Euparaves (5)

From the previous post:
The most parsimonious alternative is that the basalmost Euparaves were feathered, long-bony-tailed, flying primitive birds. And the ground-dwelling oviraptors and alvarezsaurids that came later, were secondarily flightless.

Is this a problem for the dinosaur to bird theory?

Indeed it is.
It is a problem because the oviraptors and alvarezsaurids are considered to be outgroups (intermediates) between dinosaurs and Euparaves. But if oviraptors and alvarezsaurids are descended from flying Euparaves, then there is no lineage from dinosaurs to Euparaves.

Basal Euparaves were four-winged. The hindlimbs were feathered and acted as additional wings (hindwings). They contributed to lift and control.
This is important because they contributed to the ability of the basal Euparaves to fly.

https://en.wikipedia.org/wiki/Tetrapterygidae

Tetrapterygidae (meaning "four-wings") is a group of four-winged dinosaurs proposed by Sankar Chatterjee in the second edition of his book The Rise of Birds: 225 Million Years of Evolution, where he included Microraptor, Xiaotingia, Aurornis, and Anchiornis.[1] The group was named after the characteristically long flight feathers on the legs of all included species, as well as the theory that the evolution of bird flight may have gone through a four-winged (or "tetrapteryx") stage, first proposed by naturalist William Beebe in 1915.[2] Chatterjee suggested that all dinosaurs with four wings formed a natural group exclusive of other paravians, and that this family was the sister taxon to the group Avialae, although most phylogenetic analyses have placed the animals of his Tetrapterygidae elsewhere in Paraves, such as Xiaotingia, Aurornis, andAnchiornis being placed in Avialae.[3]

https://www.google.ca/url?sa=t&rct=j&q=&esrc=s&source=web&cd=4&cad=rja&uact=8&ved=0ahUKEwjNgIW0-o3KAhXBKh4KHZZZBVwQFgg1MAM&url=http%3A%2F%2Fwww.asbweb.org%2Fconferences%2F2012%2Fabstracts%2F529.pdf&usg=AFQjCNGssPFCAJ3XMsXRzpb6B37h1wR_Mg&sig2=_L19RDPlM7wYTfgSd8Flvg&bvm=bv.110151844,d.dmo
HINDWING FUNCTION IN FOUR-WINGED FEATHERED DINOSAURS

The evolution of powered flight in birds remains a contentious issue in vertebrate paleontology. The diminutive predatory dinosaur Microraptor gui preserves evidence of extensive, lift-generating feathers on each manus and forearm, but also preserves evidence of lift-generating feathers associated with the hindlimbs, effectively forming a pair of “hindwings”.

Euparaves (4)

Notice that in the study below, Pedopenna is included with Epidexipteryx and Oviraptors within basal Euparaves. Other flying primitive birds such as Eosinopteryx, Aurornis, Xiaotingia and Anchiornis etc should also be included (with Pedopenna, Epidexipteryx and Oviraptors) as basal Euparaves. However, in this cladogram they are shown as troodontids. (They have been classified differently in various studies).

Let's now consider whether the basalmost Euparaves were ground-dwelling or whether they were flying in the trees. Pedopenna, Epidexipteryx and the other flying  primitive birds such as Eosinopteryx, Aurornis, Xiaotingia and Anchiornis etc were all flying in the trees.
So if the basalmost Euparaves were ground-dwelling, then they must all have BECOME flying primitive birds AFTER splitting from the dinosaur line. Not only that, but they must all have developed like that convergently, to become like the other flying birds that came later. That is not parsimonious. 

The most parsimonious alternative is that the basalmost Euparaves were feathered, long-bony-tailed, flying primitive birds. And the ground-dwelling oviraptors and alvarezsaurids that came later, were secondarily flightless.


For reference:
https://en.wikipedia.org/wiki/Pedopenna

Pedopenna was originally classified as a paravian, the group of maniraptoran dinosaurs that includes both deinonychosaurs and avialans (the lineage including modern birds), but some scientists have classified it as a true avialan more closely related to modern birds than to deinonychosaurs.[3]

Powered flight, flapping flight, gliding (2)

Basal Euparaves could fly by flapping their wings, but they lacked the muscle structure to take off from the ground. In other words, they could fly but they were not capable of "powered flight".
Some of the literature gives the impression that they could only glide and parachute. 
But they could fly. They just were not capable of "powered flight" (they could not take off from the ground).
And basal Euparaves could fly even though they did not have asymmetric feathers. 

