Updated human evolution video now extends back to Cambrian chordates

A human evolution video
listing and describing the ancestors of humans going back to Devonian tetrapods has been removed and updated to include recently added fish and chordates in our lineage going back to the Cambrian. Click to view this new 12:40 video on YouTube:

Gone are the more famous ‘transitional tetrapods’
Acanthostega and Ichthyostega. In their place are more direct transitional tetrapods, like Koilops and Trypanognathus. These taxa share more traits with their flat, small-lobed ancestors, Panderichthys and flat, small-limbed descendants, like Trimerorhachis, leading to frogs and reptiles… and ultimately mammals and humans.

Gone is the more famous ‘basal reptile’,
Hylonomus. In its place are the amphibian-like reptiles, Gephyrostegus and Silvanerpeton. The latter nests as the last common ancestors of all amniotes in the large reptile tree (LRT, 1691 taxa), the data source for the current list of human ancestors in the video.

Figure 1. From the Beginning - The Story of Human Evolution was published by Little Brown in 1991 and is now available as a FREE online PDF from DavidPetersStudio.com

Figure 1. From the Beginning – The Story of Human Evolution available as a FREE online PDF from DavidPetersStudio.com. Click here to view.

All of this interest
in evolution and human ancestry stems from research during the production of the book From the Beginning (Peters 1991). Back then it took 36 discrete steps from DNA to Homo to tell our story. While unprecedented for its time, that story can now be told more accurately with the addition of 50 more taxa based on knowledge gained in the last nine years while working on and constantly  improving ReptileEvolution.com.

Proviso: This nearly 30-year-old book includes both Ichthyostega and Hylonomus, so it is no longer up-to-date. That’s how science works, falsifying and building upon past hypotheses.


Dinosaurs Rediscovered, new book by Dr. Michael Benton

FIgure 1. Dinosaurs rediscovered by MJ Benton book cover.

FIgure 1. Dinosaurs rediscovered by MJ Benton book cover.

Dr. Stephen Brusatte wrote
on the cover: “If you want to know how we know what we know about dinosaurs, read this book!”

‘Amazon Customer’ wrote
at the book’s website, “Nice production, but highly biased and speculative.” (more below)

Dr. MJ Benton is professor of vertebrate paleontology and head of the Palaeobiology Research Group at the U of Bristol, England. He has written more than fifty books, including the standard textbooks in palaeontology.

From the intro:
“One by one the speculations about evolution, locomotion, feeding, growth, reproduction, physiology, and, finally, color have fallen to the drive of transformation. A new breed of dinosaur palaeobiologist replace the older ones, and they have applied a hard eye to the old speculations. Smart lateral thinking, new fossils, and new methods of computation have stormed the field.”

Funny though,
Scleromochlus (Fig. 2) is not mentioned. Benton 1999 promoted this genus close to the origin of pterosaurs and in the book he maintains that pterosaurs remain close to the origin of dinosaurs with no further explanation. Evidently Scleromochlus is no longer in favor. Nearly 20 years ago Peters 2000 invalidated the pterosaurs-close-to-dinosaurs = ornithodire hypothesis by testing Benton’s cladogram and three others by simply adding taxa overlooked and poorly scored by Benton and other prior authors. But let’s move on…

Figure 3. According to the AMNH, Scleromochlus is "one of the closest early cousins of pterosaurs." Oddly, they gave it the skull of Longisquama. Note the vestigial hands. These cannot elongate to become wings and pedal digit 5 is a vestige that cannot elongate to match basal pterosaurs.

Figure 2. According to the AMNH, Scleromochlus is “one of the closest early cousins of pterosaurs.” Oddly, they gave it the skull of Longisquama. Note the vestigial hands. These cannot elongate to become wings and pedal digit 5 is a vestige that cannot elongate to match basal pterosaurs.

Chapter 1 — Origin of the Dinosaurs
Even in 2019, Benton writes, “One thing is known for sure: the dinosaurs originated during the Triassic period, between 252 and 201 million years ago. Nearly everything else is uncertain.” This is not exactly a teaser, because it does not jive with what Benton writes earlier (Benton 1999) and later (see below).

