Sangster redescribes Dimorphodon macronyx

Dr Sangster 2022 redescribes
one of the first known pterosaurs, Dimorphodon macronyx (Buckland 1829, Figs 1, 2) in a new monograph. I’m guessing this follows and is largely based on Sangster 2001, her ?unpublished dissertation titled “Anatomy, functional morphology and systematics of Dimorphodon”.

Wonder what’s new in the last 21 years? Or the last 93?

The skull of Dimorphodon macronyx.
Figure 1. The skull of Dimorphodon macronyx. Above: in situ. Middle: Restored. Below: Palatal view. Colors reflect the times (2011) when bones lacked standardized colors. Colors have not been updated here.

The abstract:
“Dimorphodon macronyx (Buckland, 1829) is one of the earliest known Jurassic pterosaurs and the first English pterosaur to have been formally described; the original specimens from the London Natural History Museum have not been fully reviewed since Richard Owen’s monograph of 1870. This monograph provides a detailed, comparative osteological description and emended diagnosis of D. macronyx, based primarily on the four key London and British Geological Survey specimens, and supplemented by the other known skeletal material. Previously undescribed palatal bones uncovered in one of the Natural History Museum specimens (NHMUK 41212) in 2002 are also described and a tentative reconstruction of the palate presented. A short review of previous phylogenetic studies on the taxonomic status of D. macronyx is provided.”

The palate of NHMUK 41212 (Figs, 1, 3) was presented here in 2011.

Figure 2. Images of Dimorphodon from The tail attributed to Dimorphodon is shown in figure 3.
Figure 2. Images of Dimorphodon from The tail attributed to Dimorphodon is shown in figure 3.

From the University of Birmingham website:
“Sarah completed her PhD on the Jurassic pterosaur Dimorphodon at the University of Cambridge in 2003.”

As you can see, sometimes publications can be painfully slow in academia, with her PhD awarded two years after the date of her dissertation and the monograph 21 years later.

Anurognathid palate
Figure 3. Anurognathid palates in phylogenetic order. All of these specimens are badly crushed so picking out details is difficult at best. In 2011 this was the first publication of several anurognathid palates all in one place.

Buckland W 1829. Proceedings of the Geological Society London, 1: 127.
Owen R 1859. On a new genus (Dimorphodon) of pterodactyle, with remarks on the geological distribution of flying reptiles.” Rep. Br. Ass. Advmnt Sci., 28 (1858): 97–103.
Nesbitt SJ and Hone DWE 2010. An external mandibular fenestra and other archosauriform character states in basal pterosaurs. Palaeodiversity 3: 225–233.
Padian K 1983. Osteology and functional morphology of Dimorphodon macronyx (Buckland) (Pterosauria: Rhamphorhynchoidea) based on new material in the Yale Peabody Museum, Postilla, 189: 1-44.
Sangster S 2001. Anatomy, functional morphology and systematics of Dimorphodon. Strata 11: 87-88.
Sangster S 2022. The osteology of Dimorphodon macronyx, a non-pterodactyloid pterosaur from the Lower Jurassic of Dorest, England. Monographs of the Palaeontological Society 175, 221 (661):1–48.


Averianov et al 2020: origin of Multituberculata

A few Middle Jurassic mammal teeth
led Averianov et al. 2022 to describe those as early multituberculates. Averinov’s team, by default, relied on dental traits, which can become convergent in unrelated taxa. They also excluded pertinent taxa.

First, build your own wide gamut cladogram from more or less complete fossils, then insert dental taxa only where it is safe to do so.

Figure 2. Multituberculates to scale. Carpolestes is the proximal outgroup taxon.
Figure 1. Multituberculates to scale. Carpolestes is the proximal outgroup taxon.

Multituberculates are close to carpolestids, plesiadapiforms and rodents
in the large reptile tree (LRT, 2119 taxa). The LRT tests taxa that other authors omit. The Averianov et al. abstract says nothing about the origin of this clade, described as, “the most diverse and abundant group of Mesozoic mammals.” Averinov saves his hypotheses on interrelationships for the text.

Figure 1. Haramiyavia reconstructed and restored. Missing parts are ghosted. Three slightly different originals are used for the base here. The last appears to be the least manipulated and it appears to fit the premaxilla better.  The fourth maxillary tooth appears to be a small canine. The groove on the dorsal premaxillary appears to be for the nasal, not the septomaxilla. Parts are taken from both mandibles
Figure 2. Haramiyavia reconstructed and restored. Missing parts are ghosted. This is the sort of pre-mammal Averinov thought had teeth similar to multituberculates. Other taxa (see figure 1) have tooth shapes closer to multis. Note the medial premaxillary incisors are not chisel-shaped. Note the last premolar is not larger than the first molar.

Averinov et al report,
“In spite of the large fossil record for Multituberculata, their origin and phylogenetic relationships are still unclear, because of their distinctive morphology that is different from that of all other mammals.”

Different from all other mammals? No. That ‘distinctive morphology’ is focused only on the posterior jaw bones, which do not shrink during ontogeny as an embryo to become ear bones. This was a reversal due to neotony, enhancing the ability of the jaws to slide posteriorly, as we learned earlier here in 2019 and here in 2016. Multis are rodents. So are aye-ayes. So are plesiadapiformes.

Figure 3. Images from Han et al. Color and white labels added. Here the malleus, incus and stapes have reverted to their pre-mammal states and configurations. Note the quadrate is in contact with the articular, as in pre-mammals as the dentary and squamosal become a sliding joint, carried by larger jaw muscles. Also note the various ectotympanic bones (yellow) also present, typical of Theria.
Figure 3. Arboroharamiya from Han et al. Color and white labels added. Here the malleus, incus and stapes have reverted to their pre-mammal states and configurations. Note the quadrate is in contact with the articular, as in pre-mammals as the dentary and squamosal become a sliding joint, carried by larger jaw muscles. Also note the various ectotympanic bones (yellow) also present, typical of Theria.