This is all very important because the dinosaur to bird theory is built on the idea that basal Euparaves could not fly. 

Let's take a break

Parachuting!

Powered flight, flapping flight, gliding

Basal Euparaves could fly by flapping their wings, but they could not take off from the ground. In other words, they capable of "powered flight" but not able to takeoff from the ground.

https://en.wikipedia.org/wiki/Paraves

.... the work of Xu et al. (2003), (2005) and Hu et al. (2009) provide examples of basal and early paravians with four wings,[11][12][13] adapted to an arboreal lifestyle who would only lose their hindwings when some adapted to a life on the ground and when avialans evolved powered flight.[14] Newer research also indicates that gliding, flapping and parachuting was another ancestral trait of Paraves, while true powered flight only evolved once, in the lineage leading to modern birds.[15]

http://www.bris.ac.uk/news/2014/february/origin-of-birds.html

"Out of all these flappers and gliders, only the birds seem to have been capable of powered flight," said co-author Mike Benton, Professor of Vertebrate Palaeontology at Bristol.

http://jeb.biologists.org/content/200/23/2987.full.pdf

The lack of a morphologically derived SC [supracoracoideus] in Late Jurassic and Early Cretaceous birds precluded a high velocity recovery stroke which undoubtedly limited powered flight in these forms. Subsequent evolution of the derived SC capable of imparting a large rotational force to the humerus about its longitudinal axis was an important step in the evolution of the wing upstroke and in the ability to supinate (circumflex) the manus in early upstroke, a movement fundamental to reducing air resistance during the recovery stroke.

https://en.wikipedia.org/wiki/Euavialae

Euavialae (meaning "true winged birds") is a group of birds which includes all avialan species more closely related to modern birds, than to the primitive, long-tailed birds Archaeopteryx and Jeholornis.[1]

http://books.google.ca/books?id=8QRKV7eSqmIC&pg=PA136&lpg=PA136&dq=dorsal+elevator+feduccia&source=bl&ots=fqR1hQfECg&sig=NAYAvTNh7USVZlO-dFmaLcJsDPQ&hl=en&sa=X&ei=UUUYVIndD8-zyATm3oDgDg&ved=0CCQQ6AEwAQ#v=onepage&q=dorsal%20elevator%20feduccia&f=false

The dorsal elevators, principally the deltoideus major, can effect the recovery stroke by themselves, as they did in Archaeopteryx. The German anatomist Maxheinz J. Sy proved this when he cut the tendons of the supracoracoideus in living crows and pigeons (1936). Sy found that pigeons were capable of normal, sustained flight; the only capacity they lost was the ability to take off from level ground.

Concerning flapping flight, also see: 
http://pterosaurnet.blogspot.ca/2015/07/primitive-birds-flying.html
http://pterosaurnet.blogspot.ca/2013/02/flight-stroke.html
http://pterosaurnet.blogspot.ca/2014/09/flight-strokes-compared.html
http://pterosaurnet.blogspot.ca/2010/05/keeled-breastbone.html
http://pterosaurnet.blogspot.ca/2014/09/flight-without-supracoracoideus.html

Euparaves (3)

Notice that Pedopenna is included with Epidexipteryx as a basal Euparaves. Those are just two of the flying basal Euparaves*.
http://www.sciencedirect.com/science/article/pii/S0960982214010471 (Brusatte et al, 2014)

* Here are other flying basal Euparaves:
Scansoriopterygids (Temporal range: Late Jurassic, 165–156 Ma)
Anchiornis (Temporal range: Late Jurassic, 161–160.5 Ma)
Aurornis (Temporal range: Late Jurassic, 160 Ma)
Xiaotingia (Temporal range: Late Jurassic, 160 Ma)
Pedopenna (Temporal range: Middle or Late Jurassic, 164 Ma)
Eosinopteryx? (Temporal range: Late Jurassic, 160 Ma)

Pterosaur and Basal Euparaves

PTEROSAUR
Pterorhynchus (Rhamphorhynchidae)

BASAL EUPARAVES
Scansoriopteryx (Scansoriopterygidae)

Note that in cladistics terms, basal Euparaves did not evolve from Pterorhynchus, but rather they share a common ancestor.
In non-cladistic terms, we could say that basal Euparaves evolved from a creature like Pterorhynchus. We can see that that is credible, from the drawings above.


