Several book reviews for ‘The Rise of the Reptiles’ by H-D Sues 2019

Figure 1. Book cover for 'The Rise of Reptiles' by HD Sues.

A new book on evolution,
‘The Rise of Reptiles’ (left), came out last August 2019. Unfortunately, it took this long to come to my attention.  After a short bio on author, Dr. Hans-Dieter Sues, several reviews follow.

According to Amazon.com, author Hans-Dieter Sues is 
“a senior scientist and curator of fossil vertebrates at the Smithsonian’s National Museum of Natural History. He is a coauthor of Triassic Life on Land: The Great Transition.”


Amazon.com called ‘The Rise of the Reptiles’,
“The defining masterwork on the evolution of reptiles.”

John Long, Finders University, called it,
“A valuable reference book―entertaining read and beautifully illustrated―about how reptiles first evolved and diversified into many lineages, including the one leading to us mammals. The Rise of Reptiles is essential reading for every evolutionary biologist.”

Michael J. Benton, University of Bristol, reported,
“The writing style is clear and easy, the illustrations are excellent, and the whole design and print quality highly attractive. There is no other book like it, and this will stand as a useful reference for many years.”

Jeremy B. Stout, Quarterly Review of Biology, called it, 
“The most complete and current compendiumon reptilian evolution and diversity to date… Few (if any) are better suited to have written this volume than Sues. His impressive research record over the past 40 years has dealt directly with many of the taxonomic groups in this volume, including (nonreptile) synapsids, parareptiles, sauropterygians, crocodylomorphs, and dinosaurs.”

After scientifically testing the various hypotheses put forth by Sues
in the Large Reptile Tree (LRT, 1680+ taxa) his book was found to promote many of the traditional myths and misconceptions that have befuddled reptile cladograms for the past decade. Virtually all of the problems in ‘The Rise of the Reptiles’ can be attributed to too few taxa in the various cited phylogenetic analyses.

Starting with outgroups, Sues reports,
“The group generally considered most closely related to amniotes is Diadectomorpha.” He cites papers from 1980, 2003 and 2007.

Using the last common ancestor method, the tested out-group to the clade Reptilia in the LRT (2020) is the Reptilomorpha. This clade includes the long-limbed taxon, Gephyrostegus, a late survivor from an earlier, probably Late Devonian, radiation. When more taxa are added diadectomorphs nest well within the Lepidosauromorpha branch of the Reptilia (Fig. 5), contra Sues 2019.

Dr. Sues “Pulls a Larry Martin” when he reports
on a few traits he notes are present in his short list of stem-amniote ‘outgroups’.

Those outgroups (diadectomorphs, which are ingroups in the LRT) were gleaned from old studies. Look what happens when you don’t have the correct outgroups and correct last common ancestors. The traits you observe are going to be wrong because you’re not even looking at the correct node. Never rely on a few traits to define a clade. Always rely on the ‘last common ancestor’ method to determine clade membership. And don’t exclude pertinent taxa!

Figure x. "Classification" of reptiles according to Sues 2019. Color overlay note differences when more taxa are added, as in the LRT.

Figure 2. “Classification” of reptiles according to Sues 2019. Color overlay note differences when more taxa are added, as in the LRT.

Sues reported on ‘the oldest known reptile, Hylonomus‘ 323-315mya,
but that was based on a pre-cladistic 1964 study. Sues then hedged that ‘undisputed’ report with two ‘recent phylogenetic analyses’ (1988 and 2006) that, “instead found these taxa [Hylonomus + Palaeothyris] closely related to diapsid reptiles.”

Using the last common ancestor method, the oldest known reptile in the LRT (2020) is Silvarnerpeton (346-323 mya). No hedging required here. All descendants laid amniotic eggs.

Figure 3. Reptilia vs. Amniota in Sues 2019 compared to the LRT.

Figure 3. Reptilia vs. Amniota in Sues 2019 compared to the LRT.

Sues cites Tsuji and Müller (2009) who defined ‘Parareptilia’ as
“the most inclusive clade containing Milleretta and Procolophon, but not Captorhinus” and that clade includes such diverse taxa as mesosaurs and pareiasaurs (Fig. 4).

If those two are related, that’s a Red Flag. In the LRT mesosaurs nest with similar pachypleurosaurs and other marine diapsids. Pareiasaurs give rise to turtles. As defined, the clade ‘Parareptilia’ is paraphyletic at worst, and a junior synonym of Reptilia at best in the LRT, which includes a wider gamut of taxa.

Figure x. Parareptilia according to Sues 2019.

Figure 4. Parareptilia according to Sues 2019.

Similar problems
attend the cladogram of ‘Eureptilia’ in Sues 2019 (Fig. 6).

More taxa solve this problem, too.

Figure x. Simplified cladogram of the Reptilia according to Sues 2019 (below in white) compared to a simplified version of the LRT using the same taxa.

Figure 5. Simplified cladogram of the Reptilia according to Sues 2019 (below in white) compared to a simplified version of the LRT using the same taxa. More taxa reveal an earlier dichotomy that creates two diapsid-grade skull morphologies. 

