Dodos are flightless vultures, not pigeons.

This is what happens
when you add taxa to the mix. The large reptile tree (LRT, 1105 taxa) does what it does without tradition, taxon exclusion, bias or tribute to any overlord professor. Dodos and solitaires (Figs. 1, 3, 5-7) are not pigeons (Fig. 2) in the LRT (Fig. 8).  Given the present list of taxa dodos nest as flightless vultures, a relationship proposed over a hundred years ago by Owen (1846), but abandoned within two years by Strickland and Mehlville (1848) in favor of the earlier pigeon hypothesis by Reinhart (1842). And that has been gospel ever since.

Figure 1. The dodo, Raphus, compared to the New World vulture, Coragyps. These two nest together apart from the pigeons in figure 1. 

Figure 1. The dodo, Raphus, compared to the New World vulture, Coragyps. These two nest together apart from the pigeons in figure 1.

Current thinking (going back to Reinhart 1842)
(Wikipedia dodo page) nests the dodo (genus: Raphus, Fig. 1) with the nicobar pigeon (genus: Caloenas, Fig. 2). If you’ve ever wondered about this, I had the same giant question mark over my head, too. Here, finally, the dodo gets tested in a large gamut taxon list and another long-standing paradigm falls by the wayside.

Figure 1. Two pigeon skulls, Caloenas and Columba. These taxa nest together apart from vultures and the dodo, Raphus.

Figure 2. Two pigeon skulls, Caloenas and Columba. These taxa nest together apart from vultures (Fig. 8) and the dodo, Raphus (figure 2). Even the palate is different here. By convergence, and beside the point, these pigeons look more like the vulture, Coragyps, and less like the dodo, Raphus (Fig. 1).

The LRT 
nests the dodo, Raphus, with the New World vulture, Coragyps (Fig. 1). So, the dodo is a flightless vulture, not a pigeon. Both are derived from soaring seabirds like Macronectes. This addition of related taxa pushes the puffin, Fratercula, and the auk, Pinguinus, off to the side, as sisters to the Coragyps/Raphus clade. They also had a common ancestor among long-ranging sea birds,

Figure 4. Dodo (Raphus) skeletal data.

Figure 3. Dodo (Raphus) skeletal data.

According to Wikipedia:
“The dodo was variously declared a small ostrich, a rail, an albatross, or a vulture, by early scientists. In 1842, Danish zoologist Johannes Theodor Reinhardt proposed that dodos were ground pigeons, based on studies of a dodo skull he had discovered in the collection of the Natural History Museum of Denmark. This view was met with ridicule, but was later supported by English naturalists Hugh Edwin Strickland and Alexander Gordon Melville in their 1848 monograph The Dodo and Its Kindred, which attempted to separate myth from reality. After dissecting the preserved head and foot of the specimen at the Oxford University Museum and comparing it with the few remains then available of the extinct Rodrigues solitaire (Pezophaps solitaria) they concluded that the two were closely related. Strickland stated that although not identical, these birds shared many distinguishing features of the leg bones, otherwise known only in pigeons.”

“Strickland and Melville pointed to the very short keratinous portion of the beak, with its long, slender, naked basal part.”

A trait shared with Coragyps by convergence.

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

Figure 4. When vultures drift north and start finding fish attractive they evolve into auks and puffins.

“Other pigeons also have bare skin around their eyes, almost reaching their beak, as in dodos.”

As in Coragyps.

Figure 4. Raphus skull compared to Coragyps, a vulture, and Caloenas, a pigeon. White arrow points to nostril.

Figure 5. Raphus skull compared to Coragyps, a vulture, and Caloenas, a pigeon. White arrow points to nostril.

“The forehead was high in relation to the beak, and the nostril was located low on the middle of the beak and surrounded by skin, a combination of features shared only with pigeons.”

And vultures.

“The legs of the dodo were generally more similar to those of terrestrial pigeons than of other birds, both in their scales and in their skeletal features.”

I can’t comment on this with available data, other than to say the description is too general and many bird legs are similar in structure, inside and out.

“Depictions of the large crop hinted at a relationship with pigeons, in which this feature is more developed than in other birds.”

Also found in vultures.

“Pigeons generally have very small clutches, and the dodo is said to have laid a single egg.”

Coragyps lays from one to three eggs.

