Was the AMNH Tanytrachelos ‘with child’?

Tanytrachelos ahynis (Olsen 1979, holotype AMNH 7496; holotype Fig. 1) Latest Triassic, 200 mya, was derived from Macrocnemus and was a sister to Langobardisaurus and Tanystropheus. All are tritosaur lepidosaurs in the lineage of the terrestrial ancestors of pterosaurs, the Fenestrasauria… all ultimately derived from an earlier sister to late-surviving Huehuecuetzpalli and Tijubina.

Figure 1. AMNH 7496 holotype of Tanytrachelos with original tracing from Olsen 1979. DGS colors added.

The AMNH specimen
(Fig. 1) preserved in ventral exposure, appears to have two halves of a leathery eggshell and an ‘exploded’ embryo, best described as several dozen tiny bones that should not be there, unless, perhaps this was a gravid adult… or something else, like gastroliths, undigested prey… hard to tell. In any case, some of the pectoral bones also have new identities here.

Figure 2. Hypothetical Tanystropheus embryo compared to Dinocephalosaurus embryo. These are the sorts and sizes of bones one should look for in any maternal Tanytrachelos.

Figure 3. Tanytrachelos hopping to match Gwyneddichnium tracks (see figure 2).

Distinct from Langobardisaurus,
Tanytrachelos has twelve cervicals, but none were gracile. The posterior cervical ribs had large heads that kept the rods far from each centrum. Heterotopic bones were present. These appear to be elongated chevrons, as in Tanystropheus. Rare hopping prints (Fig. 2) match the size and shape of Tanytrachelos pedes.

Figure 4. The sternal complex of several other tritosaurs. Tanytrachelos is closer to Tanystropheus, not quite like any of these related taxa, but all are informative.

The elliptical sternum
of Tanytrachelos was wide, as in Langobardisaurus (Fig. 3), but the clavicle remained gracile, as in Huehuecuetzpalli (Fig. 3). The humerus was slightly bowed. Metacarpal I aligned with the others. Metatarsal III was the longest. Digit III was the longest as in Langobardisaurus tonelloi.

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References
Olsen PE 1979. A new aquatic eosuchian from the Newark Supergroup Late Triassic-Early Jurassic) of North Carolina and Virginia. Postilla 176: 1-14.
Smith AC 2011. Description of Tanytrachelos ahynis and its implications for the phylogeny of Protorosauria. PhD dissertation. Virginia Polytechnic Institute and State University.

 

One more trait linking hippos to mysticetes (baleen whales)

Vibrassae
are sensory organs found in hippos and baleen whales. Odontocete whales lack such vibrissae.

Figure 1. Mysticete (right whale) vibrissae compared to hippo vibrissae. Odontocete whales lack such structures.

Earlier the large reptile tree (LRT, 1644+ taxa) showed that odontocete members of the traditional clade Cetacea are not related to mysticetes, which arise from a clade of mesonychids, hippos and desmostylians. Odontocetes arise from tenrecs.

A Jurassic squid choking hazard for Rhamphorhynchus

Hoffmann et al. 2020 reported in no uncertain terms,
“Pterosaurs ate soft-bodied cephalopods (Coleoidea).”

Immediately after, Hoffmann et al. dialed it back a bit,
when they wrote, “Here, we report the first evidence of a failed predation attempt
by a pterosaur on a soft-bodied coleoid cephalopod.”

Based on size alone,
the squid (PIMUZ 37358) was more than a mouthful according to this ‘to scale’ diagram (Fig. 1)…at least more than a stomachful.

Ask yourself:
could a Rhamphorhynchus of that size (none were larger) eat a squid of that size? Did the pterosaur fail at predation? Or did it change its mind after biting the squid out of curiosity or boredom and losing a tooth in the process?

Figure 1. Plesioteuthis squid in situ with tooth. Reconstructions of Plesioteuthis (above) and the n81 specimen attributed to the largest known Rhamphorhynhcus, which has a matching tooth. The question is: could that pterosaur eat that squid? Or did it change its mind after biting the squid? At the very top is the hard tissue gladius of the squid to scale. That’s a hard part that would have been especially hard to swallow.