Dr. Sues has no idea how reptiles diverged
from their Viséan (or earlier) initial radiation. This could have been repaired if he had simply added taxa to his own analysis, rather than relying on published academic papers published decades ago with the same flaws.

Figure x. The 'Eureptilia' according to Sues 2019. This is a paraphyletic clade when more taxa are included, as in the LRT.

Figure 6. The ‘Eureptilia’ according to Sues 2019. This is a paraphyletic clade when more taxa are included, as in the LRT. Since the invalid clade ‘Sauria’ includes lepidosaurs and archosaurs it is half colored blue.

Other than cladograms, Sues 2019
also presents photos and diagrams (Fig. 7). Unfortunately some diagrams don’t match the fossils, leading to confusion at best.

Figure x. Figure from Sues 2019 showing Youngina capensis and a diagram of the same, that does not match the fossil. DGS color overlay added for comparison.

Figure 7. Figure from Sues 2019 showing Youngina capensis and a diagram of the same, that does not match the fossil. DGS color overlay added for comparison. What is the specimen number for this specimen?

Exposing and overturning old and new reptile mythology
is what PterosaurHeresies.Wordpress.com is all about as it supports the website www.ReptileEvolution.com and its centerpiece, the growing online cladogram at: www.ReptileEvolution.com/reptile-tree.htm. A wide-gamut cladogram is a powerful tool providing evidence against invalid traditional hypotheses that exclude pertinent taxa.

I cannot recommend this book.


References
Sues HD 2019. The Rise of Reptiles: 320 Million Years of Evolution.
Johns Hopkins University Press, Baltimore. xiii + 385 p.; ill.; index.
ISBN: 9781421428673 (hc); 9781421428680 (eb).

 

Fact checking dino books

As stated from the beginning
of this blog, I am more interested in clade origins and relationships than any other paleo topic (like diet or asteroid impact). So the following review of recent and important dino books will focus on what they say about the origin of dinos: i.e. what happens when taxon exclusion meets the LRT.

Figure 1. Dinosaurs a Concise Natural History by Weishampel and Fastovsky, editions 1, 2 and 3.

Figure 1. Dinosaurs a Concise Natural History by Weishampel and Fastovsky, editions 1, 2 and 3.

Dinosaurs: a concise natural history (Weishampel and Fastovsky 2009)
This book defines the living reptiles as: turtles + diapsids (including living birds)

That is outdated
according to the large reptile tree (LRT, 1240 taxa) where mammals are also reptiles, and diapsids are not monophyletic. According to the LRT reptiles can be defined as any archosauromorph (e.g. Eldeceeon, mammals, ichthyosaurs, archosaurs), plus any lepidosauromorph (e.g. Urumqia, captorhinids, turtles, lepidosaurs), their last common ancestor (e.g. the amphibian-like reptiles Silvanerpeton, Gephyrostegus) and all descendants.

In the W+F book there are two main clades of Diapsids: Lepidosauromorpha and Archosauromorpha.

This is outdated.
See above.

In the W+F book there are five main clades of Archosauromorpha: Rhynchosauria, Prolacertiformes, Crurotarsi (including Crocodylia), Pterosauria, Dinosauria.

This is outdated.
Their way-too-simplified-cladogram is what makes paleontology and evolution look bad due to taxon exclusion. No taxa display a gradual accumulation of traits in the W+F cladogram. You can’t have super specialized rhynchosaurs at the base of any large clade. They are terminal taxa arising from sphenodontids in the LRT. Pterosaurs share nothing in common with their purported W+F sisters, crocs and dinos. Pterosaurs actually arise from fenestrasaur, tritosaur lepidosaurs. Another terminal taxon, Tanystropheus, the poster-child for prolacertiforms is not related to Prolacerta in the LRT, but arises from Huehuecuetzpalli, Macrocnemus and Langobardisaurus, among others, not far from pterosaurs. Derived taxa in this clade share a nearly identical foot morphology.

The W+F book presents two hypotheses of dinosaur origins:

  1. pterosaurs + Lagosuchus + dinosaurs (Gauthier and Padian) = Ornithodira.
  2. Lagosuchus, Lagerpeton, Pseudolagoschus, Marasuchus (Sereno) = dinosaur precursors. Pterosaurs are more distantly related.

These two hypotheses are both outdated
Pterosaurs are lepidosaurs, unrelated to dinosaurs. Lagerpeton is a sister to the chanaresuchid, Tropidosuchus. Marasuchus is a theropod, not a dinosaur outgroup. In the LRT bipedal crocs are the proximal outgroups to dinosaurs.

________________________________________

Figure 2. Rise and fall of the dinosaurs by Brusatte 2018.

Figure 2. Rise and fall of the dinosaurs by Brusatte 2018.

The rise and fall of the Dinosaurs. Brusatte 2018. Don’t confuse this 2018 book with ‘The rise and fall of the dinosaur’ (singular, not plural) by Wallace 1987.

Unfortunately
Brusatte clings to his wrong-from-the-start hypothesis (Brusatte et al. 2011) that the first dinosaurs are represented by Prorotodactylus (Fig. 3) footprints in the Early Triassic (251 mya). Prorotodactylus tracks were actually made by tritosaur lepidosaurs, like Macrocnemus, with a small, symmetrical, 5-fingered manus and an asymmetric pes with a short digit 5.