“Like pigeons, the dodo lacked the vomer and septum of the nostrils, and it shared details in the mandible, the zygomatic bone, the palate, and the hallux.”

You can see through both nares in Coragyps, too.  The rest of this description is too generalized to comment on, other than to say the LRT recovers more dodo traits shared with the vulture, Coragyps than with the pigeon, Colaenas.

“The dodo differed from other pigeons mainly in the small size of the wings and the large size of the beak in proportion to the rest of the cranium.”

Same with Coragyps.

Not too many skeletal traits
listed by these authors. The LRT employs only skeletal traits. Not sure why vultures were excluded from dodo cladograms. And if they were not excluded, one has to wonder why dodos did not nest with Old World vultures. The long list of synapomorphies easily overwhelms any list of pigeon traits.

Raphus cucullatus (Linnaeus 1758; recently extinct; 1m tall) is the flightless dodo, endemic to the island of Maritius, east of Madagascar. Traditionally the closest living relative is the Nicobar pigeon, Caloenas. Here Raphus nests with Pezohaps and Coragyps, a New World vuluture.

The dodo has a less famous flightless relative
more closely related to it than to Coragyps. Meet Pezophaps solitaire (Fig. 6), the flightless solitaire, from the island of Rodrigues, east of Madagascar. It was also recently discovered and recently decimated by humans.

Figure 5. Swan-sized, Pezophaps, the solitaire, is the closest dodo relative. It is likewise closer to New World vultures than to African pigeons.

Figure 6. Swan-sized, Pezophaps, the solitaire, is the closest dodo relative. It is likewise closer to New World vultures than to African pigeons.

Pezophaps solitaria (Gmelin 1789; recently extinct; 70cm tall) is the flightless solitaire, from the island of Rodrigues, east of Madagascar. Here it nests with the dodo, Raphus, and both are related to the New World vulture, Coragyps.

Figure 6. Pezophaps skull. This is a very robust skull for any bird or any theropod dinosaur. 

Figure 7. Pezophaps skull. This is a very robust skull for any bird or any theropod dinosaur.

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 Raphus, the dodo, the head and neck lack feathers.

Unlike the flightless ratites
these flightless birds kept a deep sternum, despite having tiny wings.

Figure 8. Subset of the LRT focusing on birds. Here Raphus, the dodo, nests with the New World vulture, Coragyps (in orange), not the pigeon, Caloenas (in green).

Figure 8. Subset of the LRT focusing on birds. Here Raphus, the dodo, nests with the New World vulture, Coragyps (in orange), not the pigeon, Caloenas (in green).

Oddly, if you’ve noticed…
dodos and solitaires both lived east of Madagascar, in the Old World. They are not closely related to Old World vultures like, Torgos. Rather, dodos and solitaires are related to New World vultures. How can that be? All hypotheses and speculations are welcome. IMHO we’re going to find a long-ranging, soaring seabird that was the last common ancestor of both dodos and New World vultures. Or we’ll find some New World vulture fossils in the Old World, out-competed or diseased out of existence, except on those tiny islands East of Madagascar.

Size matters, sometimes.
Both the dodo and the solitaire are closer in size to soaring sea birds and vultures than to pigeons. I think the historical confusion arose because, for reasons unknown, pigeons do indeed share several traits with New World vultures (including dodos) by convergence (Figs. 1, 2).

At this point, I look for further citations
Meuer et al. 2014 report in an abstract that only one complete dodo skeleton is known from a single individual. Others are chimeric combinations of incomplete specimens. They report, “Dodo cranial morphology is characterized by a lengthening and heightening of the maxilla, a concomitant antero-posterior compression of the cranium and a dorsally expanded frontal region. There is no ossified nasal septum, and both specimens lack an ossified vomer. Although the orbital region forms a large part of the cranium, it is reduced in size relative to extant Columbiformes. The occipital region is flat, wide and oriented vertically. The foramen magnum and the occipital condyle are located in a posterior position on the skull. This arrangement is similar to that of the closely related Solitaire of Rodriguez, Pezophaps solitaria, but differs from extant Columbiformes, including the dodo’s closest living relative, the Nicobar Pigeon (Caloenas nicobarica).