Benton reports
that he raised traditional eyebrows back in 1983 when he suggested the old standard model of one group/clade giving way to another should be replaced with a scenario in which new clades only appeared and/or radiated after an extinction event. This view makes great sense and is supported by strong evidence. Ironically, Benton reports, “This new idea of mine was probably quite annoying for the established paleontologists.” Now that he’s older, the tables have turned and it’s Benton’s turn to be annoyed. Philosophically he has taken the place of his 1983 opponent and mentor, Dr. Alan Charig in that Benton now refuses to consider, test or replace invalid scenarios with new ones.

Let’s not forget…
in his unbiased youth, Benton 1985 used an early form of phylogenetic analysis to show that pterosaurs were sister taxa to lepidosaurs, closer to lizards than to dinosaurs by a long shot. Now that this hypothesis has become heterodox, he and others have avoided it ever since by selective deletion of pertinent taxa.

Figure 2. Cladogram from Benton 1985 in which he nests pterosaurs closer to lepidosaurs than to dinosaurs and other archosaurs.

Figure 2. Cladogram from Benton 1985 in which he nests pterosaurs closer to lepidosaurs than to dinosaurs and other archosaurs. Lots of confusion here due to taxon exclusion going back to the advent of Reptilia (= Amniota).

A subchapter follows
on the lepidosaur rhynchosaur, Hyperodapedon (Benton 1983), where Benton first published on taxa he was given to worth with, but made the phylogenetic mistake of lumping rhynchosaurs with archosauromorphs. This was due to taxon exclusion.

The next subchapter, “What was the first dinosaur?”
Benton correctly identifies one of the first dinosaurs as Herrerasaurus. That agrees with the large reptile tree (LRT, 1562 taxa) which uses the last common ancestor method for determining clade member inclusion.

Basal bipedal dinosauriformes, from Lagerpeton through Marasuchus, Lewisuchus, Asilisaurus, Sacisaurus and Silesaurus.

Figure 3. Basal bipedal ‘dinosauriformes’, from Lagerpeton through Marasuchus, Lewisuchus, Asilisaurus, Sacisaurus and Silesaurus, according to Nesbitt (2011). The LRT does not support this listing or sequence.

Then Benton reports on the poposaur
dinosaur-mimic, Silesaurus (Fig. 3), the Early Triassic ichnite Prorotodactylus, and another poposaur dinosaur-mimic, Asilisaurus (Fig. 3). Benton reports, “The discovery of Asilisaurus unequivocally re-dated the origin of dinosaurs back from 230 to 245 million years ago, or older.” There is little to differentiate Asilisaurus from Silesaurus. Both remain poposaurs and dinosaur-mimics, unrelated to the dinosaurs, except through basal bipedal crocodylomorphs, which Benton avoids. So, taxon exclusion strikes Benton, once again.

Quote here, an anonymous, but well-educated, review from Amazon.com:
“Dinosaurs Rediscovered is an engagingly written and highly personalized account of dinosaurs, generally covering the field’s perceived advances from 1980 to the present. The publisher Thames & Hudson did an outstanding job in producing the book, formatting, and in the selection of paper.

“The author notes that the field transformed from 1984 onwards by cladistic methods, and the resulting phylogenetic trees or cladograms have thus become the “basis” for evaluating evolutionary models and all things dinosaurian, including anatomical reconstructions, physiology, behaviour, etc. The work described is rather restricted, with most emphasis given to the University of Bristol’s vertebrate palaeontologists, often ignoring important discoveries from other groups, and regrettably ignoring most conflicting evidence. The most egregious is the complete omission of any discussion of the persisting problem of dinosaur/avian digital homology.

“Benton begins with the discovery (in his laboratory) of microsomes known as melanosomes from SEMs of fibers from the back of the small theropod Sinosauropteryx, that were described as “proto-feathers” back in 1998. However, there was never any evidence that the fibers had any feather affinity, and many who studied the specimens found an external coating of small tubercular scales above the layer of fibers —- since prepared away and lost! It is clear, however, that the fibers called proto-feathers or “dino-fuzz” were beneath the skin and therefore not feathers. Too, as South African palaeontologist Lingham-Soliar showed in several important papers (not cited) the structures called melanosomes cannot be interpreted from the scanning electron micrograph (p. 8) as being within the fibers. Speculation!