Averianov et al report,
“The most intriguing finding from the discovery of the new Middle Jurassic multituberculate material is the fact that multituberculates show affinities not with Haramiyida as a whole, but specifically with Euharamiyida.” The LRT tests two purported haramiyids. After testing Arboroharamiya (Fig 3) nests with Carpolestes, between rodents and multituberculates in the LRT. Haramiyavia (Fig 2) is a pre-mammal that nests with Brasilodon and Sinoconodon.

Averianov is otherwise notable
for publishing on Azhdarcho and other eastern Europe pterosaurs. Click here for prior reports.

Averianov A et al (8 co-authors) 2020. Multituberculate mammals from the Middle Jurassic of Western Siberia, Russia, and the origin of Multituberculata. Papers in Palaentology 7(2): DOI:10.1002/spp2.1317


SVP 2021 abstracts – 06: Multituberculate bone histology more similar to placentals
A post-dentary reversal between rodents and multituberculates

The origin of archosaurs, crocodylomorphs and dinosaurs revisited

The addition of new data presented yesterday
for the basal crocodylomorph, Junggarsuchus, brought new understanding to the origin of the Archosauria and Dinosauria in the large reptile tree (LRT, 2119 taxa).

The basal poposaur,
Turfanosuchus (Fig 1) remains the proximal ancestor to the Archosauria.

The former basalmost archosaur in the LRT,
the PVL 4597 specimen, moved to the base of the small bipedal crocodylomorph clade that includes Dibothrosuchus, Gracilisuchus, Scleromochlus and others.

The new basalmost member of the Crocodylomorpha,
Lewisuchus (Fig 1), is also the proximal sister to the same basalmost member of the Dinosauria, Herrerasaurus. Phylogenetic miniaturization is still present at and near this node where taxa are small, fast, bipedal and likely transitioned to a warm-blooded metabolism.

Figure 1. Origin of dinosaurs with Turfanosuchus, Lewiisuchus and Herrerasaurus. Note the phylogenetic miniaturization preceding the origin of the Archosauria in Turfanosuchus. Somehow these taxa rarely make it into origin of dinosaurs studies due to cherry-picking taxa. Let your software tell you which taxa.

There are no pterosaurs in this transition.
Pterosaurs nest elsewhere, with small, bipedal, flapping, lepidoaur, tanytropheid fenestrasaurs.

There are no lagerpetids in this transition.
Lagerpeton nests elsewhere, with mid-sized, bipedal, running on two toes proterochampsids close to Tropidosuchus.

There are no silesaurids in this transition.
Silesaurus nests within Poposauria, derived from Turfanosuchus in a differernt direction.

Archosauria is comprised
of just Dinosauria and Crocodylomorpha. That’s all.

Archosauriformes extend back to the late Permian
where they split into Euarchosauriformes (= Youngoides UC1528, Euparkeria and descendants) and Pararchosauriformes (= Youngina AMNH 5561, Proterosuchus and descendants).

Ornithodira and Avemetatarsalia
are junior synonym for Reptilia in the LRT because they each include dinos and pteros. Thesee appear on opposite sides of the first reptilian dichotomy in the Viséan (Early Carboniferous) documented here a decade ago and called ‘The Big Kahuna.‘ To replicate this tree topology workers are going to have to add taxa back to the late Devonian taxon, Tulerpeton.

Why did ‘Avemetatarsalia’ subsume the earlier ‘Ornithodira’?
Because professor MJ Benton 1999 came up with ‘Avemetatarsalia’ in his Scleromochlus paper AND he writes university level vertebrate paleontology textbooks. Professors have to teach what’s in the textbook, even if it’s been invalid since Peters 2000. (That’s how paleontology works.)

The definition of Avemetatarsalia
is “all archosaurs more closely related to birds than to crocodilians.” Benton thought tiny Scleromochlus was close to dinosaurs and pterosaurs. Ironically Scleromochlus is a basal bipedal crocodylomorph in the LRT (subset Fig 2). Taxon exclusion is the problem here. Workers have been unwilling to test taxa that have been in the literature for pterosaur origins for the last 22 years (e.g. Peters 2000). That’s why the LRT is here: to test taxa that have never been tested together and to present results without academic filters and interference.

Still waiting
for one more PhD to run an analysis with a similar taxon list and their own character list to confirm, refute or correct the LRT or a subset of it. Other workers did this a few years ago with Chilesaurus and Diandongosuchus by ?convergence. The lack of curiosity and the unwillingness to test more pertinent taxa in the academic community is still puzzling. As this blogpost documents, cladograms are not set in stone, even if printed in expensive and venerated textbooks. New data from old taxa and new taxa can modify tree topologies. That’s a good thing, always.

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
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.


New data on the basal crocodylomorph Junggarsuchus

Junggarsuchus sloani
(Clark et al. 2004; Ruebenstahl et al. 2022; IVPP V14010; Fig 1) nests among the basalmost crocodilomorphs in the large reptile tree (LRT, 2119 taxa, subset Fig 2), close to Pseudhesperosuchus and likely just as bipedal. Junggarsuchus was originally considered a sphenosuchian nesting between Dibothrosuchus and Protosuchus and basal to extant crocodilians. The LRT includes more basal taxa.

Figure 1. Junggarsuchus recolored from Ruebenstahl et al. 2022.

When new data comes in,
however it comes in, embrace it. Correct mistakes. Make whatever you have better. Help others correct their mistakes.

Figure 2. With the new data the nesting of Junggarsuchus did not change, but the new data and a review of prior skulls moved a few taxa to new nesting sites in this subset of the LRT. By coloring relevant nodes here it is easy to see which taxa were bipedal, which were aquatic and which were marine. Only aquatic taxa survive today.

In this case,
corrections were made and Lewisuchus + Saltoposuchus moved to the basal node within the Crocodylomorpha. The PVL 4597 specimen moved away from the basal Archosauria node to a more derived node basal to the Gracilisuchus clade of bipedal crocs.

