Hone and Benton 2007 – part 4
The first three parts of this 4-part deep dive
into Hone and Benton 2007 = HB 007 are linked here, here and here.
Hone and Benton 2007
attempted to un-discover the origin of pterosaurs from tiny tanystropheids (Fig 1) described in Peters 2000. HB 007 was written by professor and paleo textbook author, MJ Benton, and his student, DWE Hone, now a PhD. This was their first paper together.
Figure x. The lineage of pterosaurs recovered from the large reptile tree. Huehuecuetzpalli. Cosesaurus. Longisquama. MPUM 6009.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg?w=460″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg?w=460″ alt=”The lineage of pterosaurs recovered from the large reptile tree. Huehuecuetzpalli. Cosesaurus. Longisquama. MPUM 6009.” class=”wp-image-6903″ style=”aspect-ratio:0.45732038834951455″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg?w=460 460w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg?w=67 67w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg?w=135 135w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2012/08/huehuecuetzpalli-pterosaur.jpg 588w” sizes=”(max-width: 460px) 100vw, 460px” />
Here is part 4,
let’s start with a quote from HB 007: “If Peters had examined the specimens and had made no biomechanical assumptions, those seven characters (330–336) would be coded differently.
This is an ironic statement in that Peters 2000 examined all the pertinent (= previously omitted) tiny tanystropheid specimens. It was a requirement for publication. Meanwhile, Hone and Benton make no mention of examining ANY of the pertinent and other specimens.
This reminds me that Hone rejected my manuscript on the ‘First flightless pterosaur‘ because I had not looked at the specimen “long enough“. How did he know how much time I spent with the specimen? How long is long enough? This sounds like gatekeeping.
Hone also rejected by corrections on a manuscript updating Cosesaurus, Sharovipteryx and Longisquama because I “did not examine the specimens twice“. How did Hone know my itinerary? Evidently Hone did not know I had 8×10 transparencies in my files? This also sounds like gatekeeping given our history (= Hone and Benton 2007).
Back to the bipedal vs quadrupedal question.
HB 007 reported, “Cosesaurus is treated as a biped by Peters (2000) with characters coded based on this assumption.”
Not an assumption. Peters (2000a,b) matched the manus and pes of Cosesaurus (Fig 1) to Early to Mid-Triassic Rotodactylus tracks in which pedal digit 5 impressed as a small posterior point. This was achieved by flexion of the fifth pedal phalanges during extension of the medial four digits. Like Tanystropheus, Sharovipteryx and pterosaurs, the first phalanx of pedal digit 5 in Cosesaurus (Fig 6) was as large and robust as the medial metatarsals. This trait is not found in any other tetrapod clade. Rotodactylus tracks can be quadrupedal, bipedal and some show the transition between the two modes. See figures here.
HB 007 did not cite Peters 2000a, 2002, which further explored this topic five to seven years before HB 007.
“Peters (2000) argues that the Australian frilled lizard (Chlamydosaurus kingii) shares remarkably similar hind limbs and is capable of bipedalism. Chlamydosaurus is indeed capable of bipedalism at high speeds, but its primary mode of locomotion is as a sprawling quadruped.”
This is incorrect. Shine and Lambeck 1989 (cited in Peters 2000) reported that they observed Chlamydosaurus knigii spending much of its day bipedally, and the rest of its day in the trees.
“To consider Cosesaurus a true biped is misleading, as while it might have been capable of bipedalism, it is not a biped in the strict sense and should be considered quadrupedal.”
This is more quibbling, taking up page space while saying nothing. Even so, Let’s follow the HB 007 logic stream here: “Cosesaurus should be considered quadrupedal”, but that does NOT in their view make it related to Triassic pterosaurs, which were also, in their view, quadrupedal. That’s a match, not a mismatch. So, where’s the argument?
Note: HB 007 overlooked the obligate biped Sharovipteryx in this argument. Pterosaurs evolved from quadrupeds to bipeds during Cosesaurus, as indicated by its morphology and matching ichnology. This is the transitional taxon, ‘the Archaeopteryx of pterosaurs.’ As experts in evolution, Hone and Benton should understand the concept of a transitional taxon.
Moreover, Peters 2000 repeated what Shine and Lambeck 1989 reported on skeletal traits in extant lepidosaurs that indicate degrees of facultative bipedalism. The anterior process of the ilium framing four sacrals in Cosesaurus is one of those traits, overlooked by HB 007. Plus matching Rotodactylus tracks.
“Cosesaurus is also given a ‘centre of balance’ again without justification and well forward of the pelvis and these biomechanical assumptions colour the coding of hindlimb characters.”