Unfortunately
Brusatte’s mistake comes from a misunderstanding of reptile relationships, probably because he was schooled using the Weishampel and Fastovsky books (see above). Brusatte was familiar with the invalid Lagerpeton hypothesis of dinosaur origins because he was an undergrad student of Paul Sereno (see above). Thus Brusatte was ready to accept the asymmetric pedal track of  Prorotodactylus track as homologous with that of Lagerpeton. Brusatte was also a student of Mike Benton. You might remember, Benton infamously promoted the bipedal croc with tiny hands and no pedal digit 5, Scleromochlus, as a pterosaur sister. Later Benton joined with his other student, David Hone (Hone and Benton 2007, 2008) in deciding to include, then exclude Peters 2000 data on fenestrasaurs. In so doing Hone gained his PhD and pterosaur origins were taken back to the dark ages. We talked about professor/student influences earlier here. In my youth I was influenced by books and professors, so I know how readily that can happen. If the influence is valid, that’s fine. If not, that’s a problem.

Porotodactylus pes

Figure 2. Porotodactylus pes and manus ichnites. Latest Early Triassic.

The actual last common ancestor of all dinos (and all crocs, too) in the LRT is the Túcuman specimen wrongly attributed to Gracilisuchus, PVL 4597 (Lecuona and Desojo 2011; Fig. 4) which has been known for 7 years.

It’s unfortunately ironic
that as much as Dr. Brusatte loves tyrannosaurs, as co-author (Lü and Brusatte 2015) he also misidentified Zhenyuanlong as a dromaeosaurid, not a tyrannosaur ancestor, as we looked at earlier here. He also listed a number of unrelated taxa, like Dilong, as tyrannosaur ancestors in an earlier magazine article discussed here. So his track record regarding dino systematics is not validated in the LRT.

Figure 6. The closest known taxa to the Dinosauria, PVL 4597, is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

Figure 4. The closest known taxon to the Dinosauria in the LRT, PVL 4597 (Lecuona and Desojo 2011, Ladinian (Late Miiddle Triassic). It is a tiny taxon (phylogenetic miniaturization) with erect hind limbs, a large and deep pelvis and a tiny calcaneal tuber.

Finally,
let’s remember that paleontology works at a snail’s pace. We’ve know that birds are dinosaurs since TH Huxley looked at Archaeopteryx in the 1860s, but workers were not keen to accept that. Then Ostrom 1969 echoed Huxley’s hypothesis with new discoveries of Deinonychus. Still to little to no progress in the paleo community.

According to
the Hartford Courant (2000), “In 1973, Ostrom broke from the scientific mainstream by reviving a Victorian-era hypothesis (see above) that his colleagues considered far-fetched: Birds, he said, evolved from dinosaurs. And he spent the rest of his career trying to prove it.” With the announcement of the first dinosaurs with feathers from China, Ostrom (then age 73) was in no mood to celebrate. He is quoted as saying, ““I’ve been saying the same damn thing since 1973, `I said, `Look at Archaeopteryx!’” Ostrom was the first scientist to collect physical evidence for the theory. Ostrom provoked a debate that raged for decades. “At first they said, `Oh John, you’re crazy,”’ Ostrom said in 1999.”

It wasn’t until dozens of Chinese specimens
appeared with feathers that paleontologists finally got out of their academic chairs and said in essence, ‘well, we better change our mind about feathers and dinosaurs, or else.’ The scaly theropods in the Jurassic Park/Jurassic World movies reflect that slow-to-accept mentality.

The Dinosaur Heresies book by Dr. Robert Bakker.

The Dinosaur Heresies book by Dr. Robert Bakker 1986.

Books should not just echo false tripes
and recirculate untenable cladograms. Books should break old paradigms and advance new and valid ideas. Case in point: The Dinosaur Heresies by Bakker 1986.

References
Bakker RT 1986. The dinosaur heresies. New theories unlocking the mystery of the dinosaurs and their extinction. Citadel Press.
Brusatte S, Niedźwiedzki G and Butler RJ 2011. Footprints pull origin and diversification of dinosaur stem-lineage deep into Early Triassic. Proceedings of the Royal Society of London, Series B, 278, 1107-1113.
Brusatte S 2015. Rise of the Tyrannosaurs. Scientific American 312:34-41. doi:10.1038/scientificamerican0515-34
Brusatte S 2018. The rise and fall of the dinosaurs. Harper Academic. author interview
Hone DWE and Benton MJ 2007. An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles. Journal of Systematic Palaeontology 5:465–469.
Hone DWE and Benton MJ 2008. 
Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.
Lü J and Brusatte SL 2015. 
A large, short-armed, winged dromaeosaurid (Dinosauria: Theropoda) from the Early Cretaceous of China and its implications for feather evolution. Scientific Reports 5, 11775; doi: 10.1038/srep11775.
Ostrom JH 1969. “Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana”Peabody Museum of Natural History Bulletin. 30:1–165.
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Wallace J 1987. The rise and fall of the dinosaur. Gallery Books
Weishampel DE and Fastovsky DB 2009: Dinosaurs: a concise natural history. 3 editions. Cambridge University Press. pdf

wiki/Zhenyuanlong

https://pterosaurheresies.wordpress.com/2016/03/16/sometimes-it-takes-the-paleo-crowd-an-epoch-to-accept-new-data/