“Plate V shows a fuzzy Sinosauropteryx with a ring tail like that of a civit or ring-tailed cat!! Then there is an outlandish image of a reconstruction of the Jurassic urvogel Anchiornis (incorrectly called a troodontid, see Pei ref below), as a terrestrial animal; but the feathers emanating from the legs and feet would have been a hindrance in ground locomotion. New fossil images (Pei et al., 2017 AMNH Bull 411, 66 pp) show claws consistent with tree-trunk climbing, similar to those of other urvogels. Plate VI shows photos of a “dinosaur tail” in amber, but there is NO evidence it is from a dinosaur and is most likely an enantiornithine bird.

“The section on dinosaur evolution is straight forward, but laden with speculation, and given the massive convergence among various archosaur lineages during the Triassic it is difficult to have full faith in the interpretations; and authors from Cambridge and the British Museum have questioned the time-honored phylogeny (pp. 82-84).

“Most of the remainder of the book is a romp through the various dinosaurian groups, with comments on everything from brains and internal organs to behaviour. Archaeopteryx is depicted as an earth-bound runner (p. 112), with open wings (like no living avian cursor – e.g. capercallie, chicken, etc.), despite the fact that Manchester’s Derek Yalden showed conclusively that the urvogel’s claws were those of a trunk-climber, quite similar in structure to those of woodpeckers and climbing mammals.

“Benton notes (122) without reservation that Sinosauropteryx “was the first dinosaur to have its feather colour determined”—-and on page 123 he shows a feathered Caudipteryx with avian wing feathers and notes “it is clearly a theropod and not a bird” in contrast to numerous papers arguing that it is a secondarily flightless bird. If not, flight feathers, a perfection of aerodynamic engineering, would have to evolve in a non-flight context, a real stretch of biological thought!

“In chapter 5 “Jurassic Park” he seems ambivalent about reconstructing dinosaurs from ancient DNA, although most would agree that it is impossible. Certainly Mary Schweitzer’s supposed discovery of T. rex blood vessels and proteins has been firmly refuted. He comments on small genome size in birds and dinosaurs, but the studies conflated the two groups, and small genome size is to be found in flying animals: bats, pterosaurs and birds. Growth studies on dinosaurs are discussed but much of that has recently been brought into question. Allosaurus (188) and Tyrannosaurus, with no evidence, are shown with a feathered coat! Diplodocus (210) is shown with neck high in the air, a posture disputed by computer-generated imaging. Benton appears to favor the model of flight origin of Dial and Heers, but such a model requires a fully developed flight apparatus, and both putative dinosaurian ancestors of birds, urvogels, and even archosaurian antecedents, all lacked the pectoral architecture to enact this model. It just will not work. Much speculation!

“Finally, although there is no citation in the text, the monolithic bibliographic listing in the section on ‘Further Reading’ is alarming; it appears highly selected to bolster the Bristolian view of dinosaurs, while ignoring any contrary views, many of which are supported by solid scientific data. Most disturbingly, the discoveries by Chinese palaeontologists, especially those at Beijing’s Institute of Paleontology and Paleoanthropology, which in reality propelled the recent revolution in our knowledge of dinosaur/bird evolution is largely ignored.”

Dr. Benton’s new book gave us old, misguided and too often invalid information. In 2019 we know better how taxa are related to one another and Benton should have known better, too. Taxon exclusion (= phylogenetic context) seems to be his number one problem because his descriptions and illustrations of specimens are typically excellent. After messing up on his first paper (removing rhynchosaurs from rhynchocephalians), Benton’s reputation and output continue to be tarnished with his latest book and many of his recent papers all due to taxon exclusion. On the other hand, and in the present climate, Dr. Benton understands there is no consequence for ignoring new hypotheses. If only he could recall what it was like for him back in 1983, trying to promote his own new scenario to the establishment.

Those paleo professionals who wrote glowing reports
for this book should also have known better, but allegiance can sometimes trump good science. Author and paleontologist, Stephen Brusatte (quoted above) was a student at Bristol, where Benton teaches.

A wide gamut phylogenetic analysis based on specimens
is a necessary ingredient before, during and after any specimen description. It remains the one and only way to minimize taxon exclusion.