Characters are either present or not present, longer than or not longer than, etc. I’ve never come acoss any analysis that was ‘coloured’. Clearly HB 007 had assumptions that did not pan out after testing. Many examples here (below) and previously.
These mistaken assumptions might have come from comparisons with basal theropod dinosaurs in which the deep chevron tail was made more massive with large caudofemoral muscles that counterbalanced the relatively short pre-pelvic area including the skull. Thus in non-avian theropods the center of balance is closer to the pelvis.
By contrast, and spelled out in Peters 2000, Cosesaurus, Sharovipteryx and pterosaurs had an attenuated tail lacking deep chevrons and thus lacking massive caudofemoral muscles. This lack of tail weight moved the center of balance anteriorly.
“Peters (2000) also suggests that the elongation of the metatarsals in both prolacertiforms and pterosaurs is an indication of digitigrade bipedality. Comparative diapsid analogues suggest that this is not the case. Lizards are mostly obligate quadrupeds and yet have elongated metatarsals and those that are capable of facultative bipedalism may have reduced metatarsals and elongate tarsals and phalanges compared to other species (e.g. Cnemidophorus tigris, Dipsosaurus dorsalis). Other digitigrade bipeds such as ornithischian and theropod dinosaurs do not always show lengthened metatarsals.”
More quibbling without presenting more parsimonious outgroup taxa. See Shine and Lambeck 1989 (cited in Peters 2000, but ignored by HB 007) for their list of traits that are present and barely present in living lizards capable of bipedal locomotion vs those that are incapable. I trusted their data in my freshman paper.
Figure 1. Sharovipteryx in situ. Click to enlarge. Here both plate and counter plate are shown along with a tracing based on both.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg?w=584″ alt=”Figure 1. Sharovipteryx in situ. Click to enlarge. Here both plate and counter plate are shown along with a tracing based on both.” class=”wp-image-17258″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg?w=143 143w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg?w=285 285w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2015/01/sharovipteryx-insitu588.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
Sharovipteryx issues raised by HB 007.
“A significant number of characters also appear to have been coded by Peters (2000) in the absence of physical evidence. Sharovipteryx has been coded for characters that are missing in the single specimen or are buried in the matrix and so cannot have been observed. Some examples of these are: the ‘lacrimal fails to meet nasal’ although this part of the skull is disarticulated, ‘twelve cervical vertebrae’ are coded although this section is obscured by matrix and the character ‘last few dorsals with fused ribs’ when some of the bones are disarticulated and the rest obscured. Furthermore, there are codings for three humerus characters, ‘loss of intermedium in carpus’, ‘ulna lacks olecranon and sigmoid notch’ and ‘manual asymmetry’ even though the arms are not preserved in the specimen.”
HB 007 did not report that they examined Sharovipteryx. So their claims are competing with those of Peters 2000 who did examine the plate and rarely photographed contemplate (Fig 2). If mistakes were made, which is possible in a freshman study, many of those mistakes were corrected in the manuscript of corrections rejected by Hone. That’s called gatekeeping.
Figure 1. Click to enlarge. The origin of the pterosaur wing and the migration of the pteroid and preaxial carpal. A. Sphenodon. B. Huehuecuetzpalli. C. Cosesaurus. D. Sharovipteryx. E. Longisquama. F-H. The Milan specimen MPUM 6009, a basal pterosaur.
” data-large-file=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg?w=584″ src=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg?w=584″ alt=”Figure 1. Click to enlarge. The origin of the pterosaur wing and the migration of the pteroid and preaxial carpal. A. Sphenodon. B. Huehuecuetzpalli. C. Cosesaurus. D. Sharovipteryx. E. Longisquama. F-H. The Milan specimen MPUM 6009, a basal pterosaur.” class=”wp-image-1415″ srcset=”https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg?w=584 584w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg?w=150 150w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg?w=300 300w, https://pterosaurheresies.wordpress.com/wp-content/uploads/2011/08/pterosaur-wing-origin1.jpg 588w” sizes=”(max-width: 584px) 100vw, 584px” />
“Peters (2000) reconstructed Sharovipteryx with highly reduced arms and he identified the three main bones (humerus, radius and ulna). He did not identify the carpus or hand and yet he was able to code some carpal characters. The bones identified by Peters as a forelimb, however, lie along a series of disarticulated and broken ribs and it is likely they are part of that series. Were this an arm, it would have had to have become detached from the pectoral girdle and moved down towards the sacrum as a single articulated piece, which is highly unlikely with such delicate bones. In any case, the true arms of Sharovipteryx have now been found buried in the matrix (R. Reisz, pers. comm., 2003) and this confirms that Peters’ (2000) supposed arm was incorrectly identified.”