The Tyrannosaur Chronicles by David Hone

A new book
by Dr. David Hone called The Tyrannosaur Chronicles is now out. He reports here, “Although there is no more famous and recognisable dinosaur than Tyrannosaurus, the public perception of the animal is often greatly at odds with the science. The major image people have of them is the iconic jeep chasing scene in the film Jurassic Park. However, because they are among the best-studied of all dinosaurs, we can say that the tyrannosaurs almost certainly had feathers and may have fought and even ate each other.”

Figure 1. The Tyrannosaur Chronicles by Dr. David Hone is a new book chronicling tyrannosaurs.

Figure 1. The Tyrannosaur Chronicles by Dr. David Hone is a new book chronicling tyrannosaurs.

I have not read the book yet, but I’ll note a possible problem gleaned from quote pulled from a review.

Kirkus Reviews reports: While correctly surmising that tyrannosaurs and other dinosaurs were carnivores, scientists erroneously assumed that they were some kind of previously unknown “giant land reptile.” Subsequent fossil discoveries in polar regions ruled out this possibility since coldblooded reptiles could not survive such extreme cold weather.”

I hope this is a misquote or I’m misreading this. It’s not news that tyrannosaurs and dinosaurs have been and will always be giant land reptiles. They nest in the clade Reptilia, no matter how cold-adapted they might have been. Hone might be going back, back in time to the first English discoveries from 50 years earlier, like Iguanodon and Megalosaurus, the first dinosaurs, which were named terrible lizards, and originally titled, “British Fossil Reptiles.”

And I hate to judge a book by its cover, but…
That small crested dinosaurs in the lower left corner is Guanlong, an ancestor not of tyrannosaurs, but of allosaurs in the large reptile tree. No word yet if Hone included the verified ancestors of tyrannosaurs, Zhenyuanlong, Tianyuraptor and Fukuiraptor.  On that note, GotScience.org evidently quotes Hone when it reports, Early tyrannosaurs had crests used for sexual display and social rank.”

Book and academic publishing is fraught with such risk and danger. Once you print it, you can’t retract or revise it. Sympathetically, I know from experience the things I would have changed about my early papers now, but was less experienced then.

Thankfully
I hear that Hone discusses feathers and such.

Amazon Reviews are universally positive:

  1. Dinosaurs are endlessly fascinating, and the massive, blood-thirsty tyrannosaurs are most popular (and scary) of the lot! Here, renowned dinosaur expert David Hone reveals their story, and how we know what we know about these most amazing of ancient reptiles. — Professor Mike Benton, University of Bristol
  2. Tyrannosaurs are probably the world’s favourite dinosaurs. But what do we really know about this group? David Hone reviews the biology, history, evolution, and behaviour of the tyrant kings – an excellent read, containing the very latest in our understanding of Tyrannosaurus rex and its closest relatives. — Dr Tom Holtz, University of Maryland
  3. Without doubt, the best book on tyrannosaurs I’ve ever read. This is an awesome dinosaur book. — Professor Xu Xing, Chinese Academy of Sciences

Do not be confused with this website:
http://traumador.blogspot.com which earlier featured ‘Traumador the tyrannosaur in the Tyrannosaurus Chronicles’ which can be silly and serious all on the same blog, explained here as:

The Tyrannosaur Chronicles is a blog written by Traumador the Tyrannosaur about his many exploits.Traumador is a tyrannosaurid who hatched from an egg that magically survived the K/Pg Extinction Event and was discovered in Alberta by Craig, an aspiring paleontologist (and the mastermind behind the blog in real life). He eventually gets a job at the Royal Tyrell Museum and things get interesting from there.

From past experience,
such as when Hone attempted to compare the two hypotheses of pterosaur origins by dropping one, or when Hone attempted to show that Dmorphodon had a mandibular fenestra, or when Hone supported the deep chord bat wing model for pterosaur wings, or when Hone flipped the wingtips of Bellubrunnus, we might be wary about what Dr. Hone puts out there. But I don’t think you can go very wrong with tyrannosaurs, the most studied dinosaur. And the reviews speak high praise.

Scathing Book Review – Pterosaurs (Witton 2013) – Wing membrane mistakes

Earlier here, here, here, here and here we looked at the new book, “Pterosaurs” by Mark Witton (2013) and the many problems, mistakes and oversights that book contains. Today we’ll look at how Witton attempts to support the traditional hypothesis of a deep chord wing membrane attached at the ankle (Fig. 1). We covered that topic earlier here.

Figure 1. From Witton (2013) the Vienna specimen of Pterodactylus with a beautifully preserved wing membrane that is shallow at the elbow, stretched between the wingtip and elbow  and includes a small fuselage fillet back to mid thigh.