For reference:
https://www.google.ca/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&uact=8&ved=0CCAQFjAAahUKEwi1-N7DrJzHAhUJkh4KHRtPAWA&url=http%3A%2F%2Fwww.ivpp.cas.cn%2Fcbw%2Fgjzdwxb%2Fxbwzxz%2F201401%2FP020140121386966325113.pdf&ei=A3bHVfXqAYmkepuehYAG&usg=AFQjCNH1PpVWciHk2GxOR_QxipkA_gqu5Q&sig2=MEwOmtQjh_PUu6OAxg9RCg&bvm=bv.99804247,d.dmo (O'Connor and Sullivan2014)

Fig. 7 Tails and pelves of derived maniraptoran theropods (modified from Persons et al., in press)
A. Anchiornis (Deinonychosauria: Troodontidae?); B. Archaeopteryx (Aves); C. Jeholornis (Aves); D. Confuciusornis (Aves); E. Gallus (Aves); F. Caudipteryx (Oviraptorosauria); G. Khaan (Oviraptorosauria); H. Epidendrosaurus (Scansoriopterygidae); I. Zhongornis (incertae sedis); J. Epidexipteryx (Scansoriopterygidae) Color key: blue short proximal caudal vertebrae; green transitional caudals; yellow elongate distal caudals; orange partially or fully fused terminal caudals (pygostyle)

Euparaves (2)

This is a cladogram from an earlier post.
It shows the placement of Euparaves, but the cladogram is still not completely correct since Euparaves is not actually connected to dinosaurs.

Euparaves (node-based) definition:
"The most recent common ancestor of Epidendrosaurus ninchengensis (Scansoriopterygidae) and Passer domesticus (the house sparrow), and all descendants thereof".

Here is the stem-based definition of Euparaves:
"The most inclusive clade containing Passer domesticus (Linnaeus 1758) but not Pterorhynchus wellnhoferi."

Here are the apomorphies of Euparaves:
possessed remiges and rectrices, that is, enlarged, stiff-shafted, closed-vaned (= barbules bearing hooked distal pennulae), pennaceous feathers arising from the distal forelimbs and tail
AND
possessed feathered wings used in flapping flight

Concerning flapping flight, also see: 
http://pterosaurnet.blogspot.ca/2015/07/primitive-birds-flying.html
http://pterosaurnet.blogspot.ca/2013/02/flight-stroke.html
http://pterosaurnet.blogspot.ca/2014/09/flight-strokes-compared.html
http://pterosaurnet.blogspot.ca/2010/05/keeled-breastbone.html
http://pterosaurnet.blogspot.ca/2014/09/flight-without-supracoracoideus.html

Pyramid reduction hypothesis

The pyramid reduction hypothesis is a credible explanation of the origin of bird fingers. It is "developmentally plausible, and is also consistent with the phalangeal reduction pattern seen in basal birds."
It is consistent with a pterosaur to bird theory.
It is inconsistent with a dinosaur to bird theory.

http://www.ivpp.ac.cn/qt/papers/201403/P020140314320497349455.pdf
http://aracnologia.macn.gov.ar/st/biblio/Xu%20and%20Mackem%202013%20Tracing%20the%20Evolution%20of%20Avian%20Wing%20Digits.pdf

(Xu and Mackem 2013)
Good description of pyramid reduction hypothesis even though Xu and Mackem do not agree with it. 

The ‘pyramid reduction hypothesis’ assumes II-III-IV identities
for neornithine manual digits and postulates the existence
of a conservative five-digit pattern with a gradual,
bilateral reduction of phalanges and metacarpals in avian
evolution [9]. One proposed mechanism postulates that an
elevation in peripheral BMPs, signaling factors that modulate
cell survival and proliferation [60,61], drove bilateral medial
and lateral digital reduction [9]. This hypothesis is developmentally
plausible, and is also consistent with the phalangeal
reduction pattern seen in basal birds [9,23]. However, it predicts
that the direct avian ancestor had a five-fingered hand
with dominant digits II, III, and IV [9] which is inconsistent
with the digital reduction data from basal theropods (e.g.,
all known basal theropods, including ceratosaurs, have a
vestigial digit IV) [5,62–64].