Figure 3. Carnufex is basically a giant Pseudhesperosuchus. Here they are compared to one another to scale and with skulls side by side. Dark gray areas are imagined on the original at bottom by Zanno et al. Click to enlarge. With a skull 4x larger than that of Pseudhesperosuchus, Carnufex was a likely 4.4 meter long bipedal killer. Note the smaller orbit and deeper jugal. Both neural arches are missing a centrum.
Figure 3. Carnufex andPseudhesperosuchus are compared to one another to scale and with skulls side by side. Dark gray areas are imagined on the original at bottom by Zanno et al. Click to enlarge. With a skull 4x larger than that of Pseudhesperosuchus, Carnufex was a likely 4.4 meter long bipedal killer. Note the smaller orbit and deeper jugal. Both neural arches are missing a centrum.

At its best, science advances step by step.
Today’s post is just one of those steps.

Clark JM, Xu X, Forster CA and Wang Y 2004. A Middle Jurassic ‘sphenosuchian’ from China and the origin of the crocodilian skull. Nature 430:1021-1024.
Ruebenstahl AA et al. 2022. Anatomy and relationships of the early diverging Crocodylomorphs Junggarsuchus sloani and Dibothrosuchus elaphros. The Anatomical Record DOI: 10.1002/ar.24949


Tethydraco: a latest Cretaceous Pteranodon

Longrich, Martill and Andres 2018 described
several North African pteranodontid pterosaurs, some of which we looked at earlier here.

Figure 1. Tethydraco compared to the UALVP 24238 specimen of Pteranodon (Dawndraco) drawn to two size scales. Pretty good match.

Tethydraco regalis is the last of these North African pterosaurs,
known from a few long bones in at least two sizes (Fig 1). The proportions indicate a volant pterosaur, with a relatively shorter tibia than the UALVP specimen. There’s not a lot more that can be said given the few bones described.

Longrich NR, Martil DM and Andres B 2018. Late Maastrichtian pterosaurs from North Africa and mass extinction of Pterosauria at the Cretaceous-Paleogene boundary. PLoS Biol 16(3): e2001663.


Small Korean pterosaur tracks from the Late Cretaceous

Jung et al 2022 describe
“a new pterosaur footprint assemblage from the Hwasun Seoyuri tracksite in the Upper Cretaceous Jangdong Formation of the Neungju Basin in Korea. The assemblage consists of many randomly oriented prints in remarkably high densities but represents a single ichnotaxon, Pteraichnus.” See figure 1.

Figure 1. Late Cretaceous pterosaur tracks D-F from Jung et al 2022. Actual size @72dpi. Colors and PILs added here. Pterodaustro embryo to scale. Embryo pes to match size of Korea tracks. The Korean tracks are 3x the size of the embryo pes, so they could be juvenile tracks. Since pterosaurs grow isometrically, and go through phylogenetic miniaturization, age cannot easily be determined from tracks.

Jung et al 2022 continue:
“Individuals exhibit a large but continuous size range, some of which, with a wingspan estimated at 0.5 m, are among the smallest pterosaurs yet reported from the Upper Cretaceous, adding to other recent finds which contradict the idea that large and giant forms entirely dominated this interval.”

Those ‘large and giant dominating forms include pteranodontids and azhdarchids.

Unusual features of the tracks, including relatively long, slender pedal digit impressions, do not match the pes of any known Cretaceous pterosaur, suggesting that the trackmakers are as yet unknown from the body fossil record.

This is incorrect. When the tracks were traced, PILs (Peters 2000a) added and traits run through analysis in the large pterosaur tree (LPT, 262 taxa) ichnite ‘e‘ (Fig 1) matched the pes of the embryo Pterodaustro (Fig 1) without decreasing resolution (= increasing MPTs).

Jung et al did not correctly trace the specimens. They did not add PILs. They did not run pedal scores through a phylogenetic analysis that included the embryo Pterodaustro. They did not include ‘The Catalog of Pterosaur Pedes for Trackmaker Identification‘ (Peters 2011) in their citation list. Rather they (and co-author David Unwin) followed Chris Bennett’s curse, “You will not be published and and if you are published you will not be cited.”

The Hwasun pterosaur footprints appear to record gregarious behavior at the exact location by individuals of different ages, hinting at the possibility that pterosaurs gathered in mixed-age groups.”

Ichnos published ‘The Catalog of Pterosaur Pedes’ and the ‘Description of Interphalangeal Lines’ (= PILs) for a reason: to help workers identify pterosaur tracks. If you’re wondering why academic workers don’t use and cite these and other pertinent pterosaur papers click this link to the Timeline of Pterosaur Origin Studies.

Jung J et al 2022. Evidence for a mixed‑age group in a pterosaur footprint assemblage
from the early Upper Cretaceous of Korea. ScientificReports 12:10707
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods.  Ichnos 7:11-41.
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification. Ichnos 18(2):114-141.

Triassic revolution?

Benton and Wu 2022 considered several factors
affecting the return of life following the Permian – Triassic extinction event, (Fig 2) universally considered the worst extinction event of all time. Pertinent abbreviations include:

PTMEPermian-Triassic Mass Extinction – 251 mya (= Siberian traps)
CPECarnian Pluvial Episode – 231 mya (= widespread arid to rainfall)
ETMEEnd Triassic Mass Extinction – 200 mya (= Mid-Atlantic volcanoes)

MMR – Mesozoic Marine Revolution, (= the increase in shell-crushing and boring predation starting in the Early Triassic.

One factor Benton and Wu omitted
was the near complete absence of Permian marine vertebrates in the fossil record (Figs 1, 2). That’s 47 million years of virtually absent taxa.

According to Benton and Wu 2022,
“Marine predatory vertebrates show spectacular and rapid diversifications in the Early and Middle Triassic, and new discoveries from China have confirmed their early start in the Triassic, but not in the Late Permian.”

Or was this a geological illusion? There was a new dawn for animals following the PTME. We know which clades survived. Where did the survivors find refuge? Unfortunately, this topic is rarely to never covered, perhaps because those rare refugia have never been discovered or described. Extinction events garner headlines and take the spotlight.

Figure 2. Chart from Benton and Wu 2022. Overlay notes the absence of Permian marine vertebrates. They had to have been present Their absence is conspicuous here, but overlooked by the authors.

