Cariama in the LRT.

Phoenicopterus chilensis (Molina 1782) is the extant flamingo, a long-legged filter-feeder with pink plumage. Here it ness with Cariama, the seriema.

Figure 2. The seriema (genus: Cariama) is the closest taxa to Yanornis in the LRT. The two resemble one another in most details, but Cariama lacks teeth, has a retracted naris and an elevated pedal digit 1.

Cariama cristatus (Linneaus 1766) is the extant seriema, a grasslands predator from South America. It flies only to escapte predators. Here it is basal to the flamingo, Phoenicopterus. At present it is easy to see why they nest together. And this is where the LRT shines.

Figure 1. Subset of the LRT focusing on birds. Here various aspects of birds are shown, including age, teeth, feeding behavior and basic clades.

If anyone can find a better match
for flamingoes, please let me know. Otherwise, you heard it here first. Meanwhile, I’m surprised to see what I learn in just a few hours has not been discovered before. This is not rocket science.

References
Linneaus C 1766. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio duodecima, reformata. Holmiae. (Laurentii Salvii).: 1-532.
Molina JI 1782. Saggio sulla Storia Naturale del Chili. Bologna, Stamperia di S. Tommaso d’Aquino. 349 pp.

wiki/Flamingo
wiki/Seriema

 

Murres and the origin of penguins

Updated January 8, 2018
with the deletion of dippers, which now nest with wrens when more taxa are added.

Updated August 11, 2021
with the revelation that the skeleton tested in 2017 was Uria aalge, mislabeled Uria lomvia below. The in vivo Uria in figure 2 is also Uria lomvia (Fig. 1).  The former still nests with penguins. The latter now nests with auks. See figure 1.

Today
we’re going to take a heretical look at the origin of penguins, those short-legged, super-insulated, flightless, fish- & squid-eaters. Some can dive for 22 minutes down to 550 meters.

According to Wikipedia
the relationships of the penguin subfamilies (order: Sphenisciformes) and the placement of penguins among the birds “is not resolved.” By contrast, in the LRT the relationship of penguins among birds is completely resolved.

Basal penguins,
like Waimanu, are known from Antarctica and New Zealand from the early Paleocene. Waimanu was flightless and likely swam with both its short wings and paddling feet. This derived bird at the K-T boundary points to a much earlier radiation of more primitive, crane-like extant birds, apparently starting just before Yanornis in the Early Cretaceous.

In the large reptile tree
(LRT, 1089 taxa, now 1898+ taxa) penguin ancestors going back to Devonian fish are recovered. However, presently and provisionally two taxa are proximal penguin sisters in the LRT and these are derived from even more basal and high-energy grebes and kingfishers.

FIgure 2. Uria aalge and Aptenodytes, two taxa in the origin of penguins. Despite their apparent differences, the LRT nests these three taxa together in a single clade.

Representing a transitional phase
Murres like Uria (Fig. 2; 45cm), traditional nest with snipes, plovers, terns, stilts, gulls puffins and auks (= Charadriiformes),. That’s a pretty diverse clade. Some of these also appear in the penguin clade of the LRT. Many workers consider murres to be unrelated to penguins, despite appearances. Murres are all north of the equator, while penguins stay south. Uria has pelican-like plumage (black and white) and is better adapted to swimming underwater (up to  4 minutes) with a longer torso and longer sternum. Digit 1 becomes a vestige and the rib cage extends nearly to the ankle. Murres live in colonies near sea waters.

Penguins like Aptenodytes (Fig. 3) traditionally nest with loons, like Gavia. Here (Fig. 1) they don’t. Penguins are flightless, trend toward larger, can swim better and seek larger prey. Finger 1 disappears. The pygostyle straightens out. The scapula grows larger. The metatarsus becomes shorter than the pedal digits. Again, these are all minor and gradual accumulations of traits.

Uria lomvia (Linneaus 1758; 45cm tall) is the extant thick-billed murre. It is a strong flyer, both in the air and underwater. Here it is basal to auks.

References
Deguine, J-C 1974. Emperor Penguin: Bird of the Antarctic. The Stephen Greene Press, Vermont.
Hackett S et al. 2008. A phylogenetic study of birds reveals their evolutionary history. Science 320:1763–1768.
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.

wiki/Penguin
wiki/Uria

African hamerkop, Antarctic petrel and American vulture

These three extant bird taxa nest together [Not anymore they don’t]
in the large reptile tree (LRT, 1089 taxa then 2024 taxa now) and it’s easy to see why when you look at their skulls (Figs. 1–3). On the inside they are more like each other than any other tested taxa. We usually see them from the outside (Figs. 4–6).

Updated December 24, 2021
with almost 950 more taxa (Fig. 9) and several bouts of housekeeping. Vultures nest with hawks. Hamerkops nest with hornbills. Petrels are now separated from hamerkops by ducks + geese.

Figure 1. Scopus umbretta, the extant African hamerkop.

Scopus umbretta (Brisson 1760, 56 cm tall) is the extant hamerkop, a mid-sized wading bird (Figs, 1, 4). In the LRT it nests with the petrel, Macronectes, Note: the maxilla is much longer in these taxa.

Figure 3. Macronectes giganteus, the extant Southern giant petrel.

Macronectes giganteus (Gmelin 1789; 56 cm tall) is the extant Southern giant petrel, a member of the Procellariidae and Procellariformes (tubenoses). Here it nests with Scopus and Pelagornis . This soaring sea bird has a wingspan up to 2m. Note the naris has shifted anteriorly along with the maxilla.

