New insights into the ornithopod manus

Duckbills,
like Edmontosaurus, and their kin are the ornithopod ornithischian dinosaurs, a clade I have been ignoring until now. Wikipedia reports, “[they] started out as small, bipedal running grazers, and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world, and dominated the North American landscape.” 

Dryosaurus, Camptosaurus, Iguanodon and Edmontosaurus are genera within this clade and each has an interesting manus (Fig. 1). When one works in phylogenetic analysis it is imperative to compare homologous digits (apples to apples). In ornithopods, those homologies appear to be masked and perhaps misinterpreted by the appearances of new phalanges and the disappearances of old phalanges. Putting them all in one image (Fig.1) clarifies all issues (even without traveling to visit the fossils firsthand!). Hopefully the data are accurate to start with.

This all started with a phylogenetic analysis
that appeared to indicate that Edmontosaurus had a manual digit 1 with an extra digit that made it look like manual digit 2. Comparisons to other ornithopods ensued. A quick look through the Internet brought B. Switek’s article (see below) to the fore.

Figure 1. Ornithopod manus. Here the hands of Dryosaurus, Camptosaurus, Iguanodon and Edmontosaurus are compared. Note the turquoise metatarsal homologies and the digit identification based on that.

Figure 1. Ornithopod manus. Here the hands of Dryosaurus, Camptosaurus, Iguanodon and Edmontosaurus are compared. Note the turquoise metatarsal homologies and the digit identifications based on that.

Science writer Brian Switek 
writing for Smithsonian.com reports,

  1. “…the great herbivore Iguanodon had prominent thumb spikes.
  2. “The peculiar false thumb of Iguanodon was originally thought to set into the dinosaur’s nose.”
  3. “But why should Iguanodon have a hand spike? “
  4. “Though my own suggestion is not any better than those I have been disappointed by, I wonder if the Iguanodon spike is a Mesozoic equivalent of another false thumb seen among animals today—the enlarged wrist bones of red and giant pandas…  the Iguanodon spike was rigid.” Unfortunately that’s as far as journalist Switek has allowed himself to go, rather than proposing the homologies and comparisons demonstrated here.

Giving credit where credit is due,
Switek may be the first to suggest the spike was not a digit. I don’t know and was not able to find out the history of the spike. Given the text from his blogpost, you can see Switek’s choice of words actually evolves from “thumb spikes” to “false thumb” to “hand spike” to “enlarged wrist bone”. Like Brian, I also lack a PhD, but that doesn’t stop us from making contributions. If I’m duplicating earlier academic efforts, please let me know so credit can be given.

Here we’ll show
that the spike is indeed a wrist element… that digit 1 in Iguanodon and related taxa have one more phalanx, making it look like digit 2.

We’ll start with
the right manus of Dryosaurus, a basal ornithopod (at least in the large reptile tree it is, where only one other ornithopod, Edmontosaurus, is currently represented). During the course of this, I want you to focus on the the homologies of metatarsals 2 and 3 (colored in turquoise). These, I think, will guide us to correct interpretations of the other elements of the various ornithopod manus.

Now back to the manus of Dryosaurus:

  1. Data comes form loose bones in a photo formed in the shape of a hand, not an in-situ articulated hand. Thus I do not know the identification or placement of the carpals
  2. Five metatarsals are present.
  3. Mt3 is the longest. Slightly shorter is mt2.
  4. Phalangeal formula is 2-3-4-3-2, but digit 1 does not appear to be tipped with a sharp ungual. Is it missing? If so, that adds a phalanx to the formula 3-3-4-3-2.
  5. Digit 3 is the longest. Slightly shorter is digit 2.
  6. Unguals are lost in digits 4 and 5.

The manus of Camptosaurus

  1. Is reduced (stumpy) by comparison to Dryosaurus
  2. Mt 1 is a disk. M1.1 is a disk
  3. M3.2 appears to fuse with m3.3
  4. m4.3 and m5.2 are lost
  5. The new phalangeal formula is 2-3-3-2-1

The manus of Iguanodon

  1. is more robust and highly modified by comparison to Dryosaurus
  2. Two wrist elements fill the wrist. Two others extend medially.
  3. Digit 1 is longer and now sports an ungual
  4. Ungual 1 is not sharp
  5. Ungual 2 is a round hoof
  6. Ungual 3 (m3.4) is lost along with m3.3
  7. Mt4 is shorter. Two tiny phalanges are added.
  8. Digit 5 is absent.
  9. The new phalangeal formula is 3-3-2-4-0

The manus of Edmontosaurus 

  1. is long and gracile by comparison to Dryosaurus.
  2. Again, digit 1 has 3 phalanges, matching digits 2–4.
  3. Digit 4 is a vestige
  4. Mt 5 is again absent
  5. As in Iguanodon, ungual 1 is not sharp and ungual 2 is a hoof
  6. The new phalangeal formula is 3-3-3-3-0.