Unfortunately, the Reisz data have not been made available and have not been published in the last 23 years. So what did Reisz uncover? Why is he waiting? False alarm? Meanwhile, Sharov identified a long manual digit 4 that extended back to the femur. That’s what I observed and traced. Nothing out of the ordinary (Fig 3). HB 007 prefers the rumor to the published data.
The forelimb elements I examined fell in line in all morphological aspects between the two sister taxa of Sharovipteryx, Cosesaurus and Longisquama (Figs 2, 3). These included a tiny pteroid and preaxial carpal, bones otherwise found only in fenestrasaurs, including pterosaurs. I also identified prepubic bones and a hyper-elongated ilium in Sharovipteryx. These traits are also restricted to fenestrasaurs including pterosaurs.
“Peters’ (2000) cladistic analyses do not follow normal practice. Outgroups are not defined, whether selected from the data, taken as all ‘0’s, or left unrooted.”
HB 007 needs to redirect this issue to Evans, Bennett and Jalil. I simply added previously omitted taxa to their earlier studies.
“Peters discusses his outgroups, stating that a number of prolacertiforms were used as pterosaurian outgroups within the study. One outgroup was apparently a species of Langobardisaurus, but since another species of that genus is included in the ingroup, it is hard to see how this can be considered an appropriate outgroup.”
Sounds like yet another quibble while avoiding the main issue: which taxa nest closer to pterosaurs: archosaurs or Cosesaurus and kin? HB 007 do not provide a solution to this overriding issue. Seems like they are not even looking for a solution, just another place to stick the knife in again and again. HB 007 do maintain their focus, but good science makes possible new hypotheses to replace old ones. HB 007 don’t want that.
“Bennett’s work (1996) would suggest that at the very least taxa from outside the prolacertiforms should be used.”
This comment is not logical. In Peters 2000 taxon lists from Evans, Jalil and Bennett were expanded to include omitted taxa.
“However, in the final trees, more basal taxa appear at the base of the trees, so it is unclear how these prolacertiforms were used as outgroups, if at all.”
Peters 2000 was a simple experiment: Add taxa to previously published studies. Plus a few insight from Chlamydosaurus. That’s all. Whenever HB 007 report something is ‘unclear’, remember, they had every opportunity to call or email for clarification.
“For his re-analysis of Evans’ work, Peters (2000) used only a heuristic algorithm (and may have done so on other analyses), which is not guaranteed to find the shortest trees available.”
Just getting my feet wet in this freshman attempt (without professor Benton’s help) at testing four previously omitted taxa in the realm of pterosaur ancestry.
Note: Hone does not present an algorithm that IS guaranteed to find the shortest tree available (given perfect scoring, of course). Is there such an algorithm? Or are they all affected by taphonomy and the human factor of bias, knowledge and judgement. Seems like more quibbling while avoiding the results. and competition for those results
“This ambiguity in coding and in cladistic method is reflected in the trees produced by Peters (2000), for example, the re-analysis of Jalil’s (1997) work produces two large polytomies with scattered taxa between them, including all the key taxa, hardly a resolution (Fig. 3).
Turns out, this is a quibble. Yes, there was loss of resolution with the addition of taxa using Jalil’s taxon list and character list. Even so, HB 007 do not report that pterosaurs nested closer to tanystropheids than to archosaurs in ALL the studies. That confession would have been against the Benton mandate to young Hone.
Readers: You’ll know when such an operation is in full swing because NO positive news comes out, and NO solutions are offered, only inconsequential quibbles.
“The choice of representative taxa is also problematic: Eudimorphodon is a poor choice as a basal pterosaur. Cladistic analysis shows it to be placed among the relatively advanced rhamphorhynchoids (Unwin 1995, 2003) and it is, therefore, not especially close to the base of the pterosaurian phylogeny. Its relative Dimorphodon is a better candidate, being more basal (Unwin 1995, 2003) and significantly more complete.”
Wish I knew about Unwin 2003 back in 1999 when I was writing the manuscript. Unwin 1995 was published as a preliminary attempt in an obscure Chinese journal. Sorry.
“Similarly, although Longisquama had been used as a putative outgroup for previous analyses (Sharov 1970) its revision as a neodiapsid (Unwin et al. 2000) was ignored by Peters and would at least suggest that it is not suitable for inclusion in analyses focused on pterosaurs and prolacertiforms.”
I knew about Sharov 1970 in 1999. I did not know about Unwin et al 2000 in 1999. For that matter, Unwin et al 2000 is not cited in HB 007. So, two unforced errors here.