In fact, the pyramid reduction hypothesis implies that either birds are not descended from theropod dinosaurs, or that some as yet to be discovered
basal theropods were five-fingered with dominant digits II, III, and IV.

And concerning the alternative homeotic shift (frameshift):

Also, it is difficult to identify plausible selective pressures that would drive this type of homeotic shift, considering that the post-frameshift adult hand would be morphologically identical to the pre-frameshift condition [69].

The frame shift and lateral shift hypotheses adopt different criteria for assessing primary homologies: the frame shift hypothesis relies on morphological information and selective spatial expression patterns of developmental genes, whereas the lateral shift hypothesis emphasizes positional information. However, the two hypotheses are similar, as both require genes that regulate digit pattern to shift their expression domains for the developing hand and the re-emergence of a fully functional digit in position 4, but differ in proposed mechanism of homeosis. 

Xu and Mackem note that the Pyramid Reduction Hypothesis "is inconsistent with the digital reduction data from basal theropods". But it is only inconsistent if birds evolved from dinosaurs. If birds did not evolve from dinosaurs, then there is no inconsistency. 
In fact, as Xu and Mackem acknowledge, one possible implication of the Pyramid Reduction Hypothesis is that "birds are not descended from theropod dinosaurs".

And here is another point:
Xu and Machem conclude that for the Pyramid Reduction Hypothesis to be correct:
"the direct avian ancestor had a five-fingered hand with dominant digits II, III, and IV".
They have overlooked the possibility that the ancestor had digits I-II-III-IV and lost digit I in the transition to primitive bird. Which is what I suggest occurred in the transition from pterosaur to primitive bird.

https://www.academia.edu/11078361/Pentadactyl_pattern_of_the_avian_wing_autopodium_and_pyramid_reduction_hypothesis

ABSTRACT
We report herein that a pentadactyl developmental pattern is evident in early wing morphogenesis of Gallus (chicken) and Struthio (ostrich). Five avascular zones (spatially predestined locations of contiguous metacarpal and phalangeal aggregation) and four interdigital vascular spaces are established by the regression patterns of autopodial vasculature. Transient vestiges of the first and fifth metacarpals are confirmed histologically and histochemically. They lie within the preaxial-most and postaxial-most avascular zones, respectively. These observations reveal conservative patterning of the avian hand and corroborate a II-III-IV metacarpal interpretation, argue for II-III-IV identity of ossified digits in birds, and favour a simple reduction rather than a homeotic shift in terms of the phenotype expressed by Hox genes in the phylogeny of the avian manus. We suggest that gradual, bilateral reduction of phalanges and metacarpals, via apoptosis mediated by BMP, occurred during the evolution of birds (Pyramid Reduction Hypothesis). This is congruent with the establishment of a central wing axis that became co-opted for coordinated movements. On the basis of evidence presented here, the direct avian ancestor is predicted to have been five-fingered with dominant digits (+ metacarpals) as follow: II, III, IV.

https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3986427/
Frame shift  and the rabbit

SUMMARY:
For years, people have been striving to reconcile the I-II-III digits of dinosaurs with the II-III-IV digits of birds.
Why?
Because they believe that birds evolved from dinosaurs.
Why do they believe that?
Because the cladistic analyses seem to support that idea.
How so?
Because they show oviraptors and alvarezsaurs as intermediates between dinosaurs and primitive birds (Paraves).
BUT recent studies show that oviraptors and alvarezsaurs are WITHIN Paraves.
So there is no cladistic analysis evidence to support the dino to bird theory.
Therefore there is no reason to strive to reconcile the I-II-III digits of dinosaurs with the II-III-IV digits of birds.
Which is good, because there is no credible way to reconcile them in the first place.


For reference:
https://www.researchgate.net/publication/260714879_Reply_to_Limusaurus_and_bird_digit_identity
Reply to “Limusaurus and bird digit identity”
Xing Xu1, James Clark2, Jonah Choiniere2, David Hone1 & Corwin Sullivan1

Morphological data from extinct theropods, even without considering Limusaurus and ceratosaurs, clearly contains two contradictory signals for the identification of tetanuran manual digits. Thus, neither our hypothesis nor the frameshift hypothesis is able to avoid a substantial number of homoplasies.