See how hey bought into the tradition without testing. Now look at all the differences these authors find between pigeons (columbids) and the dodo:

“In these columbids, the occipital region is more rounded and both the foramen magnum and occipital condyle are located ventrally. The fossa temporal is of the dodo is deep and narrow, and the quadratum is X-shaped. Both the mandible and cranium are only gently curved dorso-ventrally. The mandibular rami are high and narrow, and contain only a single mandibular foramen. The medial mandibular process is large and triangular, and together with articulatory processes on the basitemporal plate, argues for a secondary articulation of the mandible with the basitemporal plate. Our study of the Thirioux specimens highlights the dodo’s peculiar cranial morphology, which likely evolved in response to a more demanding and specialized lifestyle and feeding mechanism than previously appreciated.”

Shapiro et al. 2002 report, “The evolutionary history of the dodo is very poorly understood.” They phylogenetically tested the dodo only against Pezophaps and 35 pigeons and doves… and they used DNA.

Switek 2011 reports, “First-hand accounts of the birds agreed that they sported plumage that was black to grey in color,”

Hume 2006 reports, “More has been written about the dodo Raphus cucullatus of Mauritius than any other bird. Many conclusions based on the available data are problematic.”

Strckland and Mehlville 1848, credit Owen 1846 with relating the dodo to the raptorial birds and Reinhart 1842 for noting the pigeon affinities with the tubular naris (Fig. 5).

Naish 2014 reviewed a book by Parish 2012. “Today it’s well known and near-universally accepted that dodos and solitaires are pigeons, deeply nested within Columbidae. Historically, however, these birds have been considered related to, or members of, ratites, gamebirds, swans, penguins, vultures, waders, and rails. Parish reviews all of these, often fanciful, suggestions (and others), using the assorted family trees and other diagrams produced by the respective authors.”

I did not have access to this book prior to posting this blog. But I have seen the evidence and I understand and attempted to show that dodos and solitaires nest with New World vultures rather than pigeons in the LRT. As readers know, I am not adverse to testing long-held paradigms and purported clades. This is just one more heresy that will someday be embraced or invalidated by others who run similar tests.

References
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.
Hume J 2006. The history of the Dodo Raphus cucullatus and the penguin of Mauritius Historical Biology, 18 (2), 65-89 DOI: 10.1080/08912960600639400
Hume JP, Cheke AS and McOran-Campbell A 2009. How Owen ‘stole’ the Dodo: academic revelry and disputed rights to a newly discovered sub fossil in nineteenth century Mauritius. Historical Biology 21:33–49. online pdf
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.
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Meijer H, Claessens L, Hume J, Jando A and Rijsdijk K 2014. A re-evaluation of cranial anatomy of the dodo (Raphus cucullatus) based on two previously undescribed specimens. Journal of Vertebrate Paleontology abstracts, 2014:186.
Naish D 2014. A review of ‘The Dodo and the Solitaire: A natural history’. Journal of Vertebrate Paleontology 34(2):489-490.
Owen R 1846. Observations on the Dodo. Proceedings of the Zoollogical Society of London 1846:51–53.
Parish JC 2012. The dodo and the solitaire: a natural history. Indiana University Press, Bloomington, IN 406 pp.
Reinhardt JT 1842. In: Kroyer H, editor. Noiere Oplysning om det I Kiobenhavn fundne Drontehoved. Vol. 4. København (Denmark): Naturhistorisk Tidskrift. p. 71.
Shapiro B et al. (7 co-authors) 2002. Flight of the Dodo. Science Brevia. 295:5560:1683.
Strickland HE and Melville AG 1848. The dodo and its kindred. London: Reeve, Benham & Reeve. p 141.
Switek B 2011. The dodo is dead, long live the dodo! Wired.com

wiki/Coragyps atratus
wiki/Dodo
wiki/Rodrigues_solitaire

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Swifts and Swallows

They look alike on the outside…
but everyone knows they are different inside. That’s been known for a long time. Swifts and swallows also nest far apart from one another in the large reptile tree (LRT, 1099 taxa).

Figure 1. Subset of the LRT focusing on extant birds and their kin. Here swifts (Apus) nest apart from swallows (Hirundo).

Figure 1. Subset of the LRT focusing on extant birds and their kin. Here swifts (Apus) nest apart from swallows (Hirundo).