Benton MJ 1983. The Triassic reptile Hyperodapedon from Elgin, functional morphology and relationships. Philosophical Transactions of the Royal Society of London, Series B, 302, 605-717.
Benton MJ 1985. Classification and phylogeny of diapsid reptiles. Zoological Journal of the Linnean Society 84: 97-164.
Benton MJ 1999. Scleromochlus taylori and the origin of the pterosaurs. Philosophical Transactions of the Royal Society London, Series B 354 1423-1446. Online pdf
Benton MJ 2019. Dinosaurs rediscovered. Thames & Hudson.
Nesbitt SJ, Sidor CA, Irmis RB, Angielczyk KD, Smith RMH and Tsuji LMA 2010. Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira. Nature 464 (7285): 95–98. doi:10.1038/nature08718. PMID 20203608.
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods.  Ichnos 7:11-41.
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.

Outdated paleontology textbooks

Following in the wake of the fading paleo textbook I grew up with,
Vertebrate Paleontology (Carroll 1988), comes more recent editions from Professor Michael Benton (3rd edition 2005; 4th edition 2014, Fig. 1).

Figure 1. Vertebrate Paleontology by M. Benton.

Figure 1. Vertebrate Paleontology by M. Benton.

Prior reviews:
“The book is a main textbook for vertebrate palaeontology and aimed at students and anyone with an interest in the evolution of vertebrates. It meets its five aims and is excellent value.” 
(Proceedings of the Open University Geological Society, 1 April 2015)

From the Amazon.com website:
“This new edition reflects the international scope of vertebrate palaeontology, with a special focus on exciting new finds from China. A key aim is to explain the science. Gone are the days of guesswork. Young researchers use impressive new numerical and imaging methods to explore the tree of life, macroevolution, global change, and functional morphology.

“The fourth edition is completely revised. The cladistic framework is strengthened, and new functional and developmental spreads are added. Study aids include: key questions, research to be done, and recommendations of further reading and web sites.

“The book is designed for palaeontology courses in biology and geology departments. It is also aimed at enthusiasts who want to experience the flavour of how the research is done. The book is strongly phylogenetic, and this makes it a source of current data on vertebrate evolution.”

A review from the perspective of the large reptile tree
Unfortunately this volume invalidates itself by taxon exclusion at many nodes. Readers are better served at ReptileEvolution.com where taxa are included and tested, not just reported on.

Dr. Benton has been caught excluding taxa
in the past (e.g. Hone and Benton 2007, 2009Yang et al. 2018) to support his own outdated and invalidated hypotheses (like Scleromochlus, the bipedal croc with tiny hands as a sister to Bergamodactylus, the basal pterosaur with giant hands). His textbook presents several falsehoods about pterosaurs (e.g. open ventral pelvis, all were quadrupedal, origin from archosaurs). The first dichotomy splitting the Reptilia into Lepidosauromorpha and Archosauromorpha is not presented, leading to many mix-ups in derived taxa. Lacking is a wide gamut specimen-based phylogenetic analysis, like the large reptile tree, a modern requirement for every textbook on this subject in the present cladistic era. Rather, a number of smaller, more focused previously published studies are presented without review or criticism.

because the Benton volume is a physical book, it cannot keep up with the weekly and daily additions of online competitors, like www.ReptileEvolution.com is able to do. The Benton book, and all such textbooks, start to become outdated the moment the authors submit their final drafts to the publishers, weeks or months before their publication dates. It’s just the nature of publishing. It cannot be avoided due to this time lag.

Popular books make similar mistakes
Naish and Barrett 2016 wrote a dinosaur book, “Informed by the latest scientific research.” Sadly, no. This book is a journalistic compendium of prior studies, many of which were invalidated by taxon exclusion. As in most traditional studies, bipedal crocs are ignored in their cladograms dealing with the origin of dinos. These authors also considered tiny bipedal Scleromochlus ancestral to pterosaurs + Dinosauromorpha (p. 34), following Benton 1999. This hypothesis of relationships was invalidated by Peters 2000 who simply added taxa ignored by 4 prior authors, including Benton 1999. We can also be disappointed that these PhD authors bought into the bogus Yi qi styliform reconstruction as a bat-winged bird amalgam without either a critical analysis or a second thought of this one-of-a-kind mistake. The authors also supported the debunked origin of birds from theropods of descending size (pp. 184–185). Authors and editors should have checked for logic errors like the following from Naish and Barrett: “The fact that a microraptor specimen preserves a fish in its belly, indicates that they were also spending time on the ground.”