Always interesting to 
uncover little paradigm busters like these. Now back to phylogenetic analysis…

The carpus (wrist) of Pterodactylus scolopaciceps

Earlier we looked at the pectoral girdle of Pterodactylus scolopaciceps  BSP 1937 I 18 (Broili 1938, P. kochi n21 of Wellnhofer 1970, 1991).. And even earlier we looked at that elusive (they say it doesn’t exist!) manual digit 5. Today, some more thoughts on that wonderful wrist… (Fig. 1).

Figure 1. The wrist of Pterodactylus scolopaciceps BSP 1937 I 18 (Broili 1938, P. kochi n21 of Wellnhofer 1970, 1991). Manual digit 5 is a vestige, but it is there.

Figure 1. The wrist of Pterodactylus scolopaciceps BSP 1937 I 18 (Broili 1938, P. kochi n21 of Wellnhofer 1970, 1991). Manual digit 5 is a vestige, but it is there.

Manual digit 5
is here. So is metacarpal 5 and distal carpal 5

Figure 1. The wrist of Pterodactylus scolopaciceps BSP 1937 I 18 (Broili 1938, P. kochi n21 of Wellnhofer 1970, 1991). Manual digit 5 is a vestige, but it is there.

Figure 2. The wrist of Pterodactylus scolopaciceps BSP 1937 I 18 (Broili 1938, P. kochi n21 of Wellnhofer 1970, 1991). Manual digit 5 is a vestige, but it is there.

Metacarpals 1-3
are not pasted onto the anterior (during flight) face of the big metacarpal 4 as tradition dictates. Here mc1-3 are in their natural positions for tetrapods, palmar side down. Only metacarpal 4 is axially rotated so the wing finger folds (flexes) and extends in the place of the hand like bird and bat wings do. That means only metacarpal 3 attaches to metacarpal 4, mc2 lies between 1 and 3 and 1 hangs out in front.

Fingers 1-3
are dislocated and axially rotated anteriorly. In life they palms of the fingers would have been ventral, just like metacarpals 1-3 — not flexing anteriorly as they do here after crushing. Note the fingers are all disarticulated at the knuckle, which was a very loose joint, enabling 90 degrees of extension dorsally (in flight) or laterally (while quadrupedal for walking. Moreover, digit 3 was able to flex in the plane of the wing, like the wing. That produces manus impressions in which digit 3 is oriented posteriorly. That’s very weird for most tetrapods, but common in pterosaurs, as it indicates the quadrupedal configuration was achieved secondarily from an initial bipedal configuration.

Of added interest here….
Note the sawtooth posterior edges of the forelimb, hand and finger four where the wing membrane was attached, fed and enervated. Note also the large extensor tendon distal to the preaxial carpal. It is rarely preserved.

The preaxial carpal and pteroid
as you might remember, are former centralia having migrated to the outside (Peters 2009). We looked at analogous migrations here.

Radius and ulna
as in birds and bats, there is no pronation or supination in the pterosaur wrist and forearm. The elements are too close together to permit this. And that’s a good thing to keep the wing in the best orientation for flight. Bats and birds don’t twist their forearms either.

As you already know, every body part that disappears
goes out with a vestige.

References
Broili F 1938. Beobachtungen an Pterodactylus. Sitz-Bayerischen Akademie der Wissenschaten, zu München, Mathematischen-naturalischenAbteilung: 139–154.
Peters D 2009. A reinterpretation of pteroid articulation in pterosaurs. Journal of Vertebrate Paleontology 29:1327-1330.
Wellnhofer P 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Abhandlungen der Bayerischen Akademie der Wissenschaften, N.F., Munich 141: 1-133.

wiki/Pterodactylus

Do ceratopsid juveniles (phylogenetically) nest together?