Rampant convergence in the bird subset of the reptile cladogram.
We’ve seen this before, recently, in penguins + murres versus auks + puffins.

Figure 2. Skull of Hirundo, the barn swallow.

Figure 2. Skull of Hirundo, the barn swallow, closer to wrens and dippers.

When you get inside their heads
Hirundo, the barn swallow (Fig. 2) and Apus, the common swift, are readily different.

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

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

Apus apus (Linneaus 1758) is the common swift. Long thought to be related to hummingbirds like Archilochus, Apus nests here with falcons and owls.

Hirundo rustica (Linneaus 1758) is the extant barn swallow. Swallows are not related to, but convergent with swifts like Apus. Here they nest with wrens and dippers, but with a shorter rostrum/larger orbit and longer wing feathers.

These direct comparisons
are teaching me something I never thought of learning, but now find fascinating.

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.

Woodpeckers nest with swifts and wrens + dippers

… and not far from murres and penguins.
This clade has evolved into a wide gamut of niches with similar skulls that share many traits.

Figure 1. Skull of Melanerpes, a woodpecker. Note the robust mandible and rostrum. The tongue bones extend over the back of the skull anchored over the nares.

Figure 1. Skull of Melanerpes, a woodpecker. Note the robust mandible and rostrum. The tongue bones extend over the back of the skull anchored over the nares.

Melanerpes aurifrons (Wagler 1829) is the extant golden-fronted woodpecker. Tongue bones extend over the back of the skull to anchor over the nares. The skull is more robust than in related forms, a trait that makes wood pecking possible. Skull and skeleton images from Digimorph.org and used with permission.

Figure 2. Melanerpes, a woodpecker in the LRT, nests with dippers, swifts and wrens.

Figure 2. Melanerpes, a woodpecker in the LRT, nests with dippers, swifts and wrens.

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

 

Stilts and hummingbirds: closer than we thought!

Sure,
one has longer legs and probes mud flats (Figs. 1, 2) while the other has tiny legs and probes flowers (Fig. 2), but now they nest close to one another in the large reptile tree (LRT, 1100 taxa, still fully resolved).

Figure 1. Photo and skeletal drawing of Himantopus, the black-necked stilt.

Figure 1. Photo and skeletal drawing of Himantopus, the black-necked stilt. 2 frames GIF movie

 

Himantopus mexicanus (Muller 1776) is the extant black-winged stilt. It is a wading bird with extraordinary long legs. Like the hummingbird, the beak is long and gracile. They eat buried invertebrates in mudflats near water.

Figure 2. Skull of Himantopus mexicanus, the black-winged stilt. Here its hummingbird affinities are apparent.

Figure 2. Skull of Himantopus mexicanus, the black-winged stilt. Here its hummingbird affinities are apparent.

And for comparison,
let’s also look at a hummingbird (Archilochus) skull (Fig. 3), sharing a long list of traits, including a long thin rostrum with a long narrow naris, large cranium and slender, arched prefrontals.

Figure 3. Hummingbird skull for comparison to the stilt in figure 2. Image courtesy of Digimorph.org and used with permission.

Figure 3. Hummingbird skull for comparison to the stilt in figure 2. Image courtesy of Digimorph.org and used with permission.

So roughly the order is:
crows > terns > stilts > hummingbirds. As in so many other novel relationships recovered by the LRT, a bit of googling indicates this hypothetical relationship has not been indicated before. And fantasy confession time: the black-necked stilt is my favorite bird. Knowing that it’s related to hummingbirds makes it all the more intriguing. 

References
Müller OF 1776. Zoologiae Danicae prodromus: seu Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Hafniae, Typiis Hallageriis. 1-274.

A flightless, swimming, polar vulture: the great auk

And the puffin
(Fig. 4) is a swimming, polar vulture, too. It’s smaller than the auk (Fig. 1)and it can still fly.

Figure 1. The great auk (genus Pinguinus) is a flightless vulture convergent with penguins.

Figure 1. The great auk (genus Pinguinus) is a flightless vulture convergent with penguins.

The great auk
(genus Pinguinus) is a recently extinct fairly large, penguin-like bird of the North Atlantic. In the large reptile tree (LRT, 1096 taxa) it nests with Coragyps, the extant black vulture.

Figure 2. Pinguinus the great auk skull.