From another point of view
that ‘spectacular and rapid diversification‘ in the Triassic was at least partly due to the current paucity of marine fossils in the Permian (Figs 1, 2) and a relative trove of marine fossils in the Early Triassic. Blank spaces (Fig 2) can still provide data based on phylogenetic bracketing with a large gamut cladogram, like the large reptile tree (LRT, 2119 taxa, subset Fig 3).

Figure 3. Subset of the LRT focusing on ichthyosaurs and kin color coded according to chronology from the early Permian to the Cretaceous.

As an example:
here (Fig 3) the currently unknown pre-Triassic ancestors of ichthyosaurs and thalattosaurs would have to have been coeval with known and related Early Permian mesosaurs. Unfortunately all we have are two Middle Triassic late-surviving basal pachypleurosaurs and pre-pachpleurosaurs, Honghesaurus and Anarosaurus. That’s too few. Way too few for 47 million years. Where are all the Early to Late Permian marine fossils?

One answer: The late-surviving representatives of missing Permian taxa are present in the Early and Middle Triassic according to the LRT. That’s a clue that primitive relatives of Early Triassic taxa were present in the Permian.

There are no large phylogenetic gaps in the LRT.
It documents a continuous gradation phyogenetically, but not chronologically due to the patchy fossil record in general. That also means Benton’s and Wu’s “spectacular and rapid diversificaiton” might actually represent a chronological illusion. In other words, with phylogenetic bracketing the wealth of Early Triassic ichthyosaurs and pre-ichthyosaurs likely arose sometime during the Early Permian. So far we’ve only found late survivors in the geologically richer Early and Middle Triassic strata. If so marine predators, including ichthyosaurs, had a slower, steadier, more typical radiation and evolution.

We haven’t found many Permian marine vertebrate fossils yet.
Perhaps they weren’t fossilized. Perhaps those strata are not present. Perhaps those strata await discovery buried under other sediments. At times like this it’s smart to remember, in general fossils from all eras are extremely rare and often restricted to tiny pin points on the Earth’s surface skipping large swathes of geological time. That’s just the way it is.

Figure 5. Atopodentatus compared to more primitive sister taxa, Adelosaurus and Claudiosaurus.
Figure 4. Middle Triassic Atopodentatus compared to more primitive sister taxa, Permian Adelosaurus and Claudiosaurus to scale.

Overlooked by Wikipedia, Benton and Wu and saved ’til now,
there are several Permian marine and semi-marine younginiforms known from the late Permian. These include Claudiosaurus. Adelosaurus (Fig 4) and related late survivors of a Carboniferous radiation. They likewise had larger, more derived Triassic ancestors, like Atopodentatus (Fig 4).

Figure 5. Giant Cymbospondylus youngorum (Sanders et al. 2021) does not appear until the Middle Triassic. Wumengosaurus is a late survivor known from Middle Triassic strata, but phylogenetically must have had its genesis in the Permian.

Benton and Wu 2022 cited
Sanders et al. 2021 who reported on a giant ichthyosaur, Cymbospondylus youngorum (Fig 5). The authors reported, “The animal existed at most 8 million years after the emergence of the first ichthyosaurs, suggesting a much more rapid size expansion that may have been fueled by processes after the Permian mass extinction.” Benton and Wu considered this “a prime example of an ‘early burst’ radiation.

That’s only 3 million years past the Early Triassic or 35 milllion years past the Early Permian, when pre-ichthyosaurs had their roots. How fast ichthyosaurs evolved is left up to the vagaries of the poor Permian marine fossil record vs the rich Triassic marine fossil record known to geology at present.

Figure 1. The naked mole rat, Heterocephalus is closer to hedgehogs than to rats.
Figure 6. The naked mole rat, Heterocephalus, has has only a few long sensory hairs that likely help it sense the tunnel walls. These provide no insulation, as in burrowing basal cynodonts first sprouting similar hair.

On land, Benton and Wu remind us,
“the Triassic was marked by a posture shift from sprawling to erect, and a shift in physiology to warm-bloodedness, with insulating skin coverings of hair and feathers.”

The authors consider the origin of hair in Permian synapsids as “an insulating pelage”. Before hair can act as insulation it must be thick enough covering a broad enough area. So the genesis of sparse and individual hairs (Fig 6) must have had some other use that enhanced survival. Perhaps the burrowing naked mole rat, Heterocephalus, can give us some insight into the origin of individual hairs first appearing in burrowing cynodonts, retained in mole rats by either a deep time reversal or neotony from ‘hairless’ newborns, or both as ontogeny recapitulates phylogeny. Yes, this is what some cynodont skin looked like.

Benton and Wu report,
“Pterosaurs, the sister group of dinosaurs, also have dermal insulating structures commonly
called pycnofibres.”

Pterosaurs are not the sister group of dinosaurs. The omitted ancestors and non-volant cousins of pterosaurs, like Sharovipteryx (Fig 7), also have dermal insulating, heat-radiating and decorative structures.

Figure 1. Note the neck skin (integument) of Sharovipteryx, a pterosaur sister.
Figure 7. Note the neck skin (integument) of Sharovipteryx, a pterosaur sister.

Benton and Wu continue,
“even if researchers balk at calling pycnofibres feathers, it does not change the fact that insulating dermal structures appeared in the first dinosaurs and the first pterosaurs, and the shared ancestry of these two clades is dated to the Early or early Middle Triassic.”

Fact? Insulating dermal structures appeared in the first dinos and pteros? Yes, for pterosaur ancestors like Cosesaurus in the Middle Triassic. No, for dinosaurs like Herrerasaurus, also in the Middle Triassic. Benton has a long history of omitting taxa related to pterosaurs recovered by Peters 2000a, b. Paleo academics at large (any exceptions?) continue to do the same.