Figure 1. Click to enlarge. From Witton (2013) the Vienna specimen of Pterodactylus with a beautifully preserved wing membrane that is shallow at the elbow, stretched between the wingtip and elbow and includes a small fuselage fillet back to mid thigh. Witton’s caption indicates the ‘in vivo’ wing membrane is not accurately represented here ‘in situ’. He considers the membrane, “somewhat desiccated, shriveled.” So, what happens when the data doesn’t fit the hypothesis? Witton blames the data. The uropatagium and toe webbing, by the way, were unaffected.

When a bad hypothesis meets good data
Witton (2013) upholds the deep chord wing membrane hypothesis. So when excellent data indicates a shallow chord with fuselage fillet (the Stromer model, Fig. 8), Witton invents excuses for it (Fig. 1), calling it “dessicated” and “shriveled.” There are no other examples of desiccated membranes in pterosaur fossils. In any case, wing membranes contained little water in life, so how could they shrink when desiccated, especially when buried underwater? Adding to the confusion, on the same page (Witton 2013: 196), he illustrates a nicely preserved uropatagium and webbed toes from the same specimen. These, strangely enough, have not suffered at all from desiccation!

To clarify things
Peters (2002) published an image (Fig. 8) of what happens when the Vienna specimen unfolds its wings. Here it is animated (Fig. 2). Witton ignored this paper.

The Vienna Pterodactylus.

Figure 2. The Vienna Pterodactylus. Click to animate. Wing membranes in situ (when folded) then animated to extend them. There is no shrinkage here or in ANY pterosaur wing membrane. There is only an “explanation” to avoid dealing with the hard evidence here and elsewhere.

The real wings, based on the excellent data provided by the Vienna specimen, demonstrate the membranes were stretched between the wingtip and elbow, not the ankle.

Note: In all pterosaurs (Fig. 2) the wings were functionally disconnected from the hind limbs (contra the traditional view upheld by Witton), which means pterosaurs would have no trouble running around on the ground, bipedally or quadrupedally.

Witton (2013) misinterprets the pterosaur wing membrane
Earlier here and elsewhere we looked at the true shape of pterosaur wing membranes. Unfortunately, Witton (2013) promotes a completely different view. And this is not just a matter of opinion or sides in an argument. This fact vs. fiction (Fig. 1).

Pterosaur wing membranes according to Witton (2013). Mistakes here include the 1. deep chord wing membrane attached at the ankles, 2. trailing edge structure, 3. single uropatagium on Rhamphorhynchus.

Figure 3. Pterosaur wing membranes according to Witton (2013). Mistakes here include the 1. deep chord wing membrane attached at the ankles, 2. trailing edge structure, 3. single uropatagium on Rhamphorhynchus.

Here's how the wing membrane in pterosaurs virtually disappeared when folded.

Figure 4. Here’s how the wing membrane in pterosaurs virtually disappeared when folded. There is absolutely no connection of the wing membrane to the ankle. Rather it goes to the elbow, as shown here, then to mid thigh as a fuselage fillet. Note how readily it disappears when folded!

The Zittel wing

Figure 5. The Zittel wing from a species of Rhamphorhynchus. Click to enlarge. Here it is more than clear that the wing is very shallow at the elbow. There’s the humerus. All that’s missing is the body. Here the wing is stretched between the wingtip and elbow. What could be more clear? I’ve added a body (Fig. 8) if you really need one.

Non-believers
Upon seeing this image (Fig. 5) promoted as a great example of a narrow-chord wing membrane, Darren Naish considered this “ambiguous” because the specimen has no torso (Fig. 8). Why do you need a torso? You have the entire wing! And it follows the Vienna Pterodactylus (Fig. 1) configuration – narrow at the elbow (and the BRI 010 specimen, see Fig. 6!). There’s no trailing membrane here that – can – extend to the ankle (Fig. 8). There’s no example of such a wing membrane anywhere, except, to some eyes, in Sordes. But that was demonstrated to be due to a disarticulated radius and ulna, which can be identified in the fossil.

Now, just to be fair and open, 
Helmut Tischlinger kindly sent me a short paper (Monninger et al. 2012) describing a new Rhamphorhynchus specimen BRI 010 (Fig. 6), in which “a linear structure runs along the trailing edige from the ankle to the tip of the wing finger.” That’s exciting. If verifiable, this would support the Witton model. Unfortunately, requests for hi-rez images of this specimen have gone unanswered. But I can present the image in low resolution from the abstracts, which are available online.

Rhamphorhynchus BRI 010. Here, unfortunately, the ankle is preserved right next to the elbow. Notably the wing membrane makes a beeline for the elbow, not curving posteriorly to meet the ankle. This is what we also see in the darkwing Rhamphorhynchus.

Figure 6. Rhamphorhynchus BRI 010. Click to enlarge. Color added. Here, the wing was reported to extend to the ankle. Unfortunately, the ankle is preserved right next to the elbow. Notably, the wing membrane does make a beeline for the elbow, not curving posteriorly to meet the knee. This is the configuration we also see in the darkwing Rhamphorhynchus.