Just let that sink in if you don’t get it right away.

Parker 2015
reports on traditional mishandlings of the evolution of reptiles considered and criticized here at ReptileEvolution.com and PterosaurHeresies. As is typical in traditional paleontology, too often sister taxa in Parker 2015 do not document a gradual accumulation of derived traits. For instance the first dichotomy in the Parker topology splits Synapsids from Sauropsids. So no amphibian-like reptiles are recognized. The next dichotomy splits Eureptilia from Anapsida / Parareptilia. So pareiasaurs nest with mesosaurs. Parker considers the origin of ichthyosaurs and turtles, “uncertain.” A wide gamut cladogram testing all possibilities has no such problem Parker splits the invalid clade Sauria into Lepidosauromorpha and Archosauromorpha, then splits Sauropterygia and Lepidosauria, then lumps Pterosauria and Dinosauria together in the invalid clade Avemetatarsalia / Ornithodira. And with Avemetatarsalia we once again return to Benton 1999, which keeps surviving like a zombie.

As others have noted,
the present day is a ‘Golden Age’ in paleontology where discoveries are being posted weekly if not daily. Paper textbooks just cannot keep up with the latest hypotheses of relationships when compared to online studies and critiques that can pop up within hours of academic publication.

Benton would not have written editions 1 and 2
unless Carroll 1988 had become obsolete. Benton would not have written editions 3 and 4 if he didn’t think the earlier editions were failing to keep with our understanding of paleontology. Given the time it takes to produce, publish and distribute giant textbooks, It may be time for textbooks to go extinct and evolve into current online information.

Benton MJ 2005. Vertebrate Paleontology 3rd Edition PDF online Wiley-Blackwell 455 pp.
Benton MJ 2014. Vertebrate Paleontology 4th Edition Wiley-Blackwell 480 pp.
Carroll RL 1988. Vertebrate Paleontology and Evolution. WH Freeman and Co. New York.
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.
Naish D and Barrett P 2016. Dinosaurs. How they lived and evolved. Smithsonian Press.  online here.
Parker S (general editor) 2015. Evolution. The whole story. Firefly Books 576 pp.
Peters D 2000. A redescription of four prolacertiform genera and implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293-336


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.

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.

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.

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.

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



Once a children’s book, now an academic paper on PeerJ.

The children’s book,
GIANTS of Land, Sea & Air – Past & Present (Fig. 1 PDF) came out in 1986 and featured all of the largest animals to scale on dozens of pages alongside humans for scale.

Figure 1. The cover of Giants, the book that launched my adult interest in dinosaurs, pterosaurs and everything inbetween.

Figure 1. The cover of Giants, the book that launched my adult interest in dinosaurs, pterosaurs and everything inbetween.

The PeerJ online academic paper
(McClain et al. 2015) featured many of the same sea creatures and many more (Fig. 2) to scale with a human swimming alongside. Lacking are any of the terrestrial, aerial and prehistoric creatures you’ll find in GIANTS.

Figure 2. Poster turned on end illustrating the largest creatures in the sea to scale from McClain et al. 2015,

Figure 2. Poster turned on end illustrating the largest creatures in the sea to scale from McClain et al. 2015,

Sorry I didn’t see this online paper when it first came out a few years ago. 
It just came to my attention today. I suppose the inspiration was by convergence, but 1986 was long enough ago that a 10-year-old reader then would have been close to forty in 2015.

I just got a thumb’s down from the PeerJ voters for wondering if any of the authors were aware of the earlier children’s book. First bad review I’ve had! Or was I not supposed to mention it at PeerJ?

McClain et al. (15 co-authors) 2015. Sizing ocean giants: patterns of intraspecific size variation in marine megafauna. PeerJ 3:e715; DOI 10.7717/peerj.715

Dinosaur books

At one time
I wanted to write and illustrate a dinosaur book. I had an idea for one (Fig. 1) and was inspired by the writers and artists of the Dinosaur Renaissance. It took several years…

Figure 1. The cover of Giants, the book that launched my adult interest in dinosaurs, pterosaurs and everything inbetween.

Figure 1. The cover of Giants, the book that launched my adult interest in dinosaurs, pterosaurs and everything inbetween.