The discovery of a second juvenile ceratopsid
(Currie et al. 2016) raised an interesting point: “In phylogenetic analysis, if all characters are coded as seen, the two juvenile ceratopsids (a partial Triceratops skull and the UALVP 52613 juvenile, Fig. 1) nest together. However, when size or age dependent characters are [not scored], the new juvenile (Chasmosaurus) specimen groups with other adult Chasmosaurus specimens.”

Figure 1. Chasmosaurus juvenile UALVP 52613 specimen.

Figure 1. Chasmosaurus juvenile UALVP 52613 specimen lacking forelimbs due to  taphoniomic loss down a nearby sinkhole.

So, does phylogenetic analysis fail us?
The new UALVP juvenile was recognized/identified as being closer to Chasmosaurus, just as the juvenile Triceratops was recognized as being closer to Triceratops, both on the basis of character traits and prior to analysis. But the Currie et al. unedited analysis takes us in another direction…

From the introduction
“The specimen comprises a nearly complete skeleton lying on its left side, lacking only the front limbs and girdle, which were lost many years ago into a large sinkhole….”

“The juvenile nature of this specimen is based on several lines of reasoning. At approximately 1.5 min total length, it is the smallest articulated ceratopsid skeleton that has ever been recovered. Immature bone textures on cranial bones (Brown et al., 2009), open neurocentral sutures throughout most of the vertebral column, incomplete fusion of sacral vertebrae, lack of fusion between caudal ribs and vertebrae, poorly formed articulations between limb bones, and many other characters confirm that this is an immature ceratopsid….”

“Of all the chasmosaurines from Dinosaur Park, it is most similar to Chasmosaurus belli and C. russelli.”

This interpretation
was made by expert and experienced assessment. The question is, why would the unedited Currie et al. analysis separate the juveniles from the adults and nest the juveniles together? They’re not exactly tadpoles or caterpillars, but they do change somewhat during maturation, following basic archosauromorph (including synapsid/mammal) growth strategies, that lepidosauromorphs (including pterosaurs) are less likely to follow.

When an adult Chasmosaurus
and the juvenile Chasmosaurus are added to the large reptile tree, using a character list NOT specific to ceratoposids, the juveniles nest with their respective adults, not with each other. And this happens despite the very few bones that represent the juvenile Triceratops (posterior face and shield only). Notably there are no other competing ceratopsid candidates in the present taxon list. All data was gleaned from online images. The adult data may be  represented by chimaera mounts and chimaera drawings. If the Currie et al analysis was restricted to just an adult and juvenile Triceratops and just an adult and juvenile Chasmosaurus, would adults nest with juveniles as they do in the large reptile tree? We don’t know because that test was not run.

Here’s how the large reptile tree divides
the Chasmosaurus adult and juvenile from the Triceratops adult and juvenile (posterior skull traits only). Please feel free to provide better data or more precise readings for any of these interpretations. Some were difficult to figure from available sources. At present I do not include traits for parietal fontanelles or horn lengths, which are the easiest two traits that most commonly separate Chasmosaurus from Triceratops and are reflected in their juveniles.

  1. skull table: C: depressed terrace, medial and lateral crests; T: convex
  2. snout in dorsal view: C: not constricted; T: constricted
  3. orbit positon: C: postorbital > preorbital; T: subequal
  4. lateral rostral shape: C: convex, smooth curve; T: double convex
  5. nasals/frontals: C: nasals >; T: subequal
  6. antorbital fenestra: C: absent; T: without mx fossa
  7. orbit/upper temporal fenestra: C: orbit not > T: orbit >
  8. orbit position/skull: C: anterior half of skull; T: not
  9. orbit shape: C: round to square: T: taller than wide
  10. upper temporal fenestrae: C: not closed or slit-like; T: closed or slit-like
  11. frontal shape: C: not wider posteriorly; T: wider posteriorly
  12. frontal shape 2: C: without posterior processes; T: with posterior processes
  13. posterior rim of parietal: C: transverse; T: anteriorly oriented or curved.
  14. parietal skull table: C: forms a sagittal crest: T: broad
  15. squamosal descent: C: mid level; T: ventral skull (ventral maxilla)
  16. skull roof fusion: C: parietal fusion only; T: frontal fusion and parietal fusion
  17. jaw joint orientation: C: descends from ventral mx; T: in line with ventral mx, after jugal arch.
  18. last maxillary tooth: C: posterior orbit; T: mid orbit
  19. mandible ventrally: C: 2-tier convex; T: straight
  20. 2nd sacral rib: C: not: T: double wide laterally
  21. manus/pes: C: subequal: T: manus smaller
  22. ilium: C: posterior process >; T: not
  23. metatarsal 1:4 ratio: C: 1 not > than half: 4 T: 1> half of 4
  24. metatarsals 2-4: C: < than half the tibia; T: not
  25. pedal 3.1 vs p2.1: C: not > T: 3.1>
  26. metatarsals 2 and 3: C: aligns with mt1; T: aligns with pedal 1.1
  27. pedal 4 length: C: subequal to mt 4; T: > mt4
  28. pedal digit 3 vs 4: C: 4 narrower than 3; T: 4 is not narrower