Benton MJ and Wu F 2022. Triassic Revolution, Frontiers in Earth Science (2022). DOI: 10.3389/feart.2022.899541
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.
Sanders et al. (6 co-authors) 2021. Early giant reveals faster evolution of large body size in ichthyosaurs than in cetaceans. Science374 (6575): DOI: 10.1126/science.abf5787


The Princeton Field Guide to Pterosaurs by Gregory Paul 2022

A while back,
paleontologist Greg Paul let me know about his new Princeton Field Guide in prep, this one about pterosaurs (Fig 1). Paul is widely acknowledged as the early leader in creating reconstructions of dinosaurs and other animals by strictly adhering to their skeletal traits and proportions.

Paul’s newst Field Guide was published June 7, 2022 with little fanfare.
I only heard about it yesterday via an off-hand blogpost comment that did not mention the book or author by name. Knowing what was in the pipeline, a keyword search was enough to find the book online. Barnes and Noble, Amazon, even Target all advertise it.

Figure 1. Cover illustration from Paul 2022. The shallow wing chord is correct. So are the uropatagia. I see little to no propatagia here as the humeri appear to extend laterally.

From the publisher’s description:
“Once seen by some as evolutionary dead-enders, pterosaurs were vigorous winged reptiles capable of thriving in an array of habitats and climates, including polar winters. The Princeton Field Guide to Pterosaurs transforms our understanding of these great Mesozoic archosaurs of the air. This incredible guide covers 115 pterosaur species and features stunning illustrations of pterosaurs ranging in size from swallows to small sailplanes, some with enormous, bizarre head crests and elongated beaks. It discusses the history of pterosaurs through 160 million years of the Mesozoic—including their anatomy, physiology, locomotion, reproduction, growth, and extinction—and even gives a taste of what it might be like to travel back to the Mesozoic. This one-of-a-kind guide also challenges the common image of big pterosaurs as ultralights that only soared, showing how these spectacular creatures could be powerful flappers as heavy as bears.”

This field guide features
115 different kinds of pterosaurs as skeletal drawings and full-color life studies. It covers pterosaur biology and the colorful history of pterosaur paleontology. Surprisingly I am listed in the ‘Acknowledgments’ section.

Let’s look inside the first few pages… The Preface
Paul is an engaging writer who draws the reader in with his literary conceit as a ‘mysterious time traveler’ both ‘startled’ and ‘delighted’. Paul described the ‘surreal’ variety in head crests.

Unfortunately Paul errs and promotes myth
1. when he describes Quetzalcoatlus (Fig 2) as, ‘the largest flying creature possible.’
2. when he calls pterosaurs distant archosaur relatives of birds.
3. when he reports, “Remaining frustratingly unresolved are the origins of pterosaurs.”

History of discovery and research section
Paul’s account is accurate and reminds one of how few and far between pterosaur papers appeared prior to the 1970s. This is when Paul estimated mass for Quetzalcoatlus (Fig 2). He reports, “My calculation that the biggest pterosaurs must have weighed much more than the biggest living ground birds (Paul 1991, 2002) was initially controversial, but it has since become widely accepted.”

If I’m not mistaken, this was Paul’s last contribution to pterosaur research. He has not participated in any phylogenetic studies, nor is he a co-author on any discoveries. Let me know if this is an error and I will correct it.

Figure 1. Click to enlarge. The largest flying and non-flying birds and pterosaurs to scale.
Figure 2. Click to enlarge. The largest flying and non-flying birds and pterosaurs to scale. Compare one leg of Quetzalcoatlus to the 175 pound ex-president to estimate its total weight.

History of discovery and research section continued.
“And the old classic Pteranodon underwent a notable revision with the realization that it had a major overbite, with the lower beak markedly shorter than the upper.”

The is true, but only for some Pteranodon species (Fig 3), not all. Paul should have known this before writing his book.

Figure 2. The Tanking-Davis specimen compared to other forms. Specimen w and specimen z appear to be the closest to the Tanking-David specimen. Specimen 'w' = Pteranodon sternbergi? USNM 12167 (undescribed). Specimen 'z' = Pteranodon longiceps? Dawndraco? UALVP 24238. Click to enlarge.
Figure 3. Click to enlarge. Some Pteranodon have a longer rostrum than mandible. Others don’t. Paul doesn’t know this. Any author writing about pterosaurs should know this.

Paul continues his history of discovery and research.
“What has not yet shown up is fossils of protopterosaurs because suitable fine-grained lake or lagoon bottom deposits… are not known”

This is incorrect. We have pterosaur ancestors back to Cambrian worms in the large reptile tree (LRT, 2119 taxa). Proximal Middle Triassic ancestors include Cosesaurus (Fig 4, Peters 2000). Relatives of the fenestrasaur radiation include Sharovipteryx and Longisquama (Fig 4). Each of these are found in such fine-grained sediments that jellyfish, insects and soft tissue are preserved or molded. Paul should have known this before writing his book.

Unfortunately Paul has never constructed a phylogenetic analysis. Like Paul, I was once a paleo artist, but I learned that a phylogenetic analysis answers so many questions and resolves every enigma with authority. No more wondering.

Click to enlarge. Squamates, tritosaurs and fenestrasaurs in the phylogenetic lineage preceding the origin of the Pterosauria.
Figure 4 from 2011. Lepidosaurs tanystropheids and fenestrasaurs in the phylogenetic lineage preceding the origin of the Pterosauria.

Paul continues his history of discovery and research.
“The evolution of human understanding of pterosaurs has not undergone as dramatic a transformation as has our view of dinosaurs over the last quarter millennium.”

This is incorrect as figure 4 shows. Paul should have known this before writing his book.

Paul 2022 – What is a pterosaur?
Paul relies on ‘the great majority of researchers’ who also throw up their hands in frustration, not willing to examine the taxa in figure 3. This is a surprising action given Paul’s earlier fame at breaking with consensus based on his own research. Instead here Paul relies on the literature. He writes, “There has been considerable speculation as to which archosaurs the pterosaurs are most closely related to.”

This is the problem. Pterosaurs are not and have never been archosaurs. So let’s stop looking there and look elsewhere (Fig 4). Like Witton before him, Paul should have known what pterosaurs were before writing his book. Presently Paul is 22 years behind the times (Peters 2000) and promoting invalid myths.