You be the judge.
Hoffmann et al. 2020 have provided the pertinent information. Above are the predator and “prey” to scale. Other Rhamphorhynchus specimens are smaller, and the tooth could have fallen from a different alveolus (a larger tooth) on a smaller specimen. Lots of variables and unknowns here. Also consider the difficulty of swallowing that long gladius, a hard part homologous with the cuttle bone in a cuttlefish.

In any case,
watch what headline you put on your paper. Here the authors went for maximum impact. If, like these authors, you have to dial it back in the second sentence of your abstract,  maybe a more conservative headline should reflect that assessment. After all, a dietary mainstay is indeed different than a curious nibble… and relative size matters.

We looked at other pterosaur choking hazards
earlier here. Pterosaurs likely swallowed their prey whole. There is no indication that they tore squids apart, creating bite-sized pieces. Likewise there is no indication that pterosaurs were able to expand their stomach to accommodate oversize prey (Fig. 1).

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References
Hoffman R, Bestwick J, Berndt G, Berndt R, Fuchs D and Klug C 2020. Pterosaurs ate soft-bodied cephalopods. http://www.nature.com/scientificreports (2020) 10:1230 | https://doi.org/10.1038/s41598-020-57731-2

The king mackerel is not a mackerel

Updated April 24, 2021
with a revision of the ray fin subset of the LRT

The king mackerel
(genus: Scomberomorus; Figs. 1, 2) enters the large reptile tree, LRT, 1643+ taxa) next to the deep sea scabbard fish (Aphanopus; Fig. 3), not the mackerel, Scomber, which entered the LRT a few days ago.

Figure 1. Scomberomorus cavalli is derived from barracuda and gives rise to seahorses and other taxa.

Scomberomorus cavalla (Cuvier 1829; 60cm) is the extant king mackeral or kingfish. King mackerals are derived from Sphyraena + Esox (barracuda and pike) in the LRT. Beside Aphanopus (Fig. 3), the king mackerel is basal to: flying fish + swordfish and  another branch: sticklebacks + sea horses.

Figure 2. Scomberomorus again, this time with the jugal highlighted in cyan.

The addition of this more plesiomorphic taxon
to the LRT makes the nesting of the more derived Aphanopus (Figs, 3, 4), a deep water taxon more understandable. Evidently, the LRT is still working, no matter what it has to work with, whether rare and autapomorphic deep sea taxa or common and plesiomorphic open ocean taxa.

Figure 3. Meter-long Aphanopus, the black scabbard fish, has a long, eel-like torso tipped with a tiny diphycercal tail.

Careful readers will note
the addition of several fish taxa (Fig. 4) shifted several traditional chondrichthyans to the lineage of bony fish, like Doliodus and Iniopteryx, helping to settle that long-standing issue of the genesis of bony fish.

For returning readers, a reminder,
the fish skulls pictured here (e.g. Fig. 2) or in ReptileEvolution.com have been given tetrapod homology colors (pink for nasals, yellow for premaxillae, etc.), which enables the homologies discussed here. Years ago these colors were not standardized, but lately this practice has been maintained.

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References
Cuvier GCLD 1829. Le Règne Animal distribué d’apres son organisation, pour servir de base a l’histoire naturelle des animaux et d’introduction a l’anatomie comparée. Avec figures dessinées d’après nature. Nouvelle édition, revue et augmentée. Tome V. Suite et fin des Insectes. Par M. Latreille. Déterville & Crochard, Paris, i-xxiv + 556pp.

wiki/Black_scabbardfish
wiki/King_mackerel
wiki/Scomber

Doliodus re-enters the LRT

Updated January 29, 2021
with a re-renesting of Doliodus as a basal bony ray fin fish.

Earlier I tried to understand
Doliodus latispinosus (Whiteaves, 1881; Maisey et al. 2018; originally Diplodus, problematicus (“problematic deceiver”) Woodward 1889; Fig. 1) and failed. Traditionally Doliodus has been considered part of the acanthodian-chondrichthyan (spiny shark to shark) transition, but the the large reptile tree (LRT 1643+ taxa) nests acanthodians far from chondrichthyans. Now Doliodus nests with basal ray-fin fish.

