Hone and Benton 2007 – revisited

Yesterday’s post
included some fact checking on Hone and Benton’s 2007 first paper as co-authors. Back then DWE Hone was an up-and-coming student of professor MJ Benton. Today Hone is a PhD.

A few years earlier
Benton (1999) wrote a paper supporting the traditional hypothesis that Scleromochluus (with tiny hands, dorsal osteoderms and lacking a pedal digit 5) was the ancestor to pterosaurs AND dinosaurs. A year later an amateur, David Peters (2000), added four previously overlooked taxa to three previously published analyses and recovered cladograms that showed a closer affinity of pterosaurs with these four tiny tanystropheid taxa, Langobardisaurus, Cosesaurus, Sharovipteryx and Longisquama.

Later work by Peters (the online large reptile tree (LRT, 2340 taxa) cemented those interrelationships as it tested all possible competing taxa and clades vying for their place in the ancestry of pterosaurs.

Today’s 2026 post dives into Hone and Benton 2007 = HB 007
breaking down some, perhaps all of the hypotheses then (and now) taught in college textbooks. Professor MJ Benton is also a college textbook author in paleontology. Today’s post also benefits from almost twenty years of hindsight, recent discoveries and new hypotheses of interrelationships.

The title of the HB 007 paper is wrong
but we didn’t know that then. Now we know pterosaurs are not archosauromorphs. They are Lepidosauromorphs in the LRT (Peters 2007). Notice the timing. Same year: 2007.

From the HB 007 introduction
“some recent publications (Evans 1988; Unwin 1995; Bennett 1996; Benton & Allen 1997; Jalil 1997; Dilkes 1998; Peters 2000) have shown that prolacertiforms and pterosaurs are the groups that have the most variable positions in phylogenetic analyses of Diapsida.”

This statement is due to the omission of the correct outgroup taxa from all prior analyses, except Peters 2000, who filled that vacuum.

HB 007 and none of the cited authors were aware that the diapsid skull architecture was convergent in Lepidosaur and Archosauromorph taxa, as recovered later by the LRT.

The LRT also split traditional Prolacerta and kin (archosauromorph diapsids) from tanystropheids (lepidosaurs).

“The prolacertiforms are the sister group of Archosauria (sensu Benton 1985) and there was a consensus that they form a clade (Evans 1988; Bennett 1996; Benton & Allen 1997; Jalil 1997; Peters 2000). Some recent analyses, however, indicate that prolacertiforms are paraphyletic. Both Dilkes (1998) and Modesto & Sues (2004, using a modified version of Dilkes’ dataset) showed that Prolacerta did not belong to the prolacertiforms (although the rest of the clade was retained).”

Interrelationships are different now in the LRT. Prolacerta, Protorosaurus and kin are basal to similar basal members of the Archosauriformes, like Proterosuchus and Euparkeria. There are many taxa between Euparkeria and the Archosauria (= crocs + dinos only).

Note the term ‘consensus’. That means HB 007 did not test this problem on their own, but relied on past authors and went along with the crowd.

Most importantly, HB 007 did not go out and visit/examine/trace/reconstruct ANY pertinent specimens. None at all. Neither were both authors pterosaur ‘experts’ prior to this paper, HB 007. “not experts’ means neither had published papers on pterosaur anatomy. or systematics before HB 007.

“Prolacerta did not belong to the prolacertiforms“. See how mangled phylogeny can get? Someone in Academia needs to call out this leap of logic. I can’t do that due to my amateur status.

“Senter (2004) also found the prolacertiforms to be paraphyletic, although here he removed the drepanosaurs to form a new clade with Longisquama.”

Adding taxa, as in the LRT, resolves this problem by minimizing taxon exclusion. Drepanosaurs led by Jesairosaurus are lepidosaurs, the sister clade to tanystropheids (including Longisquama + pterosaurs) let by Huehuecuetzpalli in the LRT.

“Most importantly, Peters (2000) has presented a heterodox view, in which the prolacertiforms are allied with the pterosaurs (see below for details). It is true that prolacertiforms have widely divergent body plans (Fig. 1), but they do appear to share a number of apomorphies. We use the term ‘prolacertiforms’ here to indicate the wider clade that includes Protorosaurus, Macrocnemus, Tanystropheus and their relatives (as in Evans 1988) and the term ‘protorosaurs’ to refer to the clade consisting of Protorosaurus, Macrocnemus, Tanystropheus and relatives, but excluding Prolacerta, as recovered by Dilkes (1998).”

“Most importantly”… hmm. Nice to hear after SC Bennett told me my work would ‘not be taken seriously’. HB 007 seems to have taken my work seriously with that preface.

BTW… Peters 1997 first advanced the hypothesis that Sharovipteryx was a proximal ancestor to pterosaurs. So this hypothesis had been brewing. More so after Longisquama and Sharovipteryx came to my home town, St. Louis, during the Russian Dinosaur Exposition. Then even more so after a visit to Barcelona to examine Cosesaurus and meet with the professor who described it in detail, P Ellenberger.