The LRT recognizes no clade ‘Neodiapsida,’ but does recover two convergent appearances of the diapsid skull architecture in the Reptilia due to taxon inclusion.
In contrast to HB 007 Longisquama nested as the closest outgroup to Triassic pterosaurs and the same when corrections were made after more experience with taxa and examination. So Longisquama IS suitable for inclusion in pterosaur analyses, contra HB 007.
“In fact, its inclusion may result in the generation of spurious trees since the outgroups used in the analysis would be more derived than Longisquama as an ingroup taxon.
When HB 007 report, “may result” the reader can assume that HB 007 are only imagining something they chose not to test. Testing is what scientists do. Quibbling is easier and faster.
The results of the re-analyses of the supermatrix suggest that the Prolacertiformes should be considered the sister group to the Archosauria. The Pterosauria are not closely related to the Prolacertiformes and should instead remain among the Archosauria and probably among the derived archosaurs.”
Let’s avoid suprageneric taxa like ‘prolaceriformes’ since tanystropheids are no longer related to the Prolacerta and Protorosaurus in the LRT, where the sister group to the Archosauria is the Poposauria, far, far, far form Prolacerta and kin. Use generic taxa, specific taxa or museum numbered specimens, please.
“However, the large amount of missing data for many taxa makes it difficult to confirm their true position.”
That depends on related taxa that might be very complete. Once again HB 007 are imagining without actually doing the work = repeating the experiment and examining specimens.
If that’s how to get a PhD in paleontology, sign me up.
“Removal of hindlimb characters has no significant effect on their position and so cannot be used as an argument for convergence and non-homology between the dinosauromorphs and pterosaurs.”
The more taxa, the more characters, the better. More taxa is better than more characters after 200+ characters. Missing pertinent taxa skew results. So Peters 2000 added taxa. The LRT added more taxa, confirming Peters 2000, not Bennett 1996.
“Finally, there are such serious questions about Peters’ (2000) cladistic methods and about his original reconstructions and character codings in certain fossil material, especially for Longisquama and Sharovipteryx, that all his analyses should be treated with great caution.”
Yes, Peters 2000 included errors, many, perhaps all corrected, in Peters unpublished here. But Hone prevented the publication of those corrections. That’s how you know the fix is in.
If HB 007 had such serious questions about methods, they should have sent those serious questions to Evans, Jalil and Bennett. I simply added taxa to their published matrices, following their methods. Very simple.
Unfortunately for HB 007, their mission: to shut out the outsider, was too late, too little, and did not provide a better solution. Only quibbles.
All observations and analyses should be treated with great caution. This is science. Everything should be tested, not simply accepted on friendship nor rejected because the author is a freeloading outsider, skipping the traditional expensive, time-consuming academic process that Bennett and Hone went through and Peters leap-frogged for free.
I hope you find it as intriguing as I do that HB 007 did not chide Bennett for recovering pterosaurs close to croc-like Proterosuchus and giant, lumbering Erythrosuchus in the absence of the tiny bipedal croc, Scleromochlus (Fig 4). Alas, this is what happens when pertinent taxa are excluded. When outsiders point this out, no one in Academia is happy about it.
Quoting from the HB 007 Acknowledgements:
“We thank Dave Unwin, Chris Bennett, Olaf Bininda-Emonds, Mark Wilkinson and two anonymous referees for extremely helpful comments on earlier versions of the manuscript.”
Note that Bennett, author of Bennett 1996, was one of the two authors competing in HB 007 for a most-parsimonious hypothesis of pterosaur outgroups and origins. Do you find it surprising to see Bennett listed here? And Peters out of the loop?
Note that Peters, author of Peters 2000, was not invited to review the manuscript prior to publication, even though Peters 2000 was labeled ‘most importantly’ in the text. Evidently the authors did not want to hear my comments prior to publication. That’s unfortunate, because early input could have resolved some of the published biases, errors and oversights. So, here, here and here are my too-late-to-matter comments in this 4-part series.
Today’s lesson:
Reviewing and refereeing manuscripts prior to publication is a privilege reserved for PhDs, no matter their experience and expertise in the subject matter. That’s why independent researchers are ‘not considered seriously’. Independents don’t referee academic papers.
I know David Unwin. Among his many achievements he promoted the myth of the deep-chord wing membrane with his superficial ‘study’ of Sordes. Unwin did not identify the posteriorly and medially displaced left wing membrane. The right wing membrane was intact and articulated. It had a slender chord stretched between the elbow and wing tip (Peters 1995, 2002).