Oviraptors and Alvarezsaurids within Euparaves

In the previous post, we saw analyses from 2009, 2011, 2013, 2014 and 2015 that show oviraptors and/or alvarezsaurids to be within Euparaves. (In other words, within the clade based on the common ancestor of Epidendrosaurus and Gallus).

What are the basalmost taxa within Euparaves? They are the Scansoriopterygidae, Anchiornis, Aurornis, Xiaotingia and Pedopenna. They are flying, arboreal, primitive birds.

On the other hand, Oviraptors and Alvarezsaurids were ground-based. They were secondarily flightless, as some previous analyses have found.

What is the significance of this? It means that Oviraptors and Alvarezsaurids are not intermediate (not outgroups) between dinosaurs and flying primitive birds.
That is like having the foundation of the dino to bird theory kicked out. There is no connection between dinosaurs and primitive birds.


As a sidenote, keep in mind also that Eudromaeosaurids are secondarily flightless, primitive birds within Euparaves. They are not intermediate (not outgroups) between dinosaurs and flying primitive birds.

Different placements of oviraptors and alvarezsaurids

Here are cladograms with oviraptors and/or alvarezsaurids within Euparaves.


https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4476167/ (Cau et al 2015)

Oviraptors within Euparaves.
Updated dataset of Brusatte et al. (2014).

http://www.nature.com/articles/nature08124.epdf?referrer_access_token=1LIOYM249T2ALXmHhUVXQtRgN0jAjWel9jnR3ZoTv0NAxxXDTxDgb7tt7vNCs5i7CDx_p1E8pIL0dPMGIw0CIZ1LRnUZIDT1a3FIDY_UW4FRwpODRDVwWg-KbK448VK63yIXiGAa_H8fA42yVK8TsNhr_ASjWKKTbM-PJCMVzpKKElR4FEstewHl9DZGaHr9&tracking_referrer=www.nature.com (2009)
Cladogram based on a subset of taxa from Xu et al 2009 study:
Both Alvarezsaurids and Oviraptors within Euparaves.

For details see here.

Cladogram based on all data except Epidexipteryx from Xu et al 2009 study.
Both Alvarezsaurids and Oviraptors within Euparaves.

http://www.nature.com/articles/nature08124.epdf?referrer_access_token=1LIOYM249T2ALXmHhUVXQtRgN0jAjWel9jnR3ZoTv0NAxxXDTxDgb7tt7vNCs5i7CDx_p1E8pIL0dPMGIw0CIZ1LRnUZIDT1a3FIDY_UW4FRwpODRDVwWg-KbK448VK63yIXiGAa_H8fA42yVK8TsNhr_ASjWKKTbM-PJCMVzpKKElR4FEstewHl9DZGaHr9&tracking_referrer=www.nature.com (2009)
Figure S7 of the Xu et al 2009 study using ALL taxa:
Alvarezsaurids within Euparaves.

http://www.scielo.br/scielo.php?script=sci_arttext&pid=S0001-37652011000100008&lng=en&nrm=iso&tlng=en Angolin and Novas (2011)
http://www.scielo.br/img/revistas/aabc/v83n1/a08fig01m.jpg  Figure 1(B)
Alvarezsaurids within Euparaves

From Agnolín and Novas  (2013):
https://en.wikipedia.org/wiki/Scansoriopterygidae#Classification

Agnolín and Novas (2013) recovered scansoriopterygids as non-paravian maniraptorans and the sister group to Oviraptorosauria.^[11]

http://www.sciencedirect.com/science/article/pii/S0960982214010471 (Brusatte et al 2014)
Oviraptors within Euparaves. (Also see Figure S2 for more details.)
Notice that Pedopenna is included with Epidexipteryx within Euparaves.

https://www.google.ca/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&uact=8&ved=0CCAQFjAAahUKEwi1-N7DrJzHAhUJkh4KHRtPAWA&url=http%3A%2F%2Fwww.ivpp.cas.cn%2Fcbw%2Fgjzdwxb%2Fxbwzxz%2F201401%2FP020140121386966325113.pdf&ei=A3bHVfXqAYmkepuehYAG&usg=AFQjCNH1PpVWciHk2GxOR_QxipkA_gqu5Q&sig2=MEwOmtQjh_PUu6OAxg9RCg&bvm=bv.99804247,d.dmo (O'Connor and Sullivan 2014)
Zhongornis and Oviraptors within Euparaves.