I got a contract to do my first book. That begat another and another. The shelf life was no more than one year for any of them. None went to second editions, though several had foreign versions. Reviews were good. Libraries stocked them. Book signings were fun, when there was advanced publicity. Every so often there was a big or small check in the mail. Now Amazon keeps them alive, if just barely. Reviews are still good…

I would not want to write and illustrate another dinosaur book. New discoveries make at least part of the text and part of the depiction of its subjects obsolete, sometimes before shelf life is over. The amount of data needed to be covered is staggering. More pages mean the price the book rises out of the ability to pay for many potential readers. With book publication, there are no ‘do-overs’ or ‘updates.’ What’s done is done. And then there are always the nagging typos. There’s a lot of work involved. And it has to be polished perfect. Editors, working for publishers, have their say. So do collaborators, if any. You have to put your life on hold to get the thing done by deadline. And when it’s done, it sits on a bookstore shelf, just one more Christmas or birthday present vying for the consumer’s eye.

It’s much better to post blogs
and nurture growing websites, like ReptileEvolution.com. These can be updated at will in one’s spare time. There are no paper or printing costs. No ships and trucks to distribute them. No bookstores to deal with. No deadlines. News can be reviewed within a day, while it’s still fresh. Everyone in the world has free access to your work. They can focus in on what they really like and ignore the rest at no cost. And one more thing (quoting Steve Jobs) that books can’t provide: animation. There’s no profit in web publishing, but money was never the front and center issue.

Figure 2. Sample animation you’ll never see in a book. The Vienna specimen of Pterodactylus (wings folded). Animation opens the wings and legs to reveal the true shape of pterosaur wings, stretched between the elbow and wingtip with a short fuselage fillet extending from elbow to mid femur.

Even so
I’m glad I went through that book phase. It had its time and place. The process led me to interact with others of like interest. Some of them are PhDs. Others are fellow artists and writers. Everyone should have a hobby to keep in touch with the world and vice versa.

I was inspired to write this blog post
after seeing parts of Walking with Dinosaurs 3D on YouTube. Click here to see it. So much talent and effort went into this— truly outstanding visuals …but the dialog was horrible, as most others agree. And there are a few new dinosaur books out now, updating older dino books. I wish them all well. Someday, perhaps decades from now, those books will either be considered cherished classics or outdated, ready to be updated. It’s all good.


A Dinosaur Year 1989 Calendar

This ‘blast from the past’ by request: 
Click here or on image to download all 13 lorez images from my 1989 Dinosaur Year calendar, published by Alfred A. Knopf, New York. Thanks for the request, Leo!

I see two copies are presently available from Amazon.com here.

Click to download PDF of cover + 12 months of 1989 Dinosaur Year Calendar pix by David Peters at 72 dpi. It's over 25 years old and you'll find mistakes here. It was a product of its time.

Click to download PDF of cover + 12 months of 1989 Dinosaur Year Calendar pix by David Peters at 72 dpi. It’s over 25 years old and you’ll find mistakes here. It was a product of its time.

The calendar is over 25 years old
and you’ll find mistakes galore. It was a product of its time and the first time I ever painted dinosaurs in settings.

This followed
the book GIANTS and A Gallery of Dinosaurs, which illustrated dinosaurs on white backgrounds, all to the same scale. Both books are available as pdf files here and as used books at several online sites.

Where are the originals?
Collectors purchased all the originals except for the Brachiosaurus family in a pond (December) because it has a razor knife cut in the sky over the mountain top, inflicted upon opening the package at the publisher. It’s hanging on the wall over my monitor as I type this and I never notice the slit.

But wait! There’s more!
Click here to connect to a FREE build-it-yourself paper Pteranodon model.
And click here to connect to a FREE build-it-yourself paper Thalassomedon model.
All you need is 8.5×11″ bristol (stiff) paper, some glue or tape and a scissors or knife. Have fun, kids!

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.

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.

The Dinosaur Heresies NYTimes Book Review from 1986

the_dinosaur_heresies200Now almost 30 years old, here’s something you might like to read (perhaps again?).
This is the NY Times book review of Dr. Robert Bakker’s ‘The Dinosaur Heresies’ from 1986. You can read the complete original here. I went to the prophesies below and marked them with a [+] or a [-] for those supported today or not and for those that are still questionable: [?].

