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Hone and Benton 2007 revisited – part 3

Thursday, May 21, 2026 16:57

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An earlier post was part one of a deep dive
into Hone and Benton 2007 (= HB 007). the paper that attempted to un-discover the origin of pterosaurs in Peters 2000b. 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.

For Part 2 click here.

Here is part 3.
Let’s start with a quote from HB 007: “The unexpected results produced by Peters (2000) and the reanalysis here, suggest that his codings should be examined more closely, particularly with reference to the pterosaurs.”

Here “unexpected” = novel, overlooked. ignored. That’s because previous to Peters (1997, 2000, 2002, 2007) pterosaur workers openly admitted they had no idea 1. how pterosaurs evolved from nonviolent ancestors, 2. how wings developed. That’s because workers omitted and ignored pertinent outgroup taxa. They did not realize basal pterosaurs were tiny bipedal tanystropheids and wings developed last (Peters 2002, not cited by HB 007).

Sadly, many pterosaur workers still believe that pterosaurs appeared without precedent, largely because of HB 007 and the widely available paleo textbook by professor MJ Benton. Academics have had the last 26 years to come up with a better = more parsimonious match for Triassic pterosaurs. Click here for their recent misguided = taxon omitting attempts at elevating lagerpetids to a pterosaur interrelationship.

As reported in part 2,
HB 007 paid little to no attention to results in Peters 2000b. They ignored Peters 2000a and 2002. Instead HB 007 sought flaws in the data. As reported in part 2, Bennett 2012 tested HB 007 and reported several typos were responsible for issues raised by the authors.

Here ‘unexpected’ = novel. That’s because previous to 1997, 2000, 2002, 2007 pterosaur workers admitted they had no idea where pterosaurs evolved from, how wings developed, etc. That’s because they were omitting and ignoring pertinent outgroup taxa.

Simply by expanding the taxon list Peters 1997 and 2000 recovered small bipedal tanystropheids, including Langobardisaurus, Cosesaurus, Sharovipteryx and Longisquama. More taxa have been added since then in the large reptile tree (LRT, 2340 taxa) and large pterosaur tree (LPT, 268 taxa).

“There are numerous methodological errors throughout the paper as well as errors in the interpretations of some specimens and the resulting codings.”

Bennett 2012 reported the errors were committed by HB 007, who included no images from Peters 2000 to compare with ‘correct’ figures. The authors looked at no specimens. NONE.

“In summary, Peters (2000) includes and excludes characters without explanation, he biases his character codings by unwarranted functional assumptions about some taxa, he includes character codings for bones that do not exist in certain specimens and there are methodological problems in the cladistic analysis. These problems are outlined below.”

HB 007 do not tell us whether or not those ‘certain specimens’ were in the original copied matrices and taxa produced by others. HB007 were vague on this matter. The ‘unwarranted functional assumptions’ are explained elsewhere in HB 007 as ‘assuming bipedalism.’

Given that Sharovipteryx was an obvious obligate biped and Cosesaurus (Fig 1) was matched precisely to Rotodactylus tracks (Fig 6), some of which transition to bipedalism, and given that flapping can only succeed after bipedalism, as in birds and bats = inverted bipeds, bipedalism is not a problem but a supporting trait with analogs in birds.

Note that HB 007 make no mention of the bipedal configuration of Scleromochlus (Fig 1) the basal bipedal crocodylomorph taxon with dorsal scutes preferred by co-author and professor MJ Benton (1999) and SC Bennett (1996) as pterosaur ancestor. That worked well enough for several publications prior to Peters 1997, 2000. In that year other taxa were described that shared MORE pterosaur traits. The search is always on for even better matches.

That’s how it works. HB 007 resisted that part of the scientific method. Instead they engaged in quibbling with mistakes of their own making (according to Bennett 2012).

Figure 1. Bipeds of the Triassic. Top to bottom: Cosesaurus, Scleromochlus, Marasuchus and Tropidosuchus. Each represents a distinct lineage of bipeds with bipedal sister taxa. Lagerpetids, recently promoted as pterosaur relatives, are most closely related to Tropidosauchus in all analyses that include both taxa.

HB 007 continue to quibble without comparing results.
“Peters (2000) deleted over two-thirds of the characters from Evans (1988), but retained many identical characters in the other analyses (e.g. his characters 6, 52, 60, 136, 142,182, 184, 245, 287): if a character holds for one analysis, thenit should be maintained for the others.”

Repeated from HB 007 revisited – part 2:
“As stated in Peters 2000, “Evans cladistically analyzed primitive diapsids, considered the Archoauria a single taxon and did not include the Pterosauria, Langobardisaurus, Longisquama or Sharovipteryx. For this paper I duplicated Evans’s data matrix relating to early diapsids and archosauromorphs (97 characters, 31 taxa, Evans’s Nodes A-H). Other characters relating to lepidosauromorphs were examined but not included because pterosaurs do not exhibit any lepiidosauromorph synapomorphies (Bennett 1996a). Benton (1985 noted two lepidosauromoph characters: 1) single ossified sternum and 2) specialized sternal attachment for the ribs. However, the sternum in pterosaurs is a fusion of other pectoral elements (Wild 1993), including a homologous sternum that is plesiomorphic. The ‘specialized sternal attachments’ are simply ossified sternal ribs which are plesiomorphic (Bennett 1996a).”