Figure 1. Doliodus from Maisey et al. 2018. Colors added. Compared to Malacosteus in figure 2.

Maisey et al. 2018 reported,
“Based on these data, Doliodus and pucapampellids both fall outside the chondrichthyan crown, but their relative phylogenetic positions on the chondrichthyan stem are unclear.

The phylogenetic position of Doliodus seems less elusive; it possessed an ‘acanthodianlike’ complex of dermal spines, including pectoral fin spines, prepectoral, admedian, and prepelvic spines, and possibly dorsal and pelvic fin spines, in conjunction with numerous ‘chondrichthyan-like’ endoskeletal features and a heterodont ‘sharklike’ dentition. Doliodus can be viewed as a quintessential component of the evolutionary transition between ‘acanthodians’ and ‘conventionally defined chondrichthyans’, leaving little doubt that the chondrichthyan total group includes ‘acanthodians’ (now widely perceived to be a paraphyletic group, populating the basal part of the chondrichthyan stem).”

This was probably a nocturnal bottom feeder
with a wide skull, wider than the narrow (as determined by the inter-fin distance) torso. Perhaps this flat taxon was transitional between low pectoral fins of most fish and the high-set pectoral fins that set iniopterygians apart.

Miller, Cloutier and Turner 2003 also reported on Doliodus.
They wrote, “This species has been truly problematic. Previously known only from isolated teeth, it has been identified as an acanthodian and a chondrichthyan [= sharks and rays]. This specimen is the oldest shark showing the tooth families in situ, and preserves one of the oldest chondrichthyan braincases. More notably, it shows the presence of paired pectoral fin-spines, previously unknown in cartilaginous fishes [= sharks and rays].” 

Check the cladogram (Fig. x).
The LRT resolves this issue clearly. This is a novel hypothesis of interrelationships based on taxon inclusion. If anyone published the same interrelationships earlier, let me know so I can promote that citation.

Later, just Turner and Miller 2004 wrote,
“The most important feature of this fossil is its paired pectoral spines. These suggest that many isolated fossil spines might have belonged to sharks rather than acanthodians as previously believed. Features of the fossil blur the distinction between acanthodians and early chondrichthyans.”

“Textbooks still parrot the conventional thinking that no fossil sharks are found before the late Devonian, but this dogma ignores work from the last three decades. The oldest microfossils definitely attributable to sharks are scales in Silurian strata (440 mya) of Siberian and Arctic Russia. 

“Early Silurian deposits in the Tarim Basin of western China have also yielded fin spines associated with sharklike scales. Are these fossils true sharks? If so, the lineage was apparently toothless for millions of years. The first indisputable shark teeth do not turn up until about 50 million years after the appearance of these first putative shark scales in the late Ordovician.”

Figure x. Current subset of the LRT focusing on ray-fin fish.

In the LRT,
big-eyed, flat-skulled Doliodus (Fig. 4) nests with other big-eyed, spiny-finned fish in the clade that leads to bony fish, not sharks. Like the unrelated Xenacanthus, Doliodus has double-tipped teeth.

Figure 3. Malacosteus niger in lateral view.

Sharks are also known from a tooth battery,
a conveyer belt lineup of teeth waiting to rotate into place, as in Dolidus. The ‘millimeter-size teeth’ of Doliodus all point toward the tongue (Fig. 4), so the next teeth in the battery rotate to this position, rather than simply ascend or descend into position, as in tetrapods.

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References
Maisey JG et al. (6 co-authors) 2018. Doliodus and Pucapampellids: Contrasting perspectives on stem chondrichthyan morphology. Chapter 5 in Evolution and Development of Fishes.
Miller RF, Cloutier R and Turner S 2003. The oldest articulated chondrichthyan from the Early Devonian period. Nature 435:501–504.
Turner S and Miller RF 2004. New ideas about old sharks. American Scientist 93:244–252.
Whiteaves JF. 1881. On some fossil fishes, Crustacea and Mollusca from the Devonian rocks at Campbellton, NB, with descriptions of five new species. Can Nat 10:93–101.
Woodward AS. 1889. Acanthodian fishes from the Devonian of Canada. Ann Mag Nat Hist 4:183–184.