I know S Chris Bennett. Among his many achievements Bennett used statistics to separate Pteranodon into genders, not realizing nearly all differences in pterosaurs are phylogenetic due to their lepidosauromorpha ancestry (Fig 1). Bennett mistook a maxilla for a scleral ring on a small, flat-headed anurognathid and thus had to invent/imagine a skull morphology different from that of all other pterosaurs with an eyeball in the front half of the skull. Bennett recovered an untenable pterosaur origin close to Scleromochlus (Fig 4) and if not Scleromochlus, aquatic Proterosuchus and giant Erythrosuchus because he ignored tiny tanystropheids = taxon exclusion.
I had to look up Olaf Bininda-Emonds. Around 2007 his website indicates “Head of Heisenberg research group investigating the evolution of chemical communication, principally in carnivore mammals (Carnivora), and also applied aspects of chemical ecology in anuran amphibians and wasps; additional research into phylogenetic methodology, including supertree construction and phylogenomics.
And: Research into phylogenetic methodology, including supertree construction and the utility of embryological heterochrony data as a source of phylogenetic information; integration of conservation biology and phylogeny; investigating aquatic adaptation in mammalian carnivores.
And: Head of the bioinformatics research group within the Lehrstuhl für Tierzucht, as part of the BMBF funded BFAM project (Bioinformatics for the Analysis of Mammalian Genomes); research into genetic basis for BSE in cattle and energy metabolism in pigs; additional research into phylogenetic methodology, including supertree construction and the utility of embryological heterochrony data as a source of phylogenetic information.
So, nothing here re: archosaurs, pterosaurs or tanystropheids.
I also had to look up Mark Wilkinson. His present website indicates, “On weekdays (and some weekends) I am a researcher in the Department of Life Sciences at the Natural History Museum, (NHM) London. My research is divided between my primary interests in the theory and methods of phylogenetic and other evolutionary inference and the biology of caecilian amphibians (Gymnophiona). I am an evolutionary biologist and I am particularly interested in how we can infer phylogeny (historical genealogical relationships). I have worked on consensus and supertrees methods, compatibility methods, character construction, detecting rogue taxa, and measures of support.”
So, again, nothing here re: archosaurs, pterosaurs or tanystropheids.
After four posts, this paragraph concludes this deep dive
into Hone and Benton 2007, a 19-year-old paper that attempted to undiscover the origin of pterosaurs – not by introducing a set of taxa closer to the origin of pterosaurs – but by nitpicking errors (of their own making, according to Bennett 2012) in the Peters 2000 paper that introduced a set of four tiny tanystropheid taxa closer to the origin of pterosaurs (Fig 1) than had been proposed previously (Fig 4). With MJ Benton as co-author tHB 007 felt they had license to skip the scientific method, perhaps because the author of Peters 2000 was an amateur, an independent, someone who could be academically bullied seven years too late.
Now that you’ve had a peek beneath the curtain of the goings-on in paleontology, I hope your manuscript survives the submissions process – if warranted and fairly judged by experts.
Not sure how my several published manuscripts and abstracts got through back then, given all the academic gatekeeping. Ironically, even though I now have moree experience and understanding, the ability to publish is worse now due to quibbling referees like DWE Hone, and better now due to ResearchGate.net AND blogposts like this one.
Thank you for your readership.
I hope all these posts help you in your journey.
Face up to all challenges. Some are only quibbles.
Correct your mistakes as often as possible.
Imagination is good, but evidence is better.
References
Bennett SC 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoolological Journal of the Linnean Society 118: 261–308.
Bennett SC 2012. The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined. Historical Biology. iFirst article, 2012, 1–19.
Benton MJ 1985. Classification and phylogeny of the diapsid reptiles. Zoological Journal of the Linnean Society 84: 97–164.
Benton MJ 1999. Scleromochlus taylori and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society London, Series B 354 1423-1446.
Clark, J, Hopson J, Hernandez R, Fastovsky D and Montellano M 1998. Foot posture in a primitive pterosaur. Nature 391: 886-889.
Evans SE 1988. The early history and relationships of the Diapsida. Pp: 221–260 in: Benton MJ (ed) The phylogeny and classifaction of the tetrapods, volume 1: Amhiphians, Reptiles, Birds. The Systematics Association special volume 35A. Clarendon Press.
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.
Jalil N-E 1997. A new prolacertiform diapsid from the Triassic of North Africa and ther interrelationships of the Prolacertiformes. Journal of Vertebrate Paleontology 17ª3):506–525.
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in TetrapodsIchnos, 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.
Peters D 1995. Wing shape in pterosaurs. Nature 374, 315-316.
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.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29: 1327-1330.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.
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].
Source: https://pterosaurheresies.wordpress.com/2026/05/22/hone-and-benton-2007-part-4/
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