Shifting the juvenile Triceratops
to the juvenile Chasmosaurus adds 12 steps. Doing the opposite adds 21 steps. Bootstrap scores are over 99-100 for the three nodes represented by the four taxa. I have not reviewed the scores or data in the Currie et al study, which obviously adds more ceratopsid traits.

Added < 24 hours after original publication Below is a new reconstruction of the Triceratops juvenile based on text measurements and an adult skull compared to the original reconstruction that does not appear to have correctly scaled the mandible to the skull elements.

Figure 4. A new reconstruction of the Triceratops juvenile with the mandible and squamosal scaled to text measurements and shaped to adult elements compared to the original (Goodwin et al.) reconstruction which appears to have shortened the mandible.

Figure 4. A new reconstruction of the Triceratops juvenile with the mandible and squamosal scaled to text measurements and shaped to adult elements compared to the original (Goodwin et al.) reconstruction which appears to have shortened the mandible.

A YouTube video, Dinosaurs Decoded, shows Mark Goodwin reassembling the juvenile Triceratops skull. Click here to watch.

_______________________

Short notes for readers and critics
“Criticism of a writer is absolutely inevitable.” — Malcolm Gladwell.
Gladwell is one of the most respected and best-selling authors in current decades. Nevertheless, this interview on YouTube quotes several critics, many with scathing barbs. So, this give and take between writers and their critics is universal and ‘inevitable.’

On the other hand,
in Science, one either can or cannot duplicate experiments and observations. It should be cut and dried, but with errors and egos on both sides, it rarely is. Even so, most people think it is better to try/experiment with/refute alternate hypotheses. Aaaaaat least that’s the editorial policy at ReptileEvolution.com where occasional lack of talent and insight is sometimes overcome by tenacity, huge blocks of data and the ability to update online blunders.

References
Currie PJ,  Holmes RB, Ryan MJ and Coy C. 2016. A juvenile chasmosaurine ceratopsid (Dinosauria, Ornithischia) from the Dinosaur Park Formation, Alberta, Canada. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1048348.

 

 

Two basal turtles added to the large reptile tree

Today
we add Niolamia (Fig. 2) and Mongolochelys (Fig. 4) to the large reptile tree (Fig. 3).

Figure 3. Meiolania is another club-tailed, short-toed turtle like Proganochelys.

Figure 3. Meiolania is a club-tailed, late-surviving basalmost turtle  related to Niolamia.

One of the basalmost hard shell turtles,
Meiolania, has a more bizarre yet older sister. Niolamia (Fig. 2) has even larger supratemporal and tabular horns, that originated with taxa like toothy Elginia from the Late Permian (Fig. 2), All known meiolanids were late-surviving members of the most basal hard-shell turtle clade with probable origins in the Early Triassic. The age of Niolamia can only be estimated (Late Cretaceous to Eocene) due to purposeful loss of formation data by its 19th century collectors. Prior turtle workers have either judiciously or accidentally avoided putting these three taxa together in the same phylogenetic analysis, despite their obvious similarities.

Figure 1. The basal turtle, Niolamia, compared to the toothed pareiasaur/turtle?, Elginia. We have no post-crania for Elginia. Figure 1. The basal turtle, Niolamia, compared to the toothed pareiasaur/turtle?, Elginia. We have no post-crania for Elginia.

Figure 2. The basal turtle, Niolamia, compared to the toothed pareiasaur/turtle?, Elginia. We have no post-crania for Elginia.

Phylogenetic analysis
confirms what any first look suggests: Niolamia and Meiolania nest with each other. In the large reptile tree these two nest as basalmost hard shell turtles. If only we knew that Elginia has a carapace and plastron (current status unknown), then it would also be the basalmost hard shell turtle, despite the teeth.