 

The sailfish enters the LRT with the swordfish, but closer to the anchovy

Updated January, 17, 2021
with new taxa and new scores nesting Istiophorus, the sailfish, with Elops, the anchovy. These give rise to the swordfish, Xiphias, and the European eel, Anguilla. All are derived from the Late Jurassic, Bavarichthys.

Traditionally the sailfish
(Istiophorous) is considered a billfish, closely related to the marlin (Makaira), and then the swordfish (Xiphias).

By contrast, 
the large reptile tree (LRT, 1641 taxa then, now 1793 taxa) nests the sailfish with the anchovy, Elops, close to the swordfish, Xiphias.

Figure 1. Istiophorus, the sailfish, nests with the cobia (Fig. 2) in the LRT, not with the swordfish.

Istiophorus platypterus (Shaw 1792 in Shaw and Nodder 1792; 3m) is the extant sailfish and a relative of Rachycentron, the extant cobia (above). The rostrum is extended, convergent with another fast, open ocean predator, the swordfish, Xiphias. The anterior dorsal fin is larger than the lateral area of the fish itself. Teeth are absent. The pectoral fins are long and slender. The anal fin is divided in two. The vertebral column is composed of relatively few, but large vertebrae.

Wikipedia reports,
“they [billfish] are also classified as being closely related to the mackerels and tuna within the suborder Scombroidei of the order Perciformes.”

By contrast,
billfish nest with anchovies in the LRT. The mackerel (genus: Scomber) also entered the LRT alongside Thunnus, the tuna, which it greatly resembles in every regard, other than size.

Figure 2. Skull of the sailfish, Istiophorus. Compare to Elops in figure 6.

Figure 3. Swordfish ontogeny (growth series). Hatchings have teeth, a short bill and an eel-like body still lacing pelvic fins.

Figure 4. Elops is the extant anchovy. Compare to Bavaricthys in figure 1 and Istiophorus in figure 5.

With the sailfish and swordfish gone, where does that leave the lonely barracuda?
In the LRT the barracuda nests with the similar remora (Remora) and cobria (Rachycentron), derived from the maui-mahi (Coryphaena).

Figure 5. Subset of the LRT focusing on ray fin fish. Eel-like taxa are highlighted.

Designed for reptiles,
the character list in the LRT is still working to separate fish as close in appearance as the swordfish and sailfish. So, please, don’t keep suggesting I expand the character list. It’s totally unnecessary.

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References
Gregory WK and Conrad GM 1937. The comparative anatomy of the swordfish (Xiphias) and the sailfish (Istiophorus). The American Museum Novitates, 952:1-25.
Kaup JJ 1826. Beiträge zu Amphibiologie und Ichthiyologie. Isis von Oken 19(1): 87-90.
Shaw G and Nodder FP 1792. Xiphias platypterus: The broadfinned swordfish. The naturalist’s miscellany, plate 88. 28 p. (Application to validate the nomen oblitum for the Indian Ocean sailfish (genus Istiophorus)

wiki/Istiophorus
wiki/Rachycentron

Kenomagnathus: what you can do with only 2 bones

Spindler 2020
reports on a new basal pelycosaur, Kenomagnathus scottae (ROM 43608; Upper Pennsylvanian, Late Carboniferous, Garnett, KS, USA; Figs 1-3) known from a single lacrimal and maxilla (with teeth) exposed in medial view (Fig. 1).

Figure 1. Kenomagnathus in situ from Spindler 2020. The halo of organic matter is interesting.

From the abstract:
“This is the oldest known diastema in synapsid evolution, and the first reported from a faunivorous member that lacks a precanine step, aside from Tetraceratops. This unique precanine morphology occurred independently from similar structures in Sphenacodontoidea.” 

See Spindler’s freehand drawing
of the ‘true diastema’ (Fig. 2). 