Even at low resolution
we can tell that this is a Rhamphorhynchus with a well-preserved wing membrane with a trailing edge directed precisely at the elbow, as in the darkwing Rhamphorhynchus. Unfortunately the ankle has swung forward to the elbow. If the narrow-chord wing is correct, then this image needs no further explanation. If the deep-chord wing is correct we should see some sort of bending or curving toward the knee of the trailing edge (as shown in Fig. 3), but there is none.

Thank goodness!
By the text of the short paper I thought I was going to have to apologize to the pterosaur experts.

Linear structure on the trailing edge?
I can’t comment on the linear structure along the trailing edge, having never seen one, but then I haven’t seen this specimen in high resolution either. We’ll have to wait.

Pteroid location
As part of the leading edge, the propatagium and pteroid are also important aspects of the pterosaur wing. Witton (2013) ignored the evidence and his own text that the pteroid articulated with the proximal carpal (radiale) and instead sometimes illustrated it in the cup of the preaxial carpal (Fig. 7), sometimes locating it on the radiale, sometimes on the distal carpal. His text does report the preaxial carpal articulated with the distal carpal, so why the imprecision? Witton reports the pteroid “probably articulated in a depression on the medial side of the preaxial carpal (Bennett 2007) and MAY have also contacted the proximal syncarpal (Peters 2009).” Actually, with the way the wrist bent, the pteroid would rarely if ever contact the preaxial carpal. They diverged from one another and were separated by the length of the proximal and distal carpals.

Dimorphodon by Mark Witton, filled with errors.

Figure 7. Dimorphodon by Mark Witton, with the pteroid articulating with the cup of the preaxial carpal.

Pteroid origin
Witton did not report the pteroid and preaxial carpal were present on Cosesaurus (Peters 2009), first observed but misidentified by Ellenberger (1993) who, unfortunately, flipped that manus over in order to have digit 2 longer than the lateral digits.

Anyway, a series of errors over time ultimately brings us to the truth of what is actually happening with the wings of pterosaurs, a configuration first envisioned by Stromer in 1910 (Fig. 8b).

The Vienna wing and the Zittel wing.

Figure 8. Click to enlarge. The Vienna wing and the Zittel wing. Both Zittel and Peters 2002 added a body to the Zittel wing and arrived at the same unambiguous conclusion. The caption within the figure provides details.

References
Monninger S, Frey E and Tischlinger H 2012. Supporting structures in the flight membrane of pterosaurs. EAVP abstracts, Teruel, España. Royo-Torres, R., Gascó, F. and Alcalá, L., coordinators. 10th Annual Meeting of the European Association of Vertebrate Palaeontologists. ¡Fundamental! 20: 1–290.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. – Historical Biology 15: 277–301.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29:1327-1330.
Witton M. 2013. Pterosaurs. Princeton University Press. 291 pages.

Scathing book review – Pterosaurs (Witton 2013) – finger mistakes

Earlier here and elsewhere in this blog we looked at the various mistakes and oversights promoted by Mark Witton (2013) in his new book, “Pterosaurs.” Today we’ll look at the configuration of the three free fingers and the many problems with Witton’s proposed configuration (Fig. 1).

Figure 1. Here Witton promotes the idea that pterosaur metacarpals were pressed back to back with the palmar side anterior, the pteroid articulating with the preaxial carpal and the elbow overextended.

Figure 1. Here Witton (2013) promotes the idea that pterosaur metacarpals were pressed back to back with their palmar sides anterior, the pteroid articulating with the preaxial carpal and the elbow overextended. None of these are supported by fossil evidence. 

Witton (2013) promotes the traditional idea that the dorsal surfaces of the three free fingers were appressed to the anterior surface (the former dorsal surface) of the axially rotated metacarpal 4. This means the free fingers would be oriented palmar side anterior during flight (Fig. 1) and extended palmar side ventro-posterior during terrestrial locomotion. Such a configuration  allows no room for the extensor tendons, especially the big one for the wing finger (Fig. 2). Moreover, posterior fingers do not match known pterosaur hand ichnites.

The actual orientation and configuration of the three fee fingers is shown below (Fig. 2).

Figure 2. The hand of the most primitive pterosaur with only metacarpal 3 attached to metacarpal 4. This provides room for extensor tendons. It also enables lateral finger extension during terrestrial locomotion and medial grappling during tree clinging. Crushing rotates the unguals anteriorly in this case. The extensor tendon process would bang into digit 2 when fully extended in the Witton model.

Figure 2. The hand of the most primitive pterosaur with only metacarpal 3 attached to metacarpal 4. This provides room for extensor tendons. It also enables lateral finger extension during terrestrial locomotion and medial grappling during tree clinging. Crushing rotates the unguals anteriorly in this case. The extensor tendon process would bang into digit 2 when fully extended in the Witton model.

The actual configuration is supported by evidence.
The present configuration provides room for the extensor tendons and permits the extensor tendon process to move unimpeded whenever fully extended in flight. With the free fingers facing palmar side down, like all other tetrapods, during terrestrial locomotion the fingers would extend laterally, matching ichnites. During tree clinging the fingers would grapple medially rather than, as Witton proposes, opening anteriorly, in a “begging” configuration.

The metacarpal “drawbridge”
The connection between metacarpals 3 and 4 produces a hinge. That’s why the metacarpus sometimes swings up during taphonomy to give the impression of the configuration that Witton proposes, as if the “drawbridge” was raised.