Figure 1. The origin and evolution of turtles. Here Meiolania and Niolamia nest as the most basal hard shell turtles. Odontochelys is the most basal soft shell turtle.

Figure 3 The origin and evolution of turtles. Here Nolamia and Mongolochelys have been added to the previous tree. Meiolania and Niolamia nest as the most basal hard shell turtles. Odontochelys is the most basal soft shell turtle. Mongolochelys and Chubutemyx (Fig. 4) do not nest with meiolanids, but as more derived turtles lacking skull ornamentation.

Sterli and de la Fuente 2011
produced the latest published literature on Niolamia and this following a thorough cleaning of the fossil. Many of the bones colored above (Fig. 1) match similar bones in Elginia, but are considered ossified scales by Sterli and de la Fuente. They also considered the shelf-like tabulars to be neo-ossifications. Their emended diagnosis notes, “an extensive contribution of the supraoccipital to the dorsal skull roof.” THAT would be very odd to see any supraoccipital as an extensive dorsal element. On more derived turtles, the supra occipital extends posteriorly as a narrow element and its dorsal contribution is minimal. Sterli and de la Fuente do not list tabulars on their list of bone abbreviations. They’re not making homologies with pareiasaurs. The supratemporal horns are considered to be ‘horns’ arising from the squamosal. We looked at the traditional misidentification of the suptratemporal and squamosal in turtles earlier here. That concept has to be adopted universally in order to make further progress on turtle systematics.

In phylogenetic analysis
Sterli and de la Fuente 2011 nest meiolanids with  Mongolochelys (Late Cretaceous) and Chubutemys (Early Cretaceous), two late-surviving basal turtles with no trace of horns (Fig. 3). These two start the process of bone fusion seen in all later turtles. In the large reptile tree (subset Fig. 2) these two taxa nest more derived than Proganochelys and Proterochersis.

Figure 3. Mongolocheys and Cubutemys nest together near, but not at, the base of the hard shell turtles. Both were considered sisters to Meiolania and Niolamia by prior workers who did not include Elginia in phylogenetic analysis.

Figure 4. Mongolocheys and Cubutemys to scale nest together near, but not at, the base of the hard shell turtles. Both were considered sisters to Meiolania and Niolamia by prior workers who did not include Elginia in phylogenetic analysis. Both have the genesis of the large post temporal fenestra that contains large jaw muscles.

The horns of meiolanids
were lost in more derived hard-shell turtles. At the same time, turtles began to become sea turtles while others gained the ability to withdraw their skull beneath the carapace in one of two ways, vertical and sideways. So what look like derived traits in an apparently aberrant late-surviving clade are actually primitive and not quite as aberrant as previously thought.  Elginia provides the blueprint or bauplan for hard-shell turtle skulls, which retain a large supratemporal, contra all prior studies. As noted earlier, meiolanids are the last turtles to retain laterally splayed forelimbs. In all other known turtles, the elbows are oriented anteriorly. The club tail is also be primitive for turtles, but we don’t have the data on known stem turtle pareiasaurs yet…

Paralleling the situation in pterosaur ancestors
like Sharovipteryx and Longisquama, the exotic, difficult to nest turtle and stem turtle taxa (Fig. 2) end up nesting together in a large gamut phylogenetic analysis.

References
Gaffney ES 1983. The cranial morphology of the extinct horned turtle, Meiolania platyceps, from the Pleistocene of Lord Howe Island, Australia. Bulletin of the AMNH 175, article 4: 361-480.
Gaffney ES 1985. The cervical and caudal vertebrae of the cryptodiran turtle, Meiolania platyceps, form the Pleistocene of Lord Howe Island, Australia. American Museum Novitates 2805:1-29.
Gaffney ES 1996. The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae. Bulletin of the AMNH no. 229.
Owen R 1882. Description of some remains of the gigantic land-lizard (Megalania prisca
Owen), from Australia. Part III.Philosophical Transactions of the Royal Society London, series B, 172:547-556.
Owen R 1888. On parts of the skeleton of Meiolania platyceps (Owen). Philosophical Transactions of the Royal Society London, series B, 179: 181-191.
Sterli J and de la Fuente M 2011. Re-Description and Evolutionary Remarks on the Patagonian Horned Turtle Niolamia argentina Ameghino, 1899 (Testudinata, Meiolaniidae). Journal of Vertebrate Paleontology 31 (6): 1210–1229. doi:10.1080/039.031.0618.