“Pterosaurs had a simple dinosaur-type ankle.”

Never, ever rely on a single character to define a clade. That’s called ‘Pulling a Larry Martin.” Pterosaurs developed this ankle-type by convergence. The old ankle fetish among paleontologists goes back several decades and is now rarely brought up. Why focus on the ankle when pterosaurs have a giant finger four? Where else do we find finger four larger than three or five? Not in archosaurs. See figure 3 where each taxon in turn share a longer suite of traits with pterosaurs including a longer finger four.

Paul describes
the extreme elongation of ‘an outer finger’, apparently unable to determine if it is the fourth following the three free fingers, all bearing the typical reptile/lepidosaur number of phalanges. Paul reports, “The chest features a large sternal plate…lacking the deep bony keel. There is no wishbone furcular“. (= conjoined clavicles).

The pterosaur sternal complex is actually composed of co-ossified clavicles, a single sternum overlapping an interclavicle (Fig 5). We’ve known that since Wild 1993. Archosaurs don’t have interclavicles. Lepidosaurs do. Paul fails to mention the strap-like scapula and locked-down stem-like coracoid that indicates bipedality and flapping, as in birds. Paul should have known this before writing his book.

Tritosaur pectoral girdles demonstrating the evolution and migration of the sternal elements to produce a sternal complex.
Figure 5. Lepidosaur pectoral girdles demonstrating the evolution and migration of the sternal elements to produce a sternal complex.

Paul mentions the prepubis,
but fails to note its significance or distribution (Fig 4). He also notes the long slender ilium, but fails to note its significance or distribution (Fig 4). Paul reports, “the foot was plantigrade.” Sometimes that was so. Other times, no (Peters 2011). Paul reports, “pterosaur eggs were soft-shelled”, but he fails to note that significance. Lepidosaurs have similar soft shells and the mother keeps them within her body longer, sometimes until just prior to hatching. Paul should have known this before writing his book.

Figure 6. Image from page 18 of Paul 2022. These are generalized taxa, not based on a strict interpretation of skeletons (Fig 7). The lowest taxon is Megalancosaurus, not Longisquama. Greg Paul, uncharacteristically, is using freehand drawings to generalize prehistoric animals rather doing what Greg Paul has taught us all: to stick to the skeleton.

Moving into phylogeny again, Paul writes,
“There does not seem to be a great division within the Pterosauria in which two or more distinct groups split apart soon after the initial appearance of the group.”

This is wrong. According to the large pterosaur tree (LPT, 262 taxa) that phylogenetic split occurred in the Late Triassic when Preondactylus (Fig 7) became the last common ancestor of the fragile-skulled, dimorphodontid-anurognathid clade of insect-eaters. On the other branch, Eudimorphodon (Fig 7) was the last common ancestor of all other pterosaurs, the robust-skulled fish-eaters. A phylogenetic analysis would have been helpful here, rather than relying on a vague notion from the extremely flawed (due to taxon exclusion) literature. That way Paul would have been able to tell his readers that the pterodactyloid grade appeared five times by convergence (4x according to Peters 2007. A 5th was recently added). Here again, Paul should have known this before writing his book.

Figure 7. Skeletal studies of the taxa in Paul’s drawing in figure 6. If you want to know what Longisquama looked like see figure 3.

Paul mistakenly writes,
“No known pterosaur was flightless.”

Actually a variety of pterosaurs were flightless. These include his favorite giant pterosaurs. Paul should have known this before writing his book. Perhaps Paul remembers his time with inventor and engineer Paul Macready who sent a mechanical Quetzalcoatlus flying, but only by cheating its proportions (Fig 8).

Figure 6. Paul MacCready's flying pterosaur model had longer wings than Q. sp., with its vestigial distal wing phalanges. Here the model and its inspiration are shown to the same length.
Figure 8. Paul MacCready’s flying pterosaur model had longer wings than Q. sp., with its vestigial distal wing phalanges. Here the model and its inspiration are shown to the same length.

Gee, I wish Greg had touched base while writing his book
just once so I could have suggested new directions and help him avoid omissions. Then I could have earned that Acknowledgements mention. As is, this book comes as a disappointing rehash of old and discredited pterosaur myths. Missing here is the spirit and drive of the brilliant, young Greg Paul, the one we leaned from so many decades ago, the one who saw things precisely, sometimes differently than “the great majority of researchers” he now leans on.

Paul GS 2022. The Princeton Field Guide to Pterosaurs. Princeton Field Guides. Online.
Peters D 2000a. Description and Interpretation of Interphalangeal Lines in Tetrapods. Ichnos 7:11-41.
Peters D 2000b. A reexamination of four prolacertiforms with implications for pterosaur phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106: 293–336.
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. 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.
Peters D 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification
Ichnos 18(2):114-141.
Wild R 1993. A juvenile specimen of Eudimorphodon ranzii Zambelli (Reptilia, Pterosauria) from the upper Triassic (Norian) of Bergamo. Rivisita Museo Civico di Scienze Naturali “E. Caffi” Bergamo 16: 95-120.

The evolution of the ‘spiracular region’ in Shuyu, a tiny galeaspid without a spiracle

Gai et al. 2022 discuss their views
on the origin of the vertebrate spiracular region in Shuyu (Gai et al 2011, Middle Silurian; Fig 1), a tiny galeaspid lacking a spiracle (Fig 2) in a clade lacking spiracles (Fig 3).

Galeaspids were the only vertebrates
to migrate their primitive immobile oral cavity to the dorsal skull (Figs 1, 3). Ecollogically galeaspids appear to have been flattened, sit-and-wait filter feeders, hiding while buried in sandy sediments with only their dorsal surface and oral cavity exposed.

Note: There is no galeaspid named Galeaspis. That’s a trilobite.