Pterosaur-fingers

Figure 3. Pterosaur wrist and hand images from Witton (2013). Neither demonstrates the palmar side forward configuration. The lower Pteranodon image is in posterior view and the stacking of the metacarpals is therefore anterior, away from the viewer, as in the Peters configuration. The wing finger of the Pteranodon would have been in the Z-axis, toward the viewer, but it was disarticulated during burial. Bennett (1991, 2000) thought metacarpal 4 was axially rotated during taphonomy. Not so. Green = flexor tendons. Red = extensor tendons.

So, where did Witton get his configuration?
From Bennett (2008), and this Santanadactylus specimen (Fig. 4, ignoring, apparently all others).

Santanadactylus hand and fingers

Figure 4. Santanadactylus hand with metacarpals preserved at a 45 degree angle to the anterior face of metacarpal 4. This demonstrates the drawbridge effect in which Bennett elevates metacarpals 1-3, raising the drawbridge. Actually, the drawbridge was down in life. Imagine what would happen to fingers 1-3 when the wing finger was extended! And where were the extensor tendons? Only lowering the drawbridge solves all these problems.

The Santandactylus specimen preserves metacarpals 1-3 raised, like a drawbridge to about 45 degrees, but this is a taphonomic artifact. Bennett’s (2008) configuration further raised the metacarpus to appress against metacarpal 4, palmar side anterior. Peters’s (2002) configuration lowers the metacarpus on its hinge, palmar side down. This provides room for the large extensor tendon and the extensor tendon process of the wing finger when it is fully extended in flight. This also matches all undisturbed pterosaur fossils (Fig. 2).

Added note:
I’m hearing that my notes hear are not swaying the experts. Please see the conversations below and check out this link to pterosaur finger flexion.

References
Bennett SC 2008. Morphological evolution of the forelimb of pterosaurs: myology and function. Pp. 127–141 in E Buffetaut and DWE Hone eds., Flugsaurier: pterosaur papers in honour of Peter Wellnhofer. Zitteliana, B28.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. – Historical Biology 15: 277–301.
Witton M. 2013. Pterosaurs. Princeton University Press. 291 pages.

Scathing Book Review – Pterosaurs by Mark Witton 2013 – part 2

I finally got the new Witton pterosaur book from Amazon.
Most of the topics you’ll read here have been posted before.

pterosaurs-wittonWith his new book, Pterosaurs, Witton (2013) continues to stick his head in the sand (or wear his professional blinders), avoiding and dismissing the best testable evidence for pterosaur origins, wing shape, take-off, phylogeny, ontogeny, morphology, gender identification and reproduction. (Which is why the Pterosaur Heresies is needed, to right these wrongs). Here’s yet another expert disfiguring pterosaurs big time.

Of  course his artwork is beautiful, flaws and all. And his writing style is friendly, informative and a joy to read, until you come up against bogus information and images. Then you wonder why has the world gone topsy-turvy, where amateurs provide better, more accurate evidence and more parsimonious explanations than professionals do?!

And it’s not just that we disagree.
I am pointing out factual errors here that can be tested by looking at specimens.

Case in point
Earlier we talked about the first few chapters of Pterosaurs in which Witton ignores the four outgroup taxa closest to pterosaurs: Langobardisaurus, Cosesaurus, Sharovipteryx and Longisquama. Witton did produce his version of Sharovipteryx, which explains much of the problem and why he dismissed it. Here it is (Fig.1). See if you can see where Witton pays little heed to accuracy.

Figure 2. This is what scientists call complete fantasy and total disregard for data. Upper images from Witton 2013, in which he simply made up the proportions of the pedal elements for Sharovipteryx. No wonder he didn't see the phylogenetic connection to pterosaurs! Below, the actual proportions traced from an 8x10 transparency taken after personal examination of the fossil. Like pterosaurs, cosesaurs, langobardisaurs,  Tanystropheus and Huehuecuetzpalli, Sharovipteryx had a short metatarsal 5 and an elongated p5.1. It's a key trait for this clade. Don't tell me pterosaurs just appeared out of nowhere. Here's the evidence of kinship.

Figure 2. This is what scientists call complete fantasy and total disregard for data. Upper images from Witton 2013, in which he simply made up the proportions of the pedal elements for Sharovipteryx. Lower image from yours truly after examining the specimen firsthand. No wonder he didn’t see the phylogenetic connection to pterosaurs! Like pterosaurs, cosesaurs, langobardisaurs, Tanystropheus and Huehuecuetzpalli, Sharovipteryx had a short metatarsal 5 and an elongated p5.1. It’s a key trait for this clade. Don’t tell me pterosaurs just appeared out of nowhere. Here’s the evidence of kinship.

I rolled my eyes so far back that I actually saw my brain.
Witton (2013) disfigured Sharovipteryx by completely imagining the proportions of the pedal elements. There’s not even a feeble attempt at accuracy here. And because Witton put his blinders on he completely missed the unique morphological similarities in the pes shared by Sharovipteryx and pterosaurs. This is why I earlier stated that Witton was ill-prepared to write a book on pterosaurs. This is not about ‘not knowing’ the correct data. This is about ‘not wanting to know’ the correct data, which has been around for forty years.