Adding the Triassic turtle Proterochersis to the large reptile tree

No surprises here.
The Late Triassic German dome-shelled turtle, Proterochersis (Fraas 1913, Szczygiellski  and Sulej 2016; ZPAL V.39/48), was added to the large reptile tree. No surprise, it nested with the other Late Triassic German dome-shelled turtle, Proganochelys. I was worried that Proterochersis would cause loss of resolution because the specimen lacks a skull, cervicals, caudals and limbs. Thus, all scores were based on the dorsal verts, ribs and girdles. And that was enough.

Proganochelys and Proterochersis, two Traissic turtles.

Figure 1. Proganochelys and Proterochersis, two Traissic turtles.

Szczygiellski and Sulej 2016
recently looked at Proterochersis together with a new Triassic turtle, Murrhardtia.

Here’s a big question
Proganochelys has a tall set of clavicles (aka epiplastra) that contacted and braced both the plastron and carapace (Gaffney 1990). Several basal dome-shelled turtles have these. In the basal dome-shelled turtle, Meiolania, Gaffney (xxxx) reports, “In the plastron the epiplastra meet on the midline and bear a short median process, apparently not homologous to that in Proganochelys and Kayentachelys, that bifurcates dorsally and articulates with the scapula. The epiplastron is a paired, curved element, meeting on the midline at the front of the plastron and forming a dorsal process. None of the specimens show a midline suture.”

Szczygiellski and Sulej 2016 reported, “the sturdy build of Proganochelys quenstedti should … be considered its own apomorphy. The presence of strong dorsal epiplastral processes contacting the carapace may be one of the consequences: although the dorsal processes themselves are interpreted by Gaffney (1990) as remnants of ancestral amniote clavicles, their additional articulation with the carapace and strengthening might have stabilized the shell, and thus serve as a more rigid point of attachment for the limb musculature (which probably was required to support the heavy body). Large dorsal epiplastral processes are present in the slightly smaller Palaeochersis talampayensis (Sterli et al., 2007), but are weaker and do not articulate with the carapace in more basal Proterochersis spp. and Keuperotesta limendorsa gen. et sp. nov. In Odontochelys semitestacea they obviously do not contact the carapace, because no suitable point of attachment was available (Li et al., 2008), but they possibly played a similar role, temporarily supporting and strengthening the limb musculature (weakened by changes in rib position), and disappeared when the torso of the animal became fully stiffened and the pectoral girdle received its derived shape.”

References
Fraas E. 1913. Proterochersis, eine pleurodire Schilderöte aus dem Keuper. Jahreshefte des Vereins für Vaterlänzische Naturkunde in Württemberg 69: 13–30.
Szczygiellski T and Sulej T 2016. Revision of the Triassic European turtles Proterochersis and Murrhardtia (Reptilia, Testudinata, Proterochersidae), with the description of new taxa from Poland and Germany. Zoological Journal of the Linnean Society 177:395-427.
Gaffney ES 1996. The postcranial morphology of Meiolania platyceps and a review of the Meiolaniidae. Bulletin of the American Museum of Naturaly Histoyr 229: 1-165.

The skull of Sclerocormus reinterpreted.

Figure 1. Large Sclerocormus and its much smaller sister, Cartorhynchus. These nest with basal sauropterygians, not ichthyosauriforms.

Figure 1. Large Sclerocormus and its much smaller sister, Cartorhynchus. These nest with basal sauropterygians, not ichthyosauriforms. The odd thing about this genus is really the short neck, not the small head.

Yesterday we looked at the new basal sauropterygian with a tiny head, Sclerocormus (Figs. 1, 2). Originally Jiang et al. 2016 considered Sclerocormus ‘a large aberrant stem ichthyosauriform,’ but their cladogram did not have the stem ichthyosauriforms recovered by the 684-taxa reptile tree, Wumengosaurus, Thaisaurus and Xinminosaurus.

Basal sauropterygians often have a tiny skull. 
Check out these examples: Pachypleurosaurus, Keichousaurus, Plesiosaurus, Albertonectes. Given this pattern, the odd thing about Sclerocormus is its short neck, not its tiny skull. The outgroup, Qianxisaurus has a skull about equal to the cervical series.