Figure 3. Coragyps atratus, the extant black vulture.

Coragyps atratus (LaMout 1853; 56 cm in length, 1.5m wingspread) is the extant black vulture and a sister to the giant petrel. Note the similar premaxilla. As in Threskiornis(below), the head and neck lack feathers.

Figure 4. Scopus, the hamerkop, in vivo.

One of the current problems in bird phylogeny
is how to tie the various orders of birds together at their roots. Which orders are related to which other orders? Here’s how the situation stands according to Wikipedea:

1. Hamerkop family: Scopidae, order: Pelecaniformes
2. Petrel family: Procellariidae, order: Procellariformes
3. Vulture family: Cathartidae: order: Accipitriformes

According to Wiki:
Pelecaniformes include the pelican, shoebills, hamerkops, ibises, spoonbills, herons, egrets and bitterns. In the LRT, SOME of these disparate taxa nest at disparate nodes. Others have not been tested yet.

Frigate birds, gannets and boobies, cormorants, darters and tropic birds used to be in this clade, but DNA and morphological studies indicate otherwise. “Recent research strongly suggests that the similarities between the Pelecaniformes as traditionally defined are the result of convergent evolution rather than common descent.” None of these have been tested yet. Most have a bare throat patch (gular patch), and the nostrils have evolved into dysfunctional slits, forcing them to breathe through their mouths.

Figure 5. Macronectes, the Southern giant petrel, in vivo.

According to Wiki
Procellariiformes include the petrel and albatross. These, in turn, are most closely related to penguins and loons. The LRT (Fig. 7) finds several taxa nest between penguins and loons, including dippers, kingfishers and hummingbirds, but we’ll save that for another day.

Figure 6. Coragyps, the black vulture, in vivo.

According to Wiki:
Accipitriformes include the New World vultures, the hawk, secretary bird and eagle, but not the falcon and owl. DNA nests falcons closer to parrots and sparrows. Accipitriformes are carnivorous with raptorial claws and a sharply hooked beak, but the same can be said of falcons and parrots. Here (Fig. 7) falcons, like Falco, nest with the terror birds and their extant relatives, not with parrots.

Well, this is embarrassing!
Yesterday’s post had roadrunners linked to herons. Everyone knows roadrunners are a type of cuckoo. Today, with the addition of Coccyzus, the cuckoo, that problem resolves itself. The LRT nests both cuckoos with the heron, Ardea (Fig. 7). This series appears to  document another example of serial phylogenetic miniaturization, with a smaller and smaller overall size coupled with shorter legs (neotony) and a return to the down curved rostrum found in the ratite ancestors of herons, like Rhynchotus.

Figure 8. Members of the cuckoo/heron clade along with a baby heron.

As We’ve seen before
DNA results do not match morphological results over larger phylogenetic distances. And the same appears to hold true for extant birds. I thought the birders had this all figured out, but apparently there is room for yet another hypothesis of relationships here. The LRT bird tree topology is, so far, staying pretty simple and logical.

Figure 9. Bird cladogram from 12/24/2021 with 950 more taxa in the LRT. Follow this latest data.

Nullius in verba

References
Brisson MJ 1760. Ornithologie, ou, Méthode contenant la division des oiseaux en ordres, sections, genres, especes & leurs variétés : a laquelle on a joint une description exacte de chaque espece, avec les citations des auteurs qui en ont traité, les noms quils leur ont donnés, ceux que leur ont donnés les différentes nations, & les noms vulgaires
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.
Latham J 1790.  Index Ornithologicus, Sive Systema Ornithologiae: Complectens Avium Divisionem In Classes, Ordines, Genera, Species, Ipsarumque Varietates (2 Volumes) (in Latin). London: Leigh & Sotheby.
LeMaout 1853. xx

wiki/Scopus
wiki/Threskiornis
wiki/Macronectes
wiki/Coragyps atratus

Roadrunner skull and surface features

These two images of the extant roadrunner
(Fig. 1; genus: Geococcyx) were so close to each other, they presented a great opportunity to match skull to surface features on one of our favorite birds.

Figure 1. GIF animation of Geococcyx skull matched to surface feathers. Distinct from other birds tested so far, the nares is far forward, apart from the antorbital fenestra.

A minimum of guesswork
and/or imagination was used in the creation of this image. Since the skull is a cast, sutures were ‘sutured’ to create a single object.

Geococcyx californum 
(Wagler 1831; up to 60 cm longl) the extant roadrunner is a small terrestrial heron and a basal neognath with a posteriorly rotated pedal digit 4, unrelated to parrots and toucans with a similar toe. Traditionally roadrunners are considrered part of the cuckoo family. When cuckoos get tested, they may also nest here. They have shorter hind legs.

Figure 2. Geococcyx the roadrunner skeleton. Note the crane-like proportions of this small land heron, probably a late-surviving Early Cretaceous member of the Euornithes.

And where does the roadrunner nest
in the large reptile tree (LRT 1087 taxa)? Between toothy Yanornis and toothless Ardea, the heron, all three at the base of the neognath birds, not surprisingly close to Sagittarius and Cariama, two other extant bird terrestrial predators with long hind limbs.

Nullius in verba

References
Wagner JG 1831. Einige Mitheilungen über Thiere Mexicos. Oken’s Isis 24:510–535.

wiki/Roadrunner