Figure 2. Pinguinus the great auk skull.

Pinguinus impennis (Linneaus 1758; standing 80cm in height) is the recently extinct great auk. Here it nests with the puffin and vultures. Convergent with penguins like AptenodytesPinguinus was flightless, but a good swimmer underwater.

It’s worthwhile to keep
the skeleton of the vulture Coragyps (Fig. 3) in mind when comparing skeletons.

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

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

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, the head and neck lack feathers.

Everyone knows
Puffins (genus Fratercula) are just small auks that can still fly, even with those short whirring wings. The earlier issue was with the next level of relationships, which are traditionally relegated to suprageneric taxa

Figure 4. The skull of the puffin (genus: Fratercula) with and without the keratin beak.

Figure 4. The skull of the puffin (genus: Fratercula) with and without the keratin beak.

Fratercula arctica (Linneaus 1758; standing 20cm in height) is the extant Atlantic or common puffin. Here it nests with the great auk and vultures, hamerkops (genus: Scopus) and gulls. Both genders have a tall, colorful beak.

According to Wikipedia
birds of prey (Telluraves) includes the taxa shown below (Fig. 5).

Figure 5. Bird relationships according to Wikipedia with comments in red.

Figure 5. Bird relationships according to Wikipedia with comments in red. I never thought the birds would be this messed up. Maybe starting with a smaller taxon list was a good idea this time. Kept things simple.

Unfortunately
The LRT (Fig. 6) does not support several of the traditional bird relationships shown on the Wiki page (Fig. 5), and a good look at the relationships will suggest the gaps between sister clades are just too big. Plus, relying on suprageneric taxa always causes problems and never pinpoints actual sister genera. Test these relationships yourself, as I have, and let me know if you recover anything different.

Figure 5. Subset of the LRT focusing on auk and puffin relatives.

Figure 6. Subset of the LRT focusing on auk and puffin relatives.

I’ve been binging on Burning Man Festival videos
on YouTube and in the spirit of their cashless, gift-giving temporary society, this blog featuring the results of my studies is my gift to you.

References
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.
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/Coragyps atratus
wiki/Atlantic_puffin
wiki/Great_auk

 

Evolution of the penguin, as told by ancestral taxa

I’m going to let this image tell the story. 

Figure 1. Penguin evolution as told by ancestral taxa back to the Late Jurassic. I could have gone further, but for that see www.ReptileEvolution.com

Figure 1. Penguin evolution as told by ancestral taxa back to the Late Jurassic. I could have gone further, but for that see http://www.ReptileEvolution.com. One of the ways you can tell the LRT is more or less correct is by showing a gradual accumulation of traits for any derived taxon, in this case the penguin. Note the phylogenetic miniaturization prior to the new body plan.

What an image like this does
is to satisfy the eye that all the data points in the large reptile tree (LRT, 1094 taxa) really do add up to a gradual accumulation of traits for any derived taxon, in this case the penguin and all taxa in between. And it’s really going to take the entire Cretaceous and Tertiary to go from Alpha to Omega here. Here (Fig. 1) there are no great morphological gaps. The size changes. The beak length changes. The neck and leg lengths change, but all of this does so gradually, radiating off in different directions every several dozen/thousand generations or whenever the niche changes.

If you need a few more details,
request the .nex file here or run through the referenced taxa, starting with Aptenodytes, the emperor penguin and go backwards… or forwards from Eosinopteryx.

Giant flightless birds: Worthy et al. 2017

Worthy et al. nest several giant flightless birds
with chickens and ducks. In the large reptile tree (LRT, 1094 taxa, subset Fig. 1) none of these giants nest with chickens and ducks. Furthermore, chickens (Gallus) and ducks (Anas) don’t nest with each other in the LRT. They don’t look like each other, so their separation makes sense.

Figure 1. Subset of the LRT focusing on birds. Here giant and tiny birds are highlighted. None nest with chickens or ducks, which don't nest with each other.

Figure 1. Subset of the LRT focusing on birds. Here giant and tiny birds are highlighted. None nest with chickens or ducks, which don’t nest with each other.