Back in the day, as HB 007 noted, tanystropheids were mixed in with protorosaurs. Many years later the LRT split them apart simply by adding taxa.

(Fig. 1)‘the widely divergent body plans”illustrated by HB 007 were nothing more than freehand silhouette tracings of in vivo (not in situ) specimens drawn by various artists. Not one bone was illustrated. Again, these two authors did not visit ANY specimens. NONE. Both the editors and referees thought that was OK. Seems like a double-standard and a wink-wink.

Note how HB 007 announce they “use the term” to indicate the same clade (sans one taxon) twice, referencing two papers, two authors. They don’t test either definition. In any case, the LRT recovers tanystropheids as far apart as possible from Protorosaurus + Prolacerta.

I am also to blame. Long before the LRT, when I was producing Peters 2000, I followed suite and considered all these taxa members of the Archosauromorpha. Only by 2007 did the lepidosaur hypothesis come out after analysis.

“The Pterosauria has been a notoriously difficult clade to place in the diapsid tree: pterosaurs appear suddenly in the fossil record and in full possession of all their highly derived characters.”

Not true. HB 007 were writing their paper on one published seven years earlier: Peters 2000 who showed a gradual evolution of pterosaur traits beginning with Langobardisaurus, then Cosesaurus, then Sharovipteryx, then Longisquama. HB 007 ignored that part of the paper.

So why did HB 007 make this statement? Why did the referees allow it?

After all, THIS is the subject of HB 007. They knew better. Perhaps the authors were having a hard time admitting to themselves that other academics had been looking in the wrong place. Peters (1997, 2000a, b, 2002, 2007) found the right place. I can see where that could be upsetting to their world view and business strategy.

“Pterosaurs have been allied to virtually every basal and crown-group archosaur clade as well as to the dinosaurs, but few characters can be found that unite them the Prolacertiformes.”

Read this carefully. HB 007 are ignoring the four tiny tanystropheid subjects of Peters 2000: Langobardisaurus, Cosesaurus, Sharovipteryx and Longisquama because these taxa are not in their list of prolacertiformes or protorosaurs spelled out above. That’s a trick. Beware of their tricks. They are avoiding the long list of increasingly pterosaur-like characters demonstrated by the four tiny tanystropheidae taxa listed in Peters 2000. They were hoping no one would notice.

“However, the trees produced by Peters after he had made his additions and recodings were more poorly resolved than the trees produced by the original authors.”

That’s another trick. “more poorly resolved’ is not the issue. The issue is: Did pterosaurs align with the four added taxa? Or did pterosaurs align with archosaurs like Scleromochlus? Peters 2000 results indicated pterosaurs nested with the four small tanystropheids when these taxa were added to matrices that originally omitted them.

On that note, the LRT is not fully resolved, but pretty well resolved. Test the LRT yourself at ‘The ReptileEvolution.com Project” at FigShare.com.

“These re-analyses by Bennett (1996), Dilkes (1998) and Peters (2000) have shown that the otherwise emerging consensus on the phylogenetic positions of pterosaurs and prolacertiforms is not uniformly supported.”

Again, there is no reason to imagine that ‘prolacertiformes’ sans Langobardisaurus, Cosesaurus, Sharovipteryx and Longisquama would be ‘uniformly supported’ vs the ‘heterodox’ Peters 2000 in which these four taxa were included.

Taxon exclusion is and always was the number one problem in paleontology, and now, dear readers, I hope you are starting to understand the brains behind this continuing problem.

“The aim of this study is to evaluate the evidence of Bennett (1996) and Peters (2000) critically and to consider the likelihood or not of a close relationship between the pterosaurs and basal archosauromorph reptiles.”

So, HB 007 set up one study that included four tiny tanystropheids vs one that did not.

Part 2 tomorrow.

References
Benton MJ 1999. Scleromochlus taylori and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society London, Series B 354 1423-1446. pdf
Hone DWE and Benton MJ 2007. An evaluation of the phylogenetic relationships of the pterosaurs among the archosauromorph reptiles. Journal of Systematic Palaeontology 5(4):465–469.
Peters 1997. A new phylogeny for the Pterosauria. Journal of Vertebrate Paleontology Abstracts of Papers. Fifty-seventh annual meeting. Chicago, Illinois.
Peters D 2000a. A redescription of four prolacertiform genera and implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293-336.
Peters D 2000b Description and Interpretation of Interphalangeal Lines in Tetrapods 
Ichnos, 7: 11-41
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist 
Historical Biology 15: 277-301.
Peters D 2007. The origin and radiation of the Pterosauria. Flugsaurier. The Wellnhofer Pterosaur Meeting, Munich 27

Source: https://pterosaurheresies.wordpress.com/2026/05/21/hone-and-benton-2007-revisited/