If you are of the opinion
that my work (Fig. 1) is flawed, check out the original paper, Sharov (1971), who made the same tracing.

And if you’re friends with Mark
Yes, he’s a great guy and tries hard, but he fkd up here. Don’t run to his defense. There is no defense for this. Earlier Witton slammed ReptileEvolution.com in general. Here, as elsewhere, I’m being surgically precise with my critique. I’m simply trying to lift the blinders off those who profess to be experts in pterosaurs. If you’re an expert, act like it. Be professional. Test ideas and observations. Don’t just follow tradition, especially when you profess to not know the answer. And for Pete’s sake, don’t make up things out of your imagination.

Note that,
even in his figure of Sharovipteryx, Witton ignores several other key traits shared with pterosaurs to the exclusion of basal archosaurs: 1) Elongated and retracted naris (long premaxilla); 2) Large orbit, or is that the antorbital fenestra?; 3) Short torso (knee can reach the shoulder); 4) Elongated ilium (capturing more than four sacrals); 5) Attenuated caudals with chevrons parallel and appressed to centra; 6) Tibia longer than femur: 7) Fibula attenuated and 8) the big one, uropatagia (soft tissue trailing the hind limbs (Witton invents most of the soft tissue in front of the femur. See Fig 3.)). Evidently Witton eschews hard evidence and phylogenetic analysis. I find it answers many, many problems.

Figure 2. Sharovipteryx mirabilis in various views. No pycnofibers added yet. Click to learn more.

Figure 3. Sharovipteryx mirabilis in various views. Trailing membrane on the hand is guesswork based on phylogenetic bracketing. Note, there is a soft tissue flap in front of the femur, but it does not connect to the torso, which, in reality is circular in dorsal view with wide flat ribs. And yes, Sharovipteryx has prepubes, a pterosaurian trait inherited from Cosesaurus.

Simply having an elongated pedal digit 5 puts Sharovipteryx and pterosaurs outside of virtually all archosauriforms (they have vestiges) and squarely in kinship with tritosaur lizards, like Huehuecuetzpalli, which shares some of the traits listed above.

Witton doesn’t like pterosaurs as highly derived lizards
Witton (2013, p. 17) reports, “There seems little similarity between the skulls of pterosaurs and the highly modified, mobile skulls of squamates, or any similarity between the trunk and limb skeletons of each group.” This is, of course, bogus data (imprecise to untrue) to draw you off. Pterosaurs are not related to squamates (Iguania and Scleroglossa), but to a third, more basal lepidosaur clade, the Tritosauria, that did not have a mobile skull and did not fuse the ankle bones. Again, putting his blinders on, and following in the footsteps of Dr. David Unwin, Witton does not introduce his readers to the following lepidosaurs: Huehuecuetzpalli, Macrocnemus, Cosesaurus and Longisquama, all of which demonstrate a gradually increasing list of pterosaur traits as detailed here.

In order to further dismiss my work, Witton references Hone and Benton (2007) which has been lauded as one of the worst papers of all time based on the fact that they set up a battle between the fenestrasaurs and archosaurs, then eliminated the fenestrasaurs from consideration and declared archosaurs the winners. They also had typos in their matrix (found by Bennett 2012) which they used to dismiss data. And there were many other problems listed here. I just want to ask Dr. Witton, “Where is the critical thinking?” I know it’s easy to cozy up to your friends’ work and difficult to accept others’, but really, you have to examine the evidence without bias.

Final pertinent note
Witton reports that my work has received little attention due to my “highly controversial techniques used in his analyses and anatomical interpretations.” At least I don’t just make the stuff up (see Fig. 1) !!!!! Dr. Witton, this is really “the pot calling the kettle black.” Please look at the specimen or get precise references next time. It will solve lots of problems and get us back on the right track.

As always, if anyone has better data, I am known to frequently make corrections wherever warranted. Just made a bunch this week.

More later.

References
Bennett SC 2012. 
The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined. Historical Biology. iFirst article, 2012, 1–19.
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification. Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605
Sharov AG 1971. New flying reptiles from the Mesozoic of Kazakhstan and Kirghizia. – Transactions of the Paleontological Institute, Akademia Nauk, USSR, Moscow, 130: 104–113 [in Russian].

Book Review – Earth Before the Dinosaurs – Steyer

 Earth Before the Dinosaurs by Sébastien Steyer

Figure 1. Earth Before the Dinosaurs by Sébastien Steyer

Quite good – until you get to the reptiles.
Then mostly bogus in that it follows Carroll (1988) in putting Diadectomorphs as the outgroup to the Amniota (when its an ingroup) and includes mesosaurs with parareptiles, which we discussed earlier, among several other misfit nestings. There is no mention of Gephyrostegus and Cephalerpeton, which nest at the base of the Reptilia in the large reptile tree. And Longisquama is pictured gliding with paired plumes outstretched horizontally.

You get the picture. This is old school data.

References
Steyer S 2012. Earth Before the Dinosaurs. Indiana University Press. 182pp.