From the Worthy et al. abstract:
“The extinct dromornithids, gastornithids and phorusrhacids are among the most spectacular birds to have ever lived, with some giants exceeding 500 kg. The affinities and evolution of these and other related extinct birds remain contentious, with previous phylogenetic analyses being affected by widespread convergence and limited taxon sampling. We recognize a robust new clade (Gastornithiformes) for the giant flightless Dromornithidae (Australia) and Gastornithidae (Eurasia, North America). This clade exhibits parallels to ratite palaeognaths in that flight presumably was lost and giant size attained multiple times South America’s largest bird, Brontornis, is not a galloansere, but a member of Neoaves related to Cariamiformes.”

Figure 2. Brontornis parts compared to Gastornis, a close match both in size and morphology.

Figure 2. Brontornis parts compared to Gastornis, a close match both in size and morphology.

Brontornis bits ‘n pieces
Apparently Brontornis is known from a big metatarsus and a big fused dentary (lower beak), perhaps not enough to nest it in the LRT, but South American terror birds (Cariamiformes according to Worthy et al., a clade not supported here, Fig. 1) have a very narrow beak, whereas Brontornis does not. Here (Fig. 2) the shape and size of Brontornis is quite similar to the giant parrot, Gastornis (formerly Diatryma).

While writing this paragraph
I was drawn to the Wiki Brontornis page, which reports (after describing Brontornis as a giant, flightless terror bird), “Recent work (Agnolin 2007, Buffetaut 2014) ]has cast doubt on the hypothesis that Brontornis is a phorusrhacid. Brontornis may actually represent an anseriform” (traditionally ducks, geese and screamers, but the LRT nests screamers apart). Not sure why the Brontornis/wiki author could not make a scientific statement with more confidence. After all, there is only one answer. The other is false.

From the Worthy et al. introductiion
“Landfowl (Galliformes) and waterfowl (Anseriformes) form a diverse and important clade (Galloanseres) that is sister to Neoaves (all other extant non-palaeognath birds).” This is what Prum 2015 recovered using DNA, but it is not what the LRT recovered (Figs. 1–4) using morphology and extinct taxa.

Worthy et al. also report, 
“These giant flightless Galloanseres show striking morphological convergence with flightless palaeognaths (ratites), especially the large extinct Aepyornithidae (elephant birds; Madagascar) and Dinornithiformes (moa; New Zealand).” The LRT recovers elephant birds (Aepyornis) with corn crakes (Crex) and moas (Dinornis) between toucans (Pteroglossus) and parrots (Ara, Figs. 1, 3-5). So Worthy et al. appear to be basing their hypotheses on very shaky ground.

While we’re on the subject of birds
here are a few clade divisions recovered by the LRT.

Figure 2. Bird clades, basal divisions.

Figure 3. Bird clades, basal divisions. Where are all the Late Cretaceous birds? They are waiting to be discovered.

Figure 2. Bird predators and omnivores compared to plant/nectar eaters.

Figure 4. Bird predators and omnivores compared to plant/nectar eaters.

Figure 4. Most basal birds have a premaxilla about the length of the maxilla. That changes in these two basal clades.

Figure 5. Most basal birds have a premaxilla about the length of the maxilla. That changes in these two basal clades. I know I’m pulling a Larry Martin here, but after the phylogenetic analysis, not before. This trait stood out as a readily visible major division at a node that has remained difficult to establish for prior analyses.

The basal radiation of extant birds
has been clouded in mystery in prior studies. Here, with fewer taxa (Figs 1-5), the radiation is quite clear and it probably occurred deep into the Early Cretaceous with a large gap sprinkled with taxa until the Tertiary and then greatly expanded with living taxa.

References
Agnolin F 2007. Brontornis burmeisteri Moreno & Mercerat, un Anseriformes (Aves) gigante del Mioceno Medio de Patagonia, Argentina. Revista del Museo Argentino de Ciencias Naturales, n.s.9, 15-25.
Buffetaut E 2014. Tertiary ground birds from Patagonia (Argentina) in the Tournouër collection of the Muséum National d’Histoire Naturelle, Paris. Bulletin de la Société Géologique de France. 185(3):207–214.
Worthy TH, Degrange FJ, Handley WD and Lee MSY 2017. The evolution of giant flightless birds and novel phylogenetic relationships for extinct fowl (Aves, Galloanseres). Royal Society Open Science 4: 170975. http://dx.doi.org/10.1098/rsos.170975

tetrapodzoology/terror-birds