Also note: Paleo fish workers don’t yet understand there were two origins for jaws in vertebrates. Galeaspids were not in the gnathostome lineage. Rather their proximal outgroup taxon, Drepanaspis (Fig 3), was also basal to placoderms. Note the gradual migration of the terminal oral cavity to the dorsal surface. Primitive galeaspids retain a transverse oral cavity and lateral eyes. Derived galeaspids developed skull processes, migrated the eyes dorsally and the oral cavity evolved to become circular, then slit-shaped at right angles to the original orientation. Very strange.

Also note: Despite their bony and immobile head plates, galeapids were not related to osteostracans, like the pre-sturgeon, Hemicyclaspis (Fig 4 frame 4). Lampreys, osteostracans and sturgeons all retained a ventral oral opening.

Figure 1. Modified from Gai et al 2022 showing the tiny galeaspid, Shuyu (Gai et al 2011), in dorsal view with the gill chambers exposed. The first gill chamber evolves to become the spiracle.

Gai et all 2011 dismiss the hypothesis
that osteostracans were the closest jawless relative of the gnathostomes because, “their massive, mineralized braincase has a median nasohypophyseal organ resembling the condition in lampreys.”

By contrast, descendants of lampreys (Pteromyzon) in the large reptile tree (LRT, 2119 taxa, subset Fig 6) split into two clades: 1) hard-headed arandaspids, galeaspids and placoderms; and 2) soft-headed Jamoytius, Birkeria, Haikouichthys, hard-headed Hemicyclaspis, semi-hard-headed Thelodus, hard-headed sturgeons and gnathostomes. Both clades evolved their own jaw. Past efforts at describing or discovering the evolution of jaws have failed due to not recognizing the dual and convergent genesis of jaws that comes from adding taxa.

Figure 2. Cownose ray feeding by dropping their cephalic lobes to increase the suction on their oyster prey.
Figure 2. Cownose ray feeding by dropping their cephalic lobes to increase the suction on their oyster prey.

The spiracle is a small opening in the temple region
of rays (Fig 2), sawfish and other flattened, bottom-dwelling, cartilaginous fish. With the ventral mouth buried in silty sediment the spiracle serves as an alternate water intake for respiration. It is also the ear hole in tetrapods. Traditionally the spiracle is homologous with the first gill opening in lamprey-grade and higher chordates (Fig 1). The spiracle is lost in hammerhead sharks and many bony fish (Fig 4). Galeaspids and kin (Figs 1, 4) do not have a spiracle. Their first gill chamber (Fig 1) is not different from the other gill chambers.

Figure 3. Proposted evolution within Galeaspida, starting with Drepanaspis, a taxon not mentioned in galeaspid studies. Note the lack of a spiracle or jaws in any of these bottom-dwelling, filter-feeding taxa. Blue arrows point to oral opening and away from gill exits. Red dots are eyeballs. Dunyu ventral view filled by expandable gular sac enabling respiration and feeding.

From the Gai et al 2022 abstract
“The spiracular region, comprising the hyomandibular pouch together with the mandibular and hyoid arches, has a complex evolutionary history”.

Perhaps more complex than Gai et al realize due to their not recognizing the dual origin of jaws (placoderms and gnathostomes by convergence) in vertebrates (Fig 1). Adding taxa resolves this issue.

Figure 4. This is figure 9 from Gai et all 2022 split to fit this format, then captions moved to the graphic, then skulls removed from the graphic, then adjusted to match the LRT topology.

From the abstract
“Here we present the first confirmed example of a complete spiracular gill in any vertebrate, in the galeaspid (jawless stem gnathostome) Shuyu. Comparisons with two other groups of jawless stem gnathostomes, osteostracans and heterostracans, indicate that they also probably possessed full-sized spiracular gills and that this condition may thus be primitive for the gnathostome stem group”.

No. Gnathostomes (= Chondrosteus (Fig 4) and descendants) developed jaws apart from placoderms. Shuyu is not a stem gnathostome. Sturgeons are pre-gnathostomes.

This contrasts with the living jawless cyclostomes, in which the mandibular and hyoid arches are strongly modified and the hyomandibular pouch is lost in the adult”.

Living lampreys are 500 million years away from that last common ancestor that ultimately split into the placoderm and gnathostome clades. So living lampreys can be highly modified from their Cambrian ancestors.

“On the basis of these findings we present an overview of spiracular evolution among vertebrates.” [See figure 4 frame 1.]

Gai et al. almost have a valid phylogenetic topology (Fig 4). It is missing just a few elements. Do not proceed on your own studies without a valid cladogram. Otherwise all your subsequent hypotheses are prone to crack and possibly crumble.

Gai et al. 2022 report,
“Our new evidence indicates that the so-called interbranchial ridges of galeaspids are actually the dorsal portion of branchial arches. They are incorporated into the neurocranium to form a massive skull as assumed in osteostracans by Stensiö.

By contrast, in the LRT (subset Fig 6) these two bone-headed fish clades (galeaspids and osteostracans) are not related. This is yet another example of convergence not recognized by the authors. Don’t make the same mistake. Create your own LRT so you, too, will have a panoramic view of all vertebrates. Don’t focus too much on one clade or another. You’ll miss something by convergence on the outskirts and the LRT will catch it.

Compared to osteostracans, the entire branchial apparatus in Shuyu retains a general vertebrate condition, thus, it is easy to identify the mandibular and hyoid arches according to their topological position and nerve innervation.”

Shuyu nests close to the base of the Vertebrata in the LRT. So it should retain a general condition.

Figure 5. Cheiracanthus was examined, scored and added to the LRT during the course of this study.

Gai et al. 2022 introduced
a well-known acanthodian in their cladogram (Fig 4), Cheiracanthus (Egerton 1861, Middle Devonian, Fig 5), not previously tested in the LRT. Here Cheiracanthus is traced, scored and nested with Homalacnathus at the base of the acanthodians, which is at the base of the clade the ultimately produced lobe-fin fish and tetrapods. So it’s a human ancestor, but not a tuna ancestor.

Figure 6. Subset of the LRT focusing on basal vertebrates. Note the dual origin of jaws in placoderms and gnathostomes.

This appears to be a novel hypothesis of interrelationships.
If not, please provide a citation so I can promote it here.