Dinosaur Mysteries
Published: November 8, 1986

THE DINOSAUR HERESIES. New Theories Unlocking the Mystery of the Dinosuars and Their Extinction. By Robert T. Bakker. Illustrated. 481 pages. William Morrow & Company. $19.95.

Mr. [not Dr.?] Bakker has a quirky, free-floating imagination, and in the course of this book – which is generously illustrated with his own charming sketches – he raises many offbeat questions: Were changes in dinosaur eating patterns responsible for the evolution of flowering plants? [+] Did pink pterodactyls exist? [?] What sort of lips did dinosaurs have? [+] Could a human being beat a tyrannosaurus at arm wrestling? [?]

Mr. Bakker, the adjunct curator at the University Museum in Boulder, Colo., has published many papers in the field of vertebrate paleontology, and his book stands as an informative layman’s introduction to the wonderful world of dinosaurs while at the same time making an impassioned case for his own – sometimes heretical – views on their endurance and extinction. ”I’d be disappointed,” he writes, ”if this book didn’t make some people angry”; and given the often fiercely polarized world of vertebrate paleontology, he’s unlikely to be let down.

As Mr. Bakker sees it, dinosaurs have been given a bad rap over the years as ”failures in the evolutionary test of time” – portrayed as small-brained, cold-blooded sluggards who couldn’t ”cope with competition from the smaller, smarter, livelier mammals.” Such portraits, he suggests, are unfair as well as scientifically inaccurate: in the first place, dinosaurs dominated history for 130 million years [+] – a remarkably long period of time that attests to a decided ability to survive (the human species, in contrast, has only been around for 100,000 years). And while Mr. Bakker acknowledges that dinosaurs were probably not brilliant thinkers [+], he makes a persuasive argument for their physiological adaptability and their prodigious energy [+] – he even speculates that tyrannosaurus could gallop about at speeds approaching 45 miles an hour.  [-] 

Much of ”The Dinosaur Heresies,” in fact, revolves around the question of whether the animals were cold-blooded (and more closely related to reptiles) or, as Mr. Bakker contends, warm-blooded (and more closely related to mammals and birds) [+]. While he occasionally stops to summarize opposing viewpoints, he is less interested in presenting an objective overview of the field than in mustering evidence to support his own theories.

He argues that gizzards and large digestive tracts in [some] dinosaurs would have compensated for their weak teeth [+], enabling them to eat high quantities of land plants, necessary to support a high metabolic rate. He argues that birds and pterodactyls – both of which would have had to evolve high-pressure hearts and lungs before flight could have been achieved  [+] – descended from dinosaurs  [+] [-], and that it’s not unlikely that these ancestor dinosaurs were already equipped for high metabolism [+]. He argues that the dinosaurs’ ”adaptations for sex and intimidation” – horns, head-butting armor and all manner of bony frills -suggest that they led active, aggressive lives, uncharacteristic of lethargic, cold-blooded animals  [+]. He argues that the growth rate of dinosaurs more closely resembles that of mammals than reptiles [+]. And, finally, he argues that dinosaurs’ porous bone tissue indicates the sort of high blood-flow rate usually associated with warm-blooded creatures [?].

On the question of the dinosaurs’ demise, Mr. Bakker sides with those paleontologists who discount new theories of mass extinction caused by some sort of cosmic catastrophe – he cites evidence suggesting the extinctions occurred not during a single ”doomsday” period but over tens of thousands of years [+] [-] [?]. In his view, the development of new sorts of dinosaurs and other animals, combined with changes in the physical and genetic environment, gradually led to their doom [+] [-].

On a side note:
I liked Dr. Bakker’s quote about making some people angry with his novel ideas based on overlooked data.

On another side note:
like our antiquated notions about dinosaurs from over 30 years ago, pterosaurs today have been given a bad rap. They are still portrayed as ungainly quadrupeds, bound by membranes that tied their legs together and tied their wings to their ankles (along with a long list of other false paradigms). The data deniers, unfortunately, are still out there, thinking that if they just turn a blind eye toward certain data and hypotheses they will go away.

As everyone knows,
this blog, Pterosaur Heresies, was intended to approach data with the same verve and testing of false traditions that Dr. Bakker demonstrated.



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.

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].