Repeating from part 2: Boy, was I wrong to believe Bennett 1996a in my freshman paper, because later studies (Peters 2007, the LRT) nested pterosaurs and tanystropheids within the Lepidosauromorpha. Back then, everyone agreed that pterosaurs were Archosauromorphs.

“Peters (2000) gave no justification for the extensive pruning of the work of Evans (1988), despite having deleted more than 150 characters out of 226 in all.”

See red type above. Repeating a false accusation does not increase validity.

“Although there are numerous characters and taxa that are irrelevant to the subject matter (and therefore can be justifiably excluded), many more that were removed could have aided the resolution of the cladograms.”

As noted in part 2, high resolution was not the goal, especially while adding to the matrices of others with their characters.

The goal of Peters 2000 was to find out which tested clades the Triassic pterosaur, Eudimorphodon (Fig 2), was attracted to given new opportunities for that attraction to happen by simply adding previously omitted and ignored taxa.

HB 007 apparently did not get this reason. Or if they did get this reason, and worked to undermine it by quibbling over mistakes of their own making (Bennett 2012), while avoiding presentation of their own results, then paleo has bigger problems than widespread and ‘acceptable’ = traditional taxon exclusion.

Readers should worry whenever the vague phrase ‘could have aided‘ are written by anyone – let alone freshmen authors guided and influenced by professors defending their own mistaken hypotheses. As scientists who follow the scientific method, HB 007 should have repeated the observations and analyses to the T rather than their attempt to second-guess their pterosaur expert peers who refereed Peters 2000 seven years earlier and found the manuscript important enough and error-free to publish in an academic journal.

Don’t rely only on citations. They might be out-dated. Do your own research.

Figure 2. The origins of Dimorphodon and Eudimorphodon find a common ancestor close to Austriadactylus (the Italian specimen) and prior to that, the basal pterosaur, MPUM 6009. All are Late Triassic except Early Jurassic Dimorphodon. The robust skull of eudimorphodontids suggests piscatory (fish eating) while the fragile skulls of dimorphodontids suggest insectivory. The enlarged naris was a legacy trait inherited from the Italian specimen of Austriadactylus.

HB 007 continue,
“A number of Peters’ character recodings are based on unjustified assumptions and reconstructions. Characters for Eudimorphodon are coded as if it were a digitigrade biped capable of both walking and running. This reconstruction of pterosaurs is discounted by many and contradicts trackway evidence (Mazin et al. 2003), the distribution of mass and the issues of balance (Pennycuick 1988) and the structure of the metatarsal/tarsal joints (Clark et al. 1998).”

The bipedal vs quadrupedal subject brought up by HB 007 was covered recently in the Laueropterus post. So some repetition here is inevitable.

Let’s talk about digitgrady and bipedalism in Triassic pterosaurs
Fig 2), ignoring for the moment that everyone reconstructs Scleromochlus (Fig 1), the sole competing taxon (according to all prior workers, as a digitigrade biped.

“The reconstruction includes a ‘hypothetical’ centre of balance, but no justification is given for its location and this was significantly anterior to the pelvis, which would cause the animal to fall forwards. If Peters had examined the specimens and had made no biomechanical assumptions, those seven characters (330–336) would be coded differently. Furthermore, in his four analyses, Peters (2000) codes certain identical data cells differently in each of his analyses (e.g. characters: ‘shape of maxillary ramus of premaxilla’, ‘nasals taper anteromedially’, ‘nasals longer than the frontals’, ‘subtemporal process of jugal’, ‘transverse processes of trunk vertebrae well developed’, and ‘relative lengths of metacarpals 3 and 4’). These different codings of the same characters are not explained.”

If so, these are quibbles.
If so, I am surprised that Evans, Bennett and Jalil had identical data cells, especially since they covered different, barely overlapping taxa.
If so, only four or five taxa (those added by Peters 2000) would have been affected. I had no interest or inclination in double-checking the published works of other more experienced paleontologists in my freshman paper. I just wanted to add taxa.
If so, these might be among the typos created by HB 007 discovered by Bennett 2012.

Figure 3. Animation of Pterodactylus matched to Crayssac tracks.

Figure 4. Click to Walking pterosaur according to Bennett. Note the forelimbs provide no forward thrust, but merely act as props. And they bear weight in this configuration. No matching to Crayssac tracks are attempted in this competing animation.

Animations
Peters matched Craysaac pterosaur ichnites to Pterodactylus, resulting in an upright configuration using the forelimbs as ski-poles to aid steadiness (Fig 3). This mode permits bipedal take-off by simply raising the wings and flapping, as in birds. When standing still the toes remain beneath the center of balance, as in birds.