What drives the stream of water carrying food and oxygen in a galeaspid? The mouth doesn’t move and it is not pointed anteriorly. The gills don’t move within the solid skull. The soft tissue atrium must be able to shrink and expand like a bellows within the solid skull. The galeaspid skull is open ventrally to permit the expansion of this gular sac (Fig 7, 8). It does not include an open mouth and gill slits, as in unrelated, but often convergent osteostracans.

Figure 7. The streamlined galeaspid, Rhegmaspis xiphoidea IVPP V 19354.3

Added the morning after publication:
Rhegmaspis (Figs. 7, 8) is a streamlined galeaspid. Nochelaspis and Platylomaspis (Fig 8) are large galeaspids often reconstructed with ventral oral and gill openings, like osteostracans. I don’t see those traits here (Fig 8). Rather the ventral portion appears to have been soft tissue, a gular sac that expanded and contracted to drive inhalation and exhalation along with water borne tiny food particles. In other words these are the only fish with a mouth on top of the skull. Nares are not present, but a pineal opening often is present. All this points to a lifestyle not of predation, but of sit-and-wait filtration.

Figure 8. Regmaspis (see figure 7) compared to Nochelaspis and Platylomaspis, two large ‘galeaspids’ with a ventral mouth and gill openings, like those in osteostracans, different from those in other galeaspids. The dorsal median naris is much smaller than in galeaspids, similar to the naris in osteostracans.

Egerton P de MG 1861. British fossils, pp 51-75 in Huxley TH (ed), Preliminary Essay Upon the Systematic Arrangement of the Fishes of the Devonian Epoch, Figures and Descriptions Illustrative of British Organic Remains. Memoirs of the Geological Survey, U.K, (Decade 10).
Gai Z et al 2011. Fossil jawless fish from China foreshadows early jawed vertebrate anatomy. Nature 476:324–327.
Gai Z. Zi M. Ahlberg PE and Donoghue PCJ 2022. The Evolution of the Spiracular Region From Jawless Fishes to Tetrapods. Font. Ecol. Evol., 19 May 2022 |


Tiny Phyllodontosuchus lufengensis enters the LRT alongside tiny Coloradisuchus

Harris et al 2000 described and named
a “leaf-toothed crocodile,” Phyllodontosuchus lufengensis (Sinermurian, Early Jurassic China, 200mya, BVP568-L12, Fig 1), based on the middle and posterior teeth. The anterior teeth remained recurved and sharp. The tiny skull is 7cm (less than 3 inches) long. The authors considered this taxon a sphenosuchian crocodylomorph without conducting an analysis.

After analysis in the LRT (subset Fig 3), Phyllodontosuchus nests elsewhere, apart from Sphenosuchus and kin.

Figure 1. Phyllodontosuchus skull from Harris et al 2000. Colors added here. Shown twice life size @72dpi.

Harris et al reported,
“Unfortunately, much of the detail in the only known skull of Phyllodontosuchus is not discernable due o nuances of preservation, including similarity of the bone to the surrounding matrix. Nevertheless, enough detail is preserved to permit diagnosis.”

Using colors (Fig 1) enough detail can be gleaned to score Phyllodontosuchus.

Harris et al thought
the premaxilla was not preserved.

Colors helped to identify the premaxilla here (yellow Fig 1). The naris is terminal and breaks the jawline, as in equally tiny Coloradisuchus (Fig 2). That can be confusing.

Harris et al thought
“no sutures are discernable in any fragment”.

Maybe they should have colored the bones between the sutures using DGS (Fig 1).

Harris et al thought
“Crushing has apparently filled in the supratemporal fenestrae, and their outlines are likewise indeterminate.”

Digital graphic segregation (Fig 1) was able to discern these structures.

The authors focused their attention on the teeth,
comparing them to herbivorous dinosaurs and other archosauriforms. Harris et al more or less correctly concluded the specimen “shows more similarities with the basal Crocodylomorpha (Sphenosuchia) than any other.”

Figure 1. Coloradisuchus skull from Martinez, Alcover and Pol 2017. Colors added.
Figure 2. Coloradisuchus half-skull from Martinez, Alcover and Pol 2017. Colors added. Shown life size @72dpi. Restoration based on Dibrothosuchus, which is off a node or two.

After reconstruction and analysis
in the large reptile tree (LRT, 2118 taxa) Early Jurassic Phyllodontosuchus nested with Late Triassic Coloradisuchus (Fig 2), a taxon not described until Martinez, Alcober and Pol 2017. These taxa nest between Gracilisuchus and Scleromochlus + Saltopus + Lagosuchus as basal bipedal members of the Crocodylomorpha. The latter two taxa are known from post-crania only, which don’t resolve with skull-only taxa Phyllodontosuchus and Coloradisuchus. In order for complete resolution in the LRT either the skull-only or the skull-less taxa must be left out.

Figure 3. Subset of the LRT focusing on Crocodylomorpha. Adding taxa and rescoring earlier entries moves a few taxa to new nesting sites here.

the tiny, bipedal crocodylomorphs, Gracilisuchus, Scleromochlus, Saltopus and Lagosuchus are not often, if ever, included in other crocodylomorph studies. The LRT, which tests taxa together that have never been tested together, indicates that traditional omission needs to change. Simply add these taxa (and more) to your croc studies to see for yourself how well they fit in. Then let us know what you get.

Martinez, Alcober and Pol 2017 mistakenly considered Coloradisuchus “a new protosuchid crocodiliform.” Unfortunately, Phyllodontosuchus, Gracilisuchus, Scleromochlus, Saltoposuchus and Lagosuchus were not mentioned in the text. Protosuchus nests elsewhere in the LRT.

Harris JD et al 2000. A new and unusual sphenosuchian (Archosauria: Crocodylomorpha) from the Lower Jurassic Lufeng Formation, People’s Republic of China. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 215(1): 47–68.
Martinez RN, Alcober OA and Pol D 2017. A new protosuchid crocodyliform (Pseudosuchia, Crocodylomorpha) from the Norian Los Colorados Formation, northwestern Argentina. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1491047.