Figure 2. Kenomagnathus maxilla and lacrimal with the rest of the skull restored in lateral view. Note the deeper jugal (cyan), though not as deep as in Ophiacodon (Figs. 3, 4). For that reason the mandible of Ophiacodon was used in this restoration. Spindler’s freehand drawing indicates a deeper orbit, shallower jugal and smaller naris along with a larger mandible.

It is worth noting
that maxillary teeth shrink toward the naris in Ophiacodon (Fig. 3). A diastema may be present in Pantelosaurus (formerly Haptodus saxonicus, Fig.3). These pertinent taxa were not illustrated in Spindler 2020.

Figure 3. Pertinent synapsid skulls to scale. The origin of the Pelycosauria + Therapsida is marked by phylogenetic miniaturization, as in so many other clade origins. Note the depth of the jugal in basal taxa here.

Spindler’s freehand restoration
increased the size of the orbit and decreased the depth of the restored jugal. So this is yet another cautionary tale highlighting the danger in using freehand drawings in scientific studies.

The shallow jugal depth in the Spindler freehand restoration
is a key oversight. When repaired (Fig. 2) the semi-deep jugal of Kenomagnathus transitionally links deeper jugal Ophiacodon (Fig. 3) to shallower jugal Pantelosaurus and Haptodus (Fig. 3) at the base of Pelycosauria + Therapsida in the large reptile tree (LRT, 1642+ taxa). While running the risk of ‘Pulling a Larry Martin’, there are so few traits to consider here (Fig. 1) and none contradict the present hypothesis of interrelationships. All that puts Kenomagnathus in the lineage of synapsids leading to therapsids, mammals, primates and humans.

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References
Spindler F 2020. A faunivorous early sphenacodontian synapsid with a diastema. Palaeontologia Electronic 23(1):a01. doi: https://doi.org/10.26879/1023
https://palaeo-electronica.org/content/2020/2905-early-sphenacodontian-diastema

Helodus: a skull without sutures

Decades prior to PAUP and MacClade,
Professor Moy-Thomas 1936 reasoned that Helodus simplex (Fig. 1; Agassiz 1838; Early Carboniferous, 300mya, 30cm long) ) was close to the ancestry of the clade Holocephalii (ratfish, chimaeras and kin; Fig. 2), which we looked at yesterday. In complete accord, the large reptile tree (LRT, 1641 taxa; subset Fig. 3) fully supports that nesting using modern software: PAUP and MacClade. So… belated well done, Professor Moy-Thomas!

Figure 1. Helodus skull drawings from Moy-Thomas 1936 show no skull sutures. Colors are applied with blending edges to show were bones are based on tetrapod homologs. Gill bars are missing from these diagrams, so were added in light blue here.

In Helodus
the skull bones are all fused together, so suture estimates are provided here (Fig. 1) based on phyogenetic bracketing. Note the tiny premaxillary teeth and complex maxillary teeth. Tabulars appear to be absent. Note the coosified cervicals / anterior dorsals extending to the notochord and first dorsal spine.

Moy-Thomas 1936
considered the anatomy of Helodus in detail. From the abstract:

1. “The skull is found to be holostylic, and to have many characters in common with the skull of the Holocephali, but in some respects is less specialized.
2. The pectoral fins, with their long metapterygium, small propterygium, and fused anterior radials, resemble very closely those of the Holocephali.
3. The pelvic and unpaired fins, and general body shape are found to resemble those of the Holocephali.
4. It is concluded that the Cochliodonts are almost certainly closely related to the ancestors of the Holocephali, and the relatively unspecialized condition of the teeth gives support to the view that the holostylic condition of the jaws is primitive for the group. It is suggested that all the Bradyodonts were holostylic, that the hyomandibular may never have been suspensory, and that they may have diverged from the true Selachii before the hyomandibular played a part in the jaw suspension.”

FIgure 2. Ratfish (chimaera) and Heterodontus to scale.

 

Taking one phylogenetic step further back from Helodus,
yesterday we looked at Heterodontus (Fig. 2), the Chondrichthyes taxon phylogenetically ancestral to Helodus and the Holocephalii.

Figure 5. Subset of the LRT focusing on basal chordates, vertebrates and bony fish not related to tetrapods. Scomber and Istiophorus are new additions to the gold clade.

References
Agassiz L 1838. Recherches Sur Les Poissons Fossiles. Tome III (livr. 11). Imprimérie de Petitpierre, Neuchatel 73-140.
Moy-Thomas JA 1936. On the Structure and Affinities of the Carboniferous Cochliodont Helodus simplex. Cambridge University Press. 73(11):488–503.

wiki/Chondrichthyes
wiki/Chimaera
wiki/Cladoselache
wiki/Chondrosteus
wiki/Symmoriida
wiki/Horn_shark
wiki/Helodus not listed in English

The origin of the Holocephali (= chimaeras, ratfish)

Updated December 14, 2020
with the addition of many more taxa (see cladograms below).

Didier 1995 reports,
“There are two hypotheses on the origin of Holocephali (Bonaparte 1832). The first and most generally accepted scenario is that holocephalans have evolved from some lineage of bradyodont sharks. The second hypothesis suggests that holocephalans are most closely related to placoderms.”

FIgure 1. Ratfish (chimaera) and Heterodontus to scale.

By contrast
in the large reptile tree (LRT, 1635+ taxa then, 1776 now, Fig. 3) the few tested holocephalians (Chimaera (Fig. 1) and Belantsea)) arise from the horn shark, Heterodontus, (Fig. 1).

Didier 1995 reports,
“The adductor muscles of Heterodontus also lie anterior to the eye and superficially they resemble chimaeroid fishes in this respect. I interpret this as a convergent feature of heterodontids and chimaeroids.” That is the only mention of Heterodontus (Fig. 1) in the text. Squalus is the outgroup taxon in Didier’s figure 46 cladogram.

Figure y. Basal Gnathostomata with the addition of Rhinochimaera.

References
Bonaparte CL 1832. Iconografia delle fauna italica per le quattro classi degli animali vertebrati. Tomo III. Pesci. Roma. [Issued in puntata (installments), without pagination; total of 556 pp., 78 pls.
Didier DA 1995. Phylogenetic Systematics of Extant Chimaeroid Fishes (Holocephali, Chimaeroidei). American Museum Novitates 3119:86pp.

Crayssac basal pterosaur tracks? …or tenrec tracks?

Earlier we looked at Mazin and Pouech 2020
who claimed they had discovered “the first non-pterodactyloid pterosaurian trackways.” At the time, only the abstract was available to discuss and criticize.

Nine years ago
Peters 2011 published anurognathid tracks, which makes them the first non-pterodactyloid pterosaurian trackways published. Notable by its exclusion, Mazin and Pouech 2020 did not cite, “A catalog of pterosaur pedes for trackmaker identification” (Peters 2011), confirming Dr. S. Christopher Bennett’s threat, “You will not be published. And if you are published, you will not be cited.”

Now that I have seen the paper and the tracks,
(Figs. 1, 2) let’s determine what sort of tetrapod made those tracks named, Rhamphichnus crayssacensis, because they don’t look like other pterosaur tracks, as workers (see below) acknowledge.

Diagnosis from Mazin and Pouech 2020:
“Quadrupedal trackway with tridactyl digitigrade manus-prints and pentadactyl plantigrade to digitigrade pes-prints. Subparallel manus digit-prints orientated anteriorly. Pentadactyl pes-prints with more or less divergent digit prints. Pedal digit V divergent and postero-laterally rejected. Manus trackway slightly to clearly wider than the pes trackway.”

Distinct from typical pterosaur manus tracks:

1. tridactyl digit prints are subparallel (rather than widely splayed)
2. digits are oriented anteriorly (rather than laterally to posteriorly)
3. digits sometimes include additional medial and lateral impressions (never seen in other pterosaur tracks)
4. no claw marks are present (that seems wrong based on Fig. 1)
5. the manus impression is just anterior to the pes impression (rather than laterally and posteriorly, as in other pterosaur tracks)

Figure 1. Images from Mazin and Pouech 2020. Some manus tracks have at least four digits.

There are many
basal and derived pterosaurs with pedal digit 2 (or 2 and 3) the longest, distinct from Triassic pterosaurs. These were all examined and rejected as potential trackmakers matching Rhamphichnus for various reasons.

I also looked at 1600+ non-pterosaur trackmakers
due to the many unexpected traits (see list above) present in the Rhamphichnus tracks.

First and foremost,
the pterosaur antebrachium (radius + ulna) could not be pronated to produce anteriorly-oriented Rhamphichnus tracks. Due to folding and flying issues, pterosaurs, like birds, do not have the ability to pronate and supinate the wing. That’s why all pterosaur manus tracks are oriented laterally with fingers at full extension, impressing into the substrate. That manus digit 3 is often rotated posteriorly is a clue to its lepidosaurian ancestry. These facts form the hypothesis of a secondarily quadrupedal configuration for some, but not all pterosaurs.

One overlooked trackmaker stood out as a good match
for Rhamphichnus: the tenrec, Tenrec (Fig. 2), a small digitigrade quadrupedal mammal currently restricted to Madagascar. The medial and lateral manual digits are shorter than 2-4, which are parallel in orientation.

Figure 2. Rhamphichnus tracks compared to a Tenrec trackmaker. The brevity of pedal digit 5 is a mismatch, but a related taxon, Leptictidium, likewise reduces pedal digit 5.

One of those Tenrec sisters,
Rhynchocyon, greatly reduces manual digits 1 and 5, but pedal digit 3 is the longest.

Another Tenrec sister,
Leptictidium (Fig. 3), has a pes with a reduced pedal digit 5, but a short digit 2, but the manus is also a good match for Rhamphichnus. So there is great variation in the pes of tenrec clade members. Still, a small tenrec-like mammal remains a more parsimonious trackmaker than any Late Jurassic pterosaur. They were able to pronate the manus!

Figure 3. Elements of Leptictidium from Storch and Lister 1985.

Due to taxon exclusion,
Mazin and Pouech 2020 did not consider alternative trackmakers for the pterosaur-beach Rhamphichnus tracks that don’t match other pterosaur tracks or extremities. Now we’re stuck with an inappropriate name for these Late Jurassic tenrec tracks.

A late Jurassic tenrec?
The large reptile tree (LRT, 1637 taxa) supports the probability that a sister to Tenrec was present in in the Late Jurassic based on the coeval presence of derived members of Glires (Multiturberculata). Placental mammal fossils remain extremely rare in the Mesozoic, but these impressions add to their chronology.

It is worth repeating, due to the subject matter,
the Crayssac pterosaur beach still includes the pes of the JME-SOS 4009 specimen attributed to Rhamphorhynchus, as mentioned earlier. Here it is again (Fig. 4).

Figure 4. Crayssac track different from all others. Inset: Pes of Rhamphorhynchus muensteri JME-SOS 4009, no. 62 in the Wellnhofer catalog

twitter.com/Mark Witton mistakenly reports:
“Turns out we’ve been over-thinking it (pedal digit 5): it just lays flat on the ground during walking, like a regular toe.”

“For one, the walking fingers face forward, not sideways, as in pterodactyloids. This seems weird, but it turns out that non-ptero wing fingers fold roughly perpendicular to the walking digits.”

These basic bungles by a PhD pterosaur worker
demonstrate the dominance of myth-making among purported experts due to accepting published results like a journalist, without testing them, like a scientist. Dr. Witton’s 2013 pterosaur book is full of similar mistakes reviewed here in a seven-part series.

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References
Mazin J-M and Pouech J 2020.The first non-pterodactyloid pterosaurian trackways and the terrestrial ability of non-pterodactyloid pterosaurs. Geobios 16 January 2020. PDF
Peters, D. 2011. A Catalog of Pterosaur Pedes for Trackmaker Identification
Ichnos 18(2):114-141. http://dx.doi.org/10.1080/10420940.2011.573605

https://pterosaurheresies.wordpress.com/2020/01/18/first-non-pterodactyloid-pterosaurian-trackways-ever-described-no/