Euparaves

In order to talk about the origin of primitive birds it is necessary to introduce a new clade name, Euparaves.
Euparaves (node-based) definition:
"The most recent common ancestor of Epidendrosaurus ninchengensis (Scansoriopterygidae) and Passer domesticus (the house sparrow), and all descendants thereof".

Here is the stem-based definition of Euparaves:
"The most inclusive clade containing Passer domesticus (Linnaeus 1758) but not Pterorhynchus wellnhoferi."

Here is the apomorphy-based definition of Euparaves:
possessed remiges and rectrices, that is, enlarged, stiff-shafted, closed-vaned (= barbules bearing hooked distal pennulae), pennaceous feathers arising from the distal forelimbs and tail
AND
possessed feathered wings used in flapping flight



Alvarezsaurids and oviraptors are taxa within Euparaves. They are secondarily flightless, primitive birds that descended from flying basal Euparaves.

Here is a sample of flying, basal Euparavians:
Scansoriopterygids (Temporal range: Late Jurassic, 165–156 Ma)
Anchiornis (Temporal range: Late Jurassic, 161–160.5 Ma)
Aurornis (Temporal range: Late Jurassic, 160 Ma)
Xiaotingia (Temporal range: Late Jurassic, 160 Ma)
Pedopenna (Temporal range: Middle or Late Jurassic, 164 Ma)

These require further analysis:
Eosinopteryx? (Temporal range: Late Jurassic, 160 Ma)
Zhongornis? (Temporal range: Early Cretaceous, 122 Ma)
Zhenyuanlong?
Archaeopteryx?
Microraptor?
Jeholornis?
Ornithomimosaurs?
Therizinosaurids?
Velociraptor?
Jixiangornis?


Note.
Oviraptors, Alvarezsaurids and Eudromaeosaurids are secondarily flightless, primitive birds within Euparaves.

"Indistinguishable from random"

Here is a study on how the published dinosaur cladograms compare to the stratigraphic (time) record. It is significant that there is a total lack of congruence, it is "indistinguishable from random", for Paraves.

Wills et al (2008):
http://sysbio.oxfordjournals.org/cgi/content/full/57/6/891 (Full article)
http://sysbio.oxfordjournals.org/cgi/content-nw/full/57/6/891/TBL1 (Table 1)

Evidence for the evolutionary history of most groups derives from two independent sources. The first is the distribution of phylogenetically informative characters or markers in extant and extinct taxa. The second is the stratigraphic or temporal sequence in which taxa occur as fossils. Neither source of data can be read uncritically, and both require interpretation. Phylogenies incorporate assumptions concerning rooting and models of evolution. The resulting trees are therefore inferences rather than data. Fossils require varying degrees of interpretation depending upon the nature of the material, and dates may be subject to large margins of error. For these reasons, it is often desirable to compare inferences by mapping cladograms onto stratigraphic range charts

"Over all 19 data sets, congruence was extremely high (Table 1). The average GER (Wills, 1999) for static first occurrence dates was 0.767, with the best data set being the Hadrosauridae (Horner et al., 2004; 0.956) and the worst being the Paraves (Turner et al., 2007; 0.558)."

" The least congruent data sets were the Paraves (Turner et al., 2007; GERt = 0.571), Stegosauria (Galton and Upchurch, 2004b; GERt = 0.611), Prosauropoda (Galton and Upchurch, 2004a; GERt = 0.720), and Ceratopsia (Xu et al., 2002; GERt = 0.755)."

Paraves is objectively calculated as the worst in terms of congruence. In fact, the article says that it is "indistinguishable from random". And that is because it is based on the incorrect dino-to-bird idea.

This is even worse when you consider the high congruence of dinosaurs in general:

Preliminary reanalysis of the 1,000 animal and plant “static” data sets utilized by Benton et al. (2000) and Wills (2007) yielded an average GER* of 0.688, with only 34% attaining a GER* > 0.975. This strongly suggests that the congruence of dinosaur phylogenies is better than that for a large sample of data sets across a range of other taxa.