By contrast, Bennett produced a series of illustrations animated here (Fig 4), that are acceptable to current academics in which the pterosaur is bent over, using its wings to support itself, and at odds with Triassic pterosaurs, pre-pterosaurs (Fig 1) and Scleromochlus (Fig 1) that don’t have Pterodactylus proportions (Figs 3, 4).

Plus, manual digit 3 is typically pointed backwards in ichnites. Not good for forward propulsion. Not bad for a ski pole. Different from all other retrapods. By this evidence many pterosaurs were secondarily quadrupedal. That’s how the manus gets ‘twisted’ (Peters 2002).

Pre-pterosaurs
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.

HB 007 did not cite Peters 2000a, 2002, which further explored this topic five to seven years before HB 007.

Back to HB 007 quotes:
“Characters for Eudimorphodon are coded as if it were a digitigrade biped capable of both walking and running. This reconstruction of pterosaurs is discounted by many and contradicts trackway evidence (Mazin et al. 2003), the distribution of mass and the issues of balance (Pennycuick 1988) and the structure of the metatarsal/tarsal joints (Clark et al. 1998).”

Hone and Benton omitted citing several sources here. Padian1983 reconstructed Dimorphodon (Fig 2) as a digitigrade biped. Peters 2000b, 2002 showed that some pterosaurs were plantigrade, others were digitigrade. Basal non-volent precursors were speedy digitigrade facultative bipeds (Cosesaurus, Fig 1) to speedy obligate bipeds (Sharovipteryx).

Furthermore the only known specimen of Eudirmorphodon (Fig 2) lacks feet and legs below the knees.

Figure 5. Dimorphodon macronyx and D. weintraubi, an anurognathid larger than its namesake.

With regard to the ‘structure of the metatarsal/tarsal joints‘ (Clark et al 1998) Hone and Benton made a mistake. The joint in question is a butt-joint between the metatarsals and proximal phalanges. This is distinct from most tetrapods. Clark et al (in a Nature cover story) maintained that butt-joint required all pterosaur feet, like their 3D example, Dimorphodon weintraubi (Fig 5), to be stiff and plantigrade.

Figuure 6. Cosesaurus matched to Rotodactylus from Peters 2000b.

By contrast, Peters 2000a showed the opposite. The proximal phalanges were also elevated in Dimorphodon weintraubi (Fig 7) and similar dimorphodontids – and Cosesaurus (Fig 6) – and those feet matched Rotodactylus and Sauria aberrante tracks (Fig 7) that did not impress the proximal phalanges.

Figure 7. A pterosaur pes belonging to a large anurognathid, “Dimorphodon weintraubi,” alongside three digitigrade anurognathid tracks and a graphic representation of the phalanges within the Sauria aberrante track.in

HB 007 wrote:
“The reconstruction includes a ‘hypothetical’ centre of balance, but no justification is given for its location and this was significantly anterior to the pelvis, which would cause the animal to fall forwards.“

Not so. In flying ‘things’, like airplanes, birds, bats, insects and pterosaurs, the center of balance must be near the chord of the wing. In pterosaurs the shoulder joint is near this marker. As in walking birds, walking pterosaur toes extended just beyond this point so the center of balance was above the central phalanges (Fig 2). Perhaps I thought this was common knowledge, but then this was the first paper on pterosaurs either Hone or Benton did and neither attempted a comparative reconstruction, but relied on out-dated and invalid citations.

Be wary and watchful if you are driven by an agenda. You’ll overlook pertinent data and someday you’ll be called to account on it.

That’s enough for now.
Part 4 is next. IMHO the worst is yet to come.

Lesson for today – don’t judge all paleontologists on the mistakes, oversights and biases of DWE Hone and MJ Benton.

It would have been much better science if these two academics had published a better outgroup for pterosaurs (= taxa that share more traits with pterosaurs), than those proposed by Peters 2000a, b, 2002. After all, that’s what Peters did and that’s their job: to come up wth solutions and novel insights – not to fling mud or raise quibbles at a peer-reviewed and published paper that shed new light on pterosaur origins with four taxa tested 4 times in analyses originally created by others and approved by other referees and editors.

Worse yet, Hone and Benton did not visit, examine and deiciipher ANY of the four tanystropheids under question. If they did so, they didn’t mention it in HB 007.

HB 007 have had 13 years, since Bennett 2012, to either confirm, modify or reject his findings of typos in their matrices. Evidently they stand firm.

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 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.
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.
Sanz JL and López-Martinez N 1984. The prolacertid lepidosaurian Cosesaurus aviceps Ellenberger & Villalta, a claimed ‘protoavian’ from the Middle Triassic of Spain. Géobios 17: 747-753.
Villaseñor-Amador D, Suárez NX and Crus JA 2021. Bipedalism in Mexican Albian lizard (Squamata) abd the locomotion type in other Cretaceous lizards. Journal of South American Earth Sciences 109:103299.

https://reptileevolution.com/pterosaur-wings.htm

Source: https://pterosaurheresies.wordpress.com/2026/05/22/hone-and-benton-2007-revisited-part-3/

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