As noted previously
the terms ‘aberrant’ or ‘engimatic’ usually translate into “somewhere along the way we made a huge mistake, but don’t know what to do about it.” For the same reason, pterosaurs are widely considered ‘aberrant’ archosaurs, Vancleavea is an ‘aberrant’ archosauriform, Daemonosaurus and Chilesaurus are aberrant theropods and caseasaurs are ‘aberrant’ synapsids. All of these taxa also nest elsewhere in the large reptile tree.

Moreover
several of the Jiang et al interpretations of the skull could not by confirmed by DGS tracings (Fig. 2). Others were just fine.

Figure 2. Sclerocormus skull as originally interpreted and reinterpreted here.

Figure 2. Sclerocormus skull as originally interpreted and reinterpreted here.

Reinterpretations

  1. Jiang et al. nasals  >  nasals + premaxillae
  2. Jiang et al. premaxilla (lower portion)   >  anterior maxilla
  3. Jiang et al. premaxilla (upper portion)  >   left dentary
  4. Jiang et al. missed the right dentary and all teeth
  5. Jiang et al. missed the occipitals (postparietals, tabulars, supra occipital)
  6. Jiang et al. maxilla   >   lacrimal
  7. Jiange et al. scapula    >  coracoid + scapula
  8. Jiang et al. mandible elements? are confirmed as actual mandible elements
  9. Jiang et al. left postfrontal   >   postorbital
  10. Jiang et al. left squamosal and postfrontal   >  left posterior mandible elements

Phylogenetically
here are the stem ichthyosaurs and a sampling if ichthyosaurs (Fig. 3). Note where hupehsuchids nest, as derived utatsusaurs and shastasaurs. Cartorhynchus and Sclerocormus (Fig. 1) do not nest here.

Figure 2. Subset of the large reptile tree focusing on ichthyosaurs. Note most of the more derived ichthyosaurs from Marek et al. 2015, are not listed here. So we're not comparing apples to apples here.

Figure 2. Subset of the large reptile tree focusing on ichthyosaurs. Note most of the more derived ichthyosaurs from Marek et al. 2015, are not listed here. So we’re not comparing apples to apples here.

References
Jiang D-Y, Motani R, Huang J-D, Tintori A, Hu Y-C, Rieppel O, Fraser NC, Ji C, Kelley NP, Fu W-L and Zhang R 2016. A large aberrant stem ichthyosauriform indicating early rise and demise of ichthyosauromorphs in the wake of the end-Permian extinction. Nature Scientific Reports online here.

A new ichthyosaur mimic: Sclerocormus

A new Nature paper
by Jiang et al. 2016 introduces Sclerocormus, a large sister to the much smaller Cartorhynchus. Like a marine Cotylorhynchus, this odd basal sauropterygian had a tiny skull not much larger than that of its much smaller, big-headed sister (Fig. 1).

Figure 1. Large Sclerocormus and its much smaller sister, Cartorhynchus. These nest with basal sauropterygians, not ichthyosauriforms.

Figure 1. Large Sclerocormus and its much smaller sister, Cartorhynchus. These nest with basal sauropterygians, not ichthyosauriforms. Click to enlarge. Note the skull size of the two are within a short range.

These two nested
with Qianxisaurus, a basal sauropterygian/pachypleurosaur, not basal ichthyosauriforms. The authors are still in the dark about ichthyosaur ancestors. You can trace them, or any taxon, back to basal tetrapods here.

Figure 1. Although the pectoral girdle was preserved just behind the skull, in all sister taxa there are about 19 cervicals and 19 dorsals. Plus the pectoral girdle itself is very wide, better suited to the widest ribs. Perhaps Cartorhynchus had a longer neck than commonly assumed.

Figure 2. Although the pectoral girdle was preserved just behind the skull, in all sister taxa there are about 19 cervicals and 19 dorsals. Plus the pectoral girdle itself is very wide, better suited to the widest ribs. Perhaps Cartorhynchus had a longer neck than commonly assumed.

The authors
report that Sclerocormus had no teeth and that the nasals extended to the tip of the rostrum. I have to disagree with both observation given the photographic data and lack of similarity in sister. They also misidentified a few bones. Their big scapula is a posterior coronoid + smaller scapula.

More coming in later posts.

References
Jiang D-Y, Motani R, Huang J-D, Tintori A, Hu Y-C, Rieppel O, Fraser NC, Ji C, Kelley NP, Fu W-L and Zhang R 2016. A large aberrant stem ichthyosauriform indicating early rise and demise of ichthyosauromorphs in the wake of the end-Permian extinction. Nature